THE FAMILIES

AND GENERA
OF VASCULAR PLANTS

Edited by K. Kubitzki

Springer-Verlag
Berlin Heidelberg GmbH
Volumes published in this series:

Volume I Pteridophytes and Gymnosperms

Edited by K.U. Kramer and P.S. Green (1990)

Volume II Flowering Plants. Dicotyledons: Magnoliid, Hamamelid and Caryophyllid

Families
Edited by K. Kubitzki, J.G. Rohwer, and V. Bittrich (1993)

Volume III Flowering Plants. Monocotyledons: Lilianae (except Orchidaceae)

Edited by K. Kubitzki {1998)

Volume IV Flowering Plants. Monocotyledons: Alismatanae and Commelinanae

(except Gramineae)
Edited by K. Kubitzki {1998)
The Families
and Genera
of Vascular Plants
Edited by K. Kubitzki

IV Flowering Plants · Monocotyledons

Alismatanae and Commelinanae (except Gramineae)

Volume Editor:
K. Kubitzki
in Collaboration with H. Huber, P.J. Rudall, P.S. Stevens, and T. Stiitzel

With 126 Figures

Springer
Professor Dr. KLAUS KUBITZKI
Institut fiir Allgemeine Botanik
und Botanischer Garten
OhnhorststraBe 18
22609 Hamburg, Germany

ISBN 978-3-642-08378-5

Library of Congress Cataloging-in-Publication Data.

Flowering plants, Monocotyledons : Alismatanae and Commelinanae

(except Gramineae) I volume editor, K. Kubitzki; in collaboration
with H. Huber ... [et al.].
p. em.- (The families and genera of vascular plants; 4)
Includes bibliographical references and index.
ISBN 978-3-642-08378-5 ISBN 978-3-662-03531-3 (eBook)
DOI 10.1007/978-3-662-03531-3
1. Monocotyledons - Classification. I. Kubitzki, Klaus, 1933-
II. Huber, Herbert, Prof. Dr. rer. nat. Ill. Series.
QK495.A14F54 1998
584'.7- dc21

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Originally published by Springer-Verlag Berlin Heidelberg New York in 1998
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Preface

When Rolf Dahlgren and I embarked on preparing this book series, Rolf took prime
responsibility for monocotyledons, which had interested him for a long time. After
finishing his comparative study and family classification of the monocots, he devoted
much energy to the acquisition and editing of family treatments for the present series.
After his untimely death, Peter Goldblatt, who had worked with him, continued to handle
further incoming monocot manuscripts until, in the early 1990s, his other obligations no
longer allowed him to continue. At that time, some 30 manuscripts in various states of
perfection had accumulated, which seemed to form a solid basis for a speedy completion
of the FGVP monocots; with the exception of the grasses and orchids which would appear
in separate volumes. I felt a strong obligation to do everything to help in publishing the
manuscripts that had been put into our hands. I finally decided to take charge of them
personally, although during my life as a botainst I had never seriously been interested in
monocots.
The beginning of my involvement coincided with the revival of interest in monocot
systematics, which became manifest in the Kew Monocot Conference and the publication
of its proceedings in 1995. The latter helped me greatly in finding access to modern
concepts in monocot classification and evolution; this made it necessary to make consid-
erable changes to all the existing contributions. The results from molecular systematics
have not only greatly influenced phylogenetic concepts but have also led to an increase in
the number of monocot families, contrary to my original aim to reduce them. It is likely
that a number of genera such as those treated here as "genera anomala" may deserve
family status; further progress in molecular systematics, implying the use of additional
genes and a denser sampling, will certainly result in changes of our phylogenetic and
taxonomic concepts.
While working on these volumes, I tried to do justice to the most recent scientific
developments and, at the same time, tried not to lose sight of the legacy left to us from
earlier generations. Therefore, I often added information from important earlier works,
especially from the German morphological literature that is largely unknown abroad. In
floral morphology, I aimed at greater precision and clarity and made in these two volumes
a consistent distinction between style and stylulus (for an explanation of the latter term,
see footnote on p. 382, Vol. III). Nevertheless, I am fully aware of possible imperfections
and inconsistencies in the more than 100 family contributions; any attempt at reducing
their number, however, would have meant an unjustifiable delay in the publication of
these volumes.
I would like to express my deep gratitude to all those who supported my work in various
ways. The directors and curators of the herbaria in Berlin, Geneva, Kew, and Munich
always responded quickly in putting critical and rare herbarium material at my disposal.
W. Barthlott, R. Brummitt, M. Chase, J. Reveal, F. Salamini, and D.W. Stevenson, among
others, provided important information as to their own or others' recent work. A.
Meerow, Nam Soak Lee, J. Kadereit, M. Sazima, T. Sttitzel, S. Verhook, Woo-Tschul Lee,
and the librarians of my own institution, of the Botanic Garden and Museum at Berlin
and of the Botanische Staatssammlung Mtinchen helped with rare literature. C. Bayer, P.E.
Berry, C.D.K. Cook, D.A. Cooke, M. Fay, P. Goldblatt, 0. Huber, M. Lock, S. Mayo, A.
Meerow, I. Nordal, H.-H. Poppendieck, S.S. Renner, J. Semir, M. Sazima, H.-J. Tillich, G.
Wiegleb, and others read and commented on manuscripts. C. Schirarend and H.-H.
Poppendieck offered important help with translations from Russian and into English,
VI Preface

respectively. C. Bayer very patiently tried to develop my understanding of inflorescence

morphology. A special word of thanks, for his unfailing help, is due to J. Bogner, who not
only reviewed various manuscripts but also provided important plant material and illus-
trations. For the generous permission to use illustrations from Vol. 6 of his Plant Life
(Leningrad 1982), I am deeply indebted to A.L. Takhtajan. I am also most grateful to all
those authors and copyright holders who permitted the use of their illustrations. Likewise,
I wish to thank H.-D. Behnke, C.D.K. Cook, E. Kohler, J. Kress, H.P. Linder, R. Mello-Silva,
S. Nilsson with E. Grafstrom, and M. Harley for making valuable light, scanning and
transmission electron microscope photographs available.
Finally, I wish to thank all contributors for generously sharing their specialist knowl-
edge and for their patience, with which they reacted to all my suggestions and requests.
Last, but not least, my thanks go to the editorial and production staff of Springer Verlag
who, as always, have done splendid work.

Hamburg, May 1998 K. KUBITZKI

Contents

CONSPECTUS OF FAMILIES TREATED IN THIS VOLUME

K. KUBITZKI . . . . . . .

GENERAL REFERENCES 5

Acoraceae J. BoGNER and S.J. MAYO 7

Alismataceae R.R. HAYNES, D.H. LEs, and L.B. HoLM-NIELSEN 11
Anarthriaceae H.P. LINDER, B.G. BRIGGS, and L.A.S. JoHNSON . 19
Aponogetonaceae H.W.E. van BRUGGEN . . . . . . . . . . 21
Araceae S.J. MAYO, J. BoGNER, and P.C. BoYCE 26

Bromeliaceae L.B. SMITH and W. TILL 74

Butomaceae C.D.K. CooK 100

Cannaceae K. KUBITZKI 103

Centrolepidaceae D.A. CooKE 106
Commelinaceae R.B. FADEN 109
Costaceae K. LARSEN 128
Cymodoceaceae J, Kuo and A.J. McCOMB 133
Cyperaceae P. GOETGHEBEUR . . . . 141

Dasypogonaceae H.T. CLIFFORD, G.J. KEIGHERY, and J.G. CoNRAN. 190

Ecdeiocoleaceae H.P. LINDER, B.G. BRIGGS, and L.A.S. JoHNSON . 195

Eriocaulaceae T. STUTZEL . . . . . . . . . . 197

Flagellariaceae 0. APPEL and C. BAYER . 208

Haemodoraceae M.G. SIMPSON . . . . . . 212

Hanguanaceae C. BAYER, 0. APPEL, and P.J. RuDALL 223
Heliconiaceae L. ANDERSSON 226
Hydatellaceae U.HAMANN 231
Hydrocharitaceae C.D.K. CooK . 234

Joinvilleaceae C. BAYER and 0. APPEL 249

Juncaceae H. BALSLEV . . . . . . . 252
Juncaginaceae R.R. HAYNES, D.H. LEs, and L.B. HoLM-NIELSEN. 260

Lemnaceae E. LANDOLT . . . . . . . . . . . . . . . . . . . . . 264

Limnocharitaceae R.R. HAYNES, D.H. LEs, and L.B. HoLM-NIELSEN . 271
Lowiaceae K. LARSEN .. 275

Marantaceae L. ANDERSSON 278

Mayacaceae D.W. STEVENSON 294
Musaceae L. ANDERSSON . . 296

Najadaceae R.R. HAYNES, L.B. HoLM-NIELSEN, and D.H. LEs . 301

VIII Contents

Palmae J. DRANSFIELD and N.W. UHL 306

Philydraceae U. HAMANN . . . . . . . 389
Pontederiaceae C.D.K. CooK . . . . . . . . . . 395
Posidoniaceae J. Kuo and A.J. McCOMB . . . 404
Potamogetonaceae R.R. HAYNES, D.H. LEs, and L.B. HoLM-NIELSEN. 408

Rapateaceae D.W. STEVENSON, M. CoLELLA, and B. BooM . . . 415

Restionaceae H.P. LINDER, B.G. BRIGGs, and L.A.S. JoHNSON .. 425
Ruppiaceae R.R. HAYNES, L.B. HOLM-NIELSEN, and D.H. LEs . 445

Scheuchzeriaceae R.R. HAYNES, D.H. LEs, and L.B. HoLM-NIELSEN. 449

Strelitziaceae L. ANDERSSON 451

Thurniaceae K. KUBITZKI 455

Typhaceae K. KUBITZKI 457

Xyridaceae R. KRAL .. 461

Zannichelliaceae R.R. HAYNES, D.H. LES, and L.B. HoLM-NIELSEN. 470

Zingiberaceae K. LARSEN, J.M. LOCK, H. MAAS, and P.J.M. MAAS 474
Zosteraceae J. Kuo and A.J. McCoMB 496

Index of Scientific Names 503

List of Contributors

ANDERSSON, L. Institute of Systematic Botany, Carl Skottsbergs Gata 22,

41319 Goteborg, Sweden
APPEL, 0. Institut flir Allgemeine Botanik und Herbarium, Universitat
Hamburg, Ohnhorststr. 18, 22609 Hamburg, Germany
BALSLEY, H. University of Aarhus, Botanical Institute, Nordlandsvej 68,
8240 Risskov, Denmark
BAYER, C. Institut fur Allgemeine Botanik und Herbarium, Universitat
Hamburg, Ohnhorststr. 18, 22609 Hamburg, Germany
BoGNER, J. Menzinger Str. 63, 80638 Munchen, Germany
BooM, B. The New York Botanical Garden, Bronx, New York 10458-
5126, USA
BoYcE, P. Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB,
UK
BRIGGS, B.G. Royal Botanic Gardens, Mrs. Macquaries Road, Sydney,
NSW 2000, Australia
BRUGGEN, H.W.E. van Linthorst Homanstraat 19, 1963 KM Hemskeerk,
The Netherlands
CLIFFORD, H.T. Queensland Museum, P.O.B. 3300, South Brisbane Qld 4101,
Australia
CoLELLA, M. The New York Botanical Garden, Bronx, New York 10458-
5126, USA
CoNRAN, J.G. Department of Botany, University of Adelaide, Adelaide
S.A. 5005, Australia
CooK, C.D.K. Botanischer Garten, Universitat Zurich, Zollikerstr. 107,
8008 Zurich, Switzerland
CooKE, D.A. 58 Gardiner Avenue, Warradale S.A. 5046, Australia
DRANSFIELD, J. Royal Botanic Gardens Kew, Richmond, Survey TWG 3AB,
UK
FADEN, R.B. Smithsonian Institution, Department of Botany NBH 166,
Washington DC 20560-0001, USA
GoETGHEBEUR, P.J. Laboratory of Plant Systematics, University of Gent,
Ledeganckstraat 35, 9000 Gent, Belgium
HAMANN, U. Deceased
HAYNES, R.R. University of Alabama, Dept. of Botany, P.O.B. 870344,
Tuscaloosa AL 35487-0344, USA
HoLM-NIELSEN, L.B. The World Bank, ESP Washington DC. 20433, USA
X List of Contributors

HUBER, H. Universitat Kaiserslautern, FB Biologie, Postfach 3049,

67653 Kaiserslautern, Germany
JOHNSON, L.A.S. Deceased
KEIGHERY, G.J. Western Australian Wildlife Research Center, Department of
Conservation and Land Management, P.O.B. 51, Wanneroo
W.A. 6065, Australia
KRAL R. Biology Department, Vanderbilt University, Box 1705, StaB,
Nashville, Tennessee 37235, USA -
KUBITZKI, K. Institut fur Allgemeine Botanik und Herbarium, Universitat
Hamburg, Ohnhorststr. 18,22609 Hamburg, Germany
Kuo,J. Centre for Microscopy and Microanalysis University of
Western Australia, Nedlands, W.A. 6009, Australia
LANDOLT, E. Geobotanisches Institut ETH, Stiftung Rubel,
Zurichbergstr. 38, 8044 Zurich, Switzerland
LARSEN, K. University of Aarhus, Botanical Institute, Nordlandsvej 68,
8240 Risskov, Denmark
LES, D.H. Department of Ecology and Evolutionary Biology, University
of Connecticut, Storrs, CT 06269-3042, USA
LINDER, H.P. Botany Department, University of Cape Town,
Rondebosch 7700, South Africa
LOCK, J.M. Herbarium, Royal Botanic Gardens, Kew, Richmond,
Surrey TW9 3AB, UK
MAAS van der Department of Plant Ecology and Evolutionary Biology,
KAMER, H. University of Utrecht, Heidelberglaan 2, 3584 CS Utrecht,
The Netherlands
MAAS, P.J.M. Department of Plant Ecology and Evolutionary Biology,
University of Utrecht, Heidelberglaan 2, 3584 CS Utrecht,
The Netherlands
MAYO, S.J. Royal Botanic Garden, Kew, Richmond, Surrey TW9 3AB, UK
McCoMB, A.J. School of Biological and Environmental Sciences, Murdoch
University, Murdoch W.A. 6150, Australia
RUDALL, P.J. Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, UK
SIMPSON, M.G. Department of Biology, San Diego State University,
San Diego, CA 92182-4614, USA
SMITH, L.B. Deceased
STEVENSON, D.W. The New York Botanical Garden, Bronx, New York 10458-
5126, USA
STUTZEL, T. Spezielle Botanik, Ruhr-Universitat, Postfach 10 21 48,
44801 Bochum, Germany
TILL, w. Institut fur Botanik und Botanischer Garten, Rennweg 14,
1030 Wien, Austria
UHL,N.W. L.H. Bailey Hortorium, 462 Mann Library,
Cornell University, Ithaca, N.Y., USA
 Conspectus of Families Treated in this Volume
Conspectus of Families Treated in this Volume
A. Acoraceae
Leaves equitant; stomata paracytic; vessels in roots and rhi-
zomes with scalariform perforation; ethereal oil cells present;
silica bodies and raphides lacking, but calcium oxalate present
as prisms; anthers introrse; endosperm development cellular;
seeds with both endosperm and perisperm, the latter starch-
free and derived from the nucellar epidermis. Only one family,
1/2-4, Northern Hemisphere Acoraceae
B. Alismatanae
Aquatic, marsh, or terrestrial herbs or climbing or epiphytic
herbs or shrubs; stomates mostly paracytic; vessels mostly
only in the roots (or absent by reduction), with scalariform
perforation; tapetum periplasmodial with uninucleate tapetal
cells; ovules bitegmic, crassinucellate, or sometimes almost
tenuinucellate; endosperm development Helobial or Nuclear;
embryo often storing and chlorophyllous
1. Arales
Herbs, sometimes climbing and woody, less often aquatic;
raphides present; flowers bractless; gynoecium syncarpous;
endosperm development cellular; fruits mostly baccate; en-
dosperm often present and with aleurone, different forms of
starch grains and oil, or lacking. Two families
Terrestrial or rarely aquatic plants with roots, stems, and
leaves and vessels in the roots; laticifers often present;
flowers numerous in a spadix that is subtended by a well-
developed spathe. 105/3300, nearly cosmopolitan, mostly
tropical Araceae
Plants leafless, free-floating, with or without 1-several
roots; vessels lacking, tracheids in the roots, or absent;
flowers only 1 or, according to interpretation, 2-3( -4) in an
inflorescence. 4/30, cosmopolitan Lemnaceae
2. Alismatales
Aquatic or marsh herbs without raphides; stems with axillary,
multiseriate hairs (intravaginal squamules) at the nodes
(lacking in Limnocharitaceae, replaced by uniseriate hairs
in Scheuchzeria); flowers with subtending bracts; gynoe-
cium choricarpous or syncarpous (pseudomonomerous);
endosperm development Helobial (Nuclear); mature seeds
nearly always without endosperm. Fourteen families
Perianth generally differentiated into sepals and petals
(except Butomus ); flowers often bracteate
2 Flowers epigynous; [inflorescence borne within a spathe;
scape often curved or spiral; ovary choricarpous or syn-
carpous; placentation laminar; staminate flowers borne
on evanescent pedicels or abscising from parent before
an thesis; pollen inaperturate ]. 17/76, cosmopolitan
Hydrocharitaceae
2 Flowers hypogynous; carpels distinct, or only basally
connate
3 Leaves linear, not differentiated into blade and petiole;
plants without secretory canals; embryo straight;
pollen sulcate. 1/1, temperate Eurasia Butomaceae
3 Leaves usually differentiated into blade and petiole;
plants lactescent; embryo horseshoe-shaped; pollen
pantoporate or inaperturate
4 Carpels 6-numerous, each with 1( -2) basal ovules,
(2-numerous ovules on marginal placentae:
Damasonium); fruit achenes (follicular). 12/80,
subcosmopolitan Alismataceae
4 Carpels 3-numerous, each with several or many
ovules in laminar placentation; fruit follicular. 3/8,
tropics of the Old and New World
Limnocharitaceae
Perianth, when present, not differentiated into sepals and
petals; bracts wanting or small and inconspicuous, except
in Scheuchzeriaceae, but perianth sometimes replaced by a
single bract
5 Ovules ( 1-)2-several in each of the (2- )3-several distinct
or basally connate carpels; fruit follicular; stamens 6 or
more; anthers extrorse
6 Emergent marsh plant; tepals 6, never petaloid; inflo-
rescence a raceme, each pedicel subtended by a bract;
pollen inaperturate, shed as dyads. 1/1, arctic and
temperate regions of Northern Hemisphere
Scheuchzeriaceae
6 Aquatic plants with floating leaf blades or wholly sub-
mersed leaves; tepals 1-3( -6), sometimes petaloid;
inflorescence a simple or basally forking spike, appar-
ently bractless; pollen sulcate. 1/45, predominantly in
the Southern Hemisphere of the Old World
Aponogetonaceae
5 Ovules solitary in each distinct carpel or in each locule
of a syncarpous ovary; fruit mostly indehiscent or only
tardily and irregularly dehiscent; stamens 6-1; [pollen
inaperturate]
7 Leaves all basal, the plants commonly largely emer-
gent, [scapose with a terminal spike or raceme; flowers
actinomorphic or slightly zygomorphic, hermaphro-
ditic, or pistillate or staminate, 3-merous, 2-merous,
or 1-merous]. 4/12, subcosmopolitan Juncaginaceae
7 Leaves, or many of them, cauline, sometimes with a
floating blade, but plants not emergent
8 Pollen spheroidal or isobilateral, nor threadlike;
plants of fresh or alkaline or brackish water (Ruppia
seldom marine)
2 Conspectus of Families Treated in this Volume
9 Flowers hermaphroditic; stamens 2-4; [carpels
(1-)4(-16), distinct]
10 Tepals 4; stamens 4; fruiting carpels sessile;
pollen ellipsoid to spheroidal. 2-3/85, subcos-
mopolitan Potamogetonaceae
10 Tepals 0; stamens 2; fruiting carpels long-
stipitate; pollen elongate-arcuate. 1/1-10,
subcosmopolitan Ruppiaceae
9 Flowers unisexual; stamen 1
11 Anthers 4-, rarely 1-sporangiate; ovary com-
pound, unilocular, surmounted by a short
style with 3-4 conspicuous stigmatic branches;
ovule basal, erect. 1/40, subcosmopolitan
Najadaceae
11 Anthers 2-, 4-, or 12-sporangiate; carpels dis-
tinct, 1-9, each with a short or elongate stylu-
lus bearing an enlarged; peltate or feathery
stigma; ovule ventral-apical, pendulous.
4/10-12, subcosmopolitan Zannichelliaceae
8 Pollen filiform, threadlike; plants marine; [flowers
naked]
12 Flowers perfect; stamens 3, distinct; gynoe-
cium of a single unicarpellate pistil;
[tanniniferous cells present]. 1/9, Australia,
Mediterranean Posidoniaceae
12 Flowers unisexual; stamens 1, or 2 and con-
nate; carpels 2
13 Gynoecium of 2 separate carpels; stamens
2, united back to back; tanniniferous cells
present. 5/16, tropical and subtropical
oceans Cymodoceaceae
13 Gynoecium a single unilocular, bicarpel-
late pistil; stamen solitary; tanniniferous
cells wanting. 3/18, temperate and sub-
tropical regions of both hemispheres
Zosteraceae
C. Commelinanae
Mostly terrestrial, herbaceous or woody, sometimes with con-
spicuous stem thickening but lacking secondary growth;
unlignified cell walls with bound UV-fluorescent coumarate
and ferulate; root epidermis often differentiated into long and
short cells; leaf epidermis with silica bodies or particles;
raphides sometimes present; stomata paracytic; epicuticular
wax of unoriented platelets or compound rodlets; endosperm
copious
1. Principes
Woody plants with vessels in roots, stems, and leaves; stomata
tetracytic; calcium oxalate as raphides, crystals, and sand;
silica bodies common; ptyxis plicate; flowers trimerous, num-
bers of stamens and carpels often increased; septal nectaries
often present; microsporogenesis mostly successive; en-
dosperm development Nuclear; endosperm with fat, oils, and
hemicellulose but lacking starch. Only one family. 190/2000,
pantropical Palmae
2. Dasypogonales
Shrubby or arborescent perennials; vessels only in the roots;
flowers solitary or in globular heads; ovules with small embryo
sac and large starch-filled chalazosperm. Only one family. 4/8,
S and SW Australia Dasypogonaceae
3. Zingiberales
Rhizomatic herbs; vessels in roots, sometimes also stems;
silica bodies present, usually tannin cells, and often raphides,
(ethereal oil cells in Zingiberaceae); leaves usually petio-
late; ptyxis supervolute; flowers in compact, thyrse-derived
spikes, epigynous, zygomorphic to asymmetric; functional
stamens 6, 5, 1, or 1/2; septal nectaries often present; pollen
sulcate (-derived) but most often inaperturate; endosperm
development Nuclear or Helobial; fruit capsular or a schizo-
carp; seeds mostly operculate, containing endosperm
and mostly copious perisperm, these starchy. Eight families,
tropical
Functional stamens 6 or 5; plants with raphide sacs
2 Ovary conspicuously prolonged into a slender, hypan-
thiumlike neck; inflorescence of repeating units without
specialized subtending bracts; flowers malodorous. 1/10,
SE Asia Lowiaceae
2 Ovary not prolonged into a hypanthiumlike neck; in-
florescence with 1-many folded or boat-shaped or spa-
the-like main bracts, each subtending or enfolding
a compact, few-flowered, monochasial cyme; flowers
nectariferous, sweet -smelling
3 Flowers functionally unisexual; leaves spiral; laticifers
present; fruit fleshy, indehiscent; [seeds not arillate ].
2/40, Old World tropics Musaceae
3 Flowers perfect; leaves distichous; laticifers lacking;
fruit capsular or schizocarpic
4 Ovules numerous in each locule; fruit capsular;
seeds arillate; median sepal abaxial. 3/6, tropical
America, Africa, Madagascar Strelitziaceae
4 Ovules solitary in each locule; fruit schizocarpic
with 3 operculate pyrenes; seeds not arillate; median
sepal adaxial. 1/200, tropical America, Melanesia
Heliconiaceae
Functional stamen 1 or 1/2, plants without raphide sacs
5 Anthers 2-thecate, often not strongly petaloid; flowers
zygomorphic; endosperm development Helobial; sepals
connate below
6 Leaves distichous, often forming pseudostems; sheaths
mostly open; plants with abundant ethereal oil and
tannin cells; labellum formed by 2 connate staminodes
from inner stamina) cycle. 50/1300, pantropical
Zingiberaceae
6 Leaves spiral on spirally contorted stems; sheaths ini-
tially closed; plants lacking ethereal oil and tannin
cells; labellum formed by 5 connate staminodes from
both cycles. 4/100, pantropical Costaceae
5 Stamen with 1 functional theca, the other anther-half
petaloid; flowers asymmetric; endosperm development
Nuclear; sepals distinct
7 Flowers borne in mirror-image pairs; stem without
mucilage canals; petiole distally pulvinate; ovules soli-
tary in the single locule or in each of the 3 locules of
the ovary; seeds mostly arillate. 31/450, pantropical
Marantaceae
7 Flowers not paired; stem with mucilage canals; petiole
not pulvinate; ovules numerous in each of the 3 locules
of the ovary; seeds not arillate. 1/10-25, tropical and
subtropical America and widely naturalized elsewhere
Cannaceae
 Conspectus of Families Treated in this Volume
4. Bromeliales
Perennial rosette plants; silica bodies and raphides present;
stomata tetracytic; roots often running down inside the
stem; leaves spiral, dilated-sheathing at the base, usually
bearing peltate trichomes; flowers hypogynous to epigy-
nous, heterochlamydeous; septal nectaries nearly always
present; fruit capsular or baccate; embryo lateral; endosperm
development Helobial; endosperm starchy. Only one family.
56/2600, neotropical, with a single species in W Africa
Bromeliaceae
Related to Commelinales or Zingiberales:
Robust terrestrial or aquatic from creeping rhizomes (with
long stolons: Hanguana malayana); vessels only in roots;
ptyxis supervolute; stomata tetracytic; pollen inaperturate.
1/5, SE Asia, Micronesia, Australia Hanguanaceae
5. Commelinales
Perennial and annual herbs; raphides widespread; silica
bodies usually absent (present in some genera of Com-
melinaceae); vessels in roots, sometimes also stems and
leaves, with scalariform or simple perforation; flowers mostly
petaloid; stamens dehiscing with longitudinal slits or with api-
cal pores; septal nectaries sometimes present; ovules anatro-
pous or atropous, bitegmic and crassinucellar; endosperm
development Helobial (Nuclear in Commelinaceae); seed coat
formed by both integuments; endosperm copious, containing
aleurone and starch. Four families
Perianth of 4 segments; raphides restricted to tapetal cells;
[calcium oxalate styloids present; flowers in spikes or com-
pound spikes, hypogynous, zygomorphic; stamen 1; necta-
ries lacking; seeds operculate; endosperm development
Helobial; endosperm nonmealy, containing starch, protein
and oil]. 4/4-5, Australia, SE Asia Philydraceae
Perianth segments 6(-3); raphides abundant
2 Leaf with a closed sheath; stamens 6(-3), up to 4 reduced
to staminodes; stomata tetra-(hexa-)-cytic; nectaries
lacking; [blade well defined, commonly somewhat succu-
lent; anthers and pollen often dimorphic]. 41/650, widely
distributed in tropical and warm-temperate regions
Commelinaceae
2 Leaf sheath not closed, often inconspicuous; stamens 6-1;
stomata paracytic; nectaries often present
3 Terrestrial perennials; leaves linear, not petiolate;
flowers hypogynous or epigynous, not in pairs, not
heterostylous; stamens 6 or 3; pollen sulcate (-derived);
septal nectaries mostly present; [the roots and
rootstocks often red; seeds glabrous or hairy]. 13/100,
N and S America, S Africa; Australia and New Guinea
Haemodoraceae
3 Helophytes or aquatics; leaves either simple and linear
or differentiated into petiole and blade; flowers hyp-
ogynous, usually paired, often triheterostylous;
stamens 6 (3, 1); pollen bisulcate or trisulcate; septal
nectaries confined to two genera. 9/33, centered in
(sub)tropical America, but widely distributed and
introduced in warmer regions Pontederiaceae
6. Xyridales
Perennial or annual herbs; silica bodies sometimes present;
raphides lacking; flowers heterochlamydeous; ovules anatro-
3
pous and crassinucellate or mostly atropous and tenuinucel-
late; endosperm development Nuclear (in Xyridaceae Nuclear
and Helobial); endosperm with aleurone and copious starch;
embryo appressed to seed coat. Three families
Aquatics or marsh plants; leaves dispersed, 1-veined, with-
out distinct sheath, often apically bidentate; [flowers soli-
tary; anthers dehiscing apically; seeds operculate; pollen
sulcate]. 1/4-10, tropical and warm-temperate America and
W Africa Mayacaceae
Terrestrial or marsh plants; leaves rosulate, apically pointed
or obtuse, with a well-differentiated sheath; veins, if dis-
tinct, more than 1
2 Flowers in button-shaped disk-like heads on thin
peduncles, the heads often compound in more com-
plex inflorescences; pollen spiraperturate; ovary 2-3-
locular, each locule with 1 atropous apical-pendulous
ovule. 10/700-1400, mainly inS America but widely dis-
tributed in tropical and warm-temperate regions
Eriocaulaceae
2 Flowers in spikes or clusters of spikes; pollen sulcate
(-derived); ovules several per locule (if 1, then anatro-
pous and crassinucellate)
3 Leaf bases symmetric; silica bodies lacking; anthers
dehiscing by longitudinal slits; styles often appen-
daged; microsporogenesis successive. 5/300, mainly in
S America, but of wide tropical and north-temperate
distribution Xyridaceae
3 Leaf bases asymmetric; epidermal cells containing
silica bodies; anthers dehiscing by (sub)apical pores;
styles not appendaged; microsporogenesis simulta-
neous. 16/80, northern S America, one sp. W African
Rapateaceae
7. Juncales
Rhizomatic or caespitose herbs; centromeres often diffuse;
silica bodies often present; raphides lacking; stomata
paracytic, rarely tetracytic; vessels usually in all vegetative
organs, with scalariform and/or simple perforation; leaves
mostly tristichous; flowers basically trimerous; perianth
scaffy, but often bristle-like or reduced; microsporogenesis
simultaneous; ovary superior, 1-3-locular, with 1-several
ovules per locule; endosperm development Helobial or
Nuclear; endosperm starchy. Three families
1 Pollen in cuneate, 4-ulcerate "cryptotetrads"; ovary 1-
ovulate; [silica bodies often present; endosperm develop-
ment nuclear; perennials and annuals of manifold growth
form]. 104/5000, cosmopolitan Cyperaceae
Pollen in tetrahedral tetrads; ovary with 3-many ovules
2 Silica bodies lacking; vascular bundles of leaves of
ordinary structure with abaxial phloem; inflorescences
various but never flowers in globose heads; usually
plants from creeping rhizomes; [endosperm formation
Helobial]. 8/350, cosmopolitan Juncaceae
2 Silica bodies present in epidermis and parenchyma; vas-
cular bundles in leaf transsection in pairs above each
other with facing phloem portions; flowers aggregated in
dense heads; perennials from upright rhizomes. 1/3,
northern S America Thurniaceae
8. Typhales
Rhizomatous marsh plants; silica bodies and calcium oxalate
including raphides present; vessels in all vegetative organs,
4 Conspectus of Families Treated in this Volume

with scalariform perforation; stomates paracytic; flowers cushion-forming plants. 3/35, centered in Australia,
unisexual, in compound inflorescences; microsporogenesis extending to SE Asia and southernmost S America
successive; pollen ulcerate; gynoecium pseudomonomerous; Centrolepidaceae
ovule 1 per locule, anatropous, pendulous; endosperm devel-
Of unknown affinity:
opment Helobial; mature seeds with mealy endosperm and a
thin perisperm. Only one family. 2/16-30, widely distributed
Typhaceae
10. Hydatellaceae

9. Poales Small herbs; silica bodies and raphides lacking; vessels in

roots and stems; staminate flowers of a single stamen; pollen
sulcate( -derived); pistillate flowers a pseudomonomerous
Grasslike plants, often rhizomatous; epicuticular wax non-
pistil with 1 pendulous ovule; endosperm formation Nuclear;
oriented scales, rarely compound rodlets; vessels with simple
endosperm in mature seeds largely replaced by starchy
and/or scalariform perforation in roots, stems, and sometimes
perisperm. 2/10, Australia, New Zealand and India
also the leaves; silica bodies often deposited in epidermal cells;
Hydatellaceae
raphides mostly present (lacking in most Restionaceae and
Centrolepidaceae); leaves distichous, differentiated into
sheath and blade, with paracytic stomata; flowers often in
spikelets combined in paniclelike inflorescences; microsporo-
genesis mostly successive (simultaneous in Flagellaria);
gynoecium syncarpous; pistil 3-1-locular, the 3 carpels
ending with free styluli or less often with a single style; ovary
locules with a single apical (in grasses lateral) ovule; ovule
atropous or (only grasses) amphitropous; pollen ulcerate, the
aperture often annulate; endosperm development Nuclear;
endosperm starchy, of compound or simple starch grains;
tannin in integument present or (Flagellaria, grasses) absent.
Seven families
Flowers hermaphroditic; anthers 4-sporangiate; fruit
indehiscent
2 Flowers in spikelets; tepals 3 or fewer, generally small
("lodicules"); ovary !-locular; embryo asymmetrical,
lateral at the micropylar end, outside the endosperm;
[epidermal cells long/short]. 660/9000, worldwide
Gramineae
2 Flowers not in spikelets; tepals 6; ovary 3-locular; embryo
lenticular, centrally placed at micropylar end of
endosperm
3 Solid-stemmed Hanas; perianth somewhat petaloid;
leaf blade circinately inrolled in bud, apically cirrhose;
sheath closed; epidermal cells similar; [microsporo-
genesis simultaneous]. 1/4. Old World tropics
Flagellariaceae
3 Hollow-stemmed, coarse herbs; perianth chaffy; leaves
plicate in bud, not cirrhose; sheath open; epidermal
cells long/short. 1/2, E Asia, Pacific Joinvilleaceae
Flowers unisexual; anthers 2-4-sporangiate; fruit dehiscent
4 Anthers 4-sporangiate; [tepals 3 + 3, bractlike, chaffy;
rushlike plants with photosynthetic culms]
5 Stem with continuous cylinder of sclerenchyma
(Hopkinsia and Lyginia) Restionaceae p.p.
5 Stem lacking a continuous cylinder of sclerenchyma
6 Ovary 2-locular; stamens 4-6; pollen not scro-
biculate. 2/2, SW Australia Ecdeiocoleaceae
6 Ovary 3-locular; stamens 3; pollen scrobiculate. 1/7,
W Australia Anarthriaceae
4 Anthers 2-sporangiate
7 Tepals 3 + 3; staminate flowers with 3 stamens; ovary
1-3-locular, with 1-3 styluli, sometimes fused into a
style; wiry, jointed herbs. 55/490, Southern Hemi-
sphere, concentrated in SW Africa and SW Australia,
extending northward to Malesia Restionaceae p.p.
7 Flowers naked; staminate flowers of but 1 stamen;
pistillate flower a solitary, !-locular pistil; tufted or
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Molecular Systematics
Chase, M.W., Soltis, D.E., Olmstead, R.G., Morgan, D., Les,
D.H., Mishler, B.D., Duvall, M.R., Price, R.A., Hills, H.G.,
Qiu, Y.-L., Kron, K.A., Rettig, J.H., Conti, E., Palmer, J.D.,
Manhart, J.R., Sytsma, K.J., Michaels, H.J., Kress, W.J.,
Karol, K.G., Clark, W.D., Hedren, M., Gaut, B.S., Jansen,
R.K., Kim, K.-J., Wimpee, C.F., Smith, J.F., Fumier, G.R.,
Strauss, S.H., Xiang, Q.-Y., Plunkett, G.M., Soltis, P.S.,
Swensen, S.M., Williams, S.E., Gadek, P.A., Quinn, C.J.,
Eguiarte, L.E., Golenberg, E., Learnjr., G.H., Graham, S.W.,
Barrett, S.C.H., Dayanandan, S., Albert, V.A. 1993.
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Chase, M.W., Duvall, M.R., Hills, H.G., Conran, J.G., Eguiarte,
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Hoot, S. 1995. Molecular systematics ofLilianae. In: Rudall,
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Commeliniflorae-Zingiberiflorae complex in monocots
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80: 987-998.
 Acoraceae 7

Acoraceae foliage leaves. The continuation shoot arises in the

axil of the leaf preceding the spathe. The last leaf
J. BoGNER and S.J. MAYO of the flowering shoot (article) is called the spathe,
following Engler (1877), although it resembles a
foliage leaf morphologically. The lower part of the
spathe appears to be congenitally adnate to the
axis of the inflorescence proper. Two separate vas-
cular systems occur in the peduncle, one corre-
sponding to the spathe and the other to the
Acoraceae Martinov, Tekhno-Bot. Slovar: 6 (1820). inflorescence axis. Ray (1988) rejected the idea
that the leaf associated with the spadix is a spathe,
Herbs, rhizome repent, much branched, lacunose, preferring to interpret it as a sympodialleaf. Ac-
with aromatic oil cells; leav,es distichous, unifacial, cording to him, the shoot of Acarus calamus
ensiform, not differentiated into petiole and shows anisophyllous sympodial growth; the con-
blade, venation strictly parallel; intravaginal tinuation shoot arises sylleptically and has a
squamules present in leaf axils; inflorescence soli- prophyll that resembles a foliage leaf.
tary, terminal, borne laterally on leaflike axis;
spathe longer than spadix, erect, persistent, ap- VEGETATIVE ANATOMY (based on French
pearing merely as extension of leaflike peduncle; 1997). Primitive vessels are present in the roots
spadix conic-digitiform or slender and tail-like, and rhizome (Carlquist and Schneider 1997). The
densely flowered throughout; flowers bisexual, roots have large, schizogenous intercellular spaces
bractless, 3-merous, perigoniate; tepals in 2 in the cortex. Idioblastic oil cells are present in the
whorls of 3, thin, fornicate; stamens in 2 whorls root cortex. The inner boundary of the stem cortex
of 3; filaments linear-elongate; anthers introrse; is delimited by an endodermis with Casparian
thecae rounded-elliptic, subopposite, opening by strips (absent in the majority of Araceae genera).
longitudinal slit; gynoecium obconic-oblong, The endodermis encircles the central cylinder, in-
equalling the tepals, 2-3-locular; placenta apical; terrupted only where the leaf traces depart. Simple
ovules several per locule, pendent, atropous; collateral axial vascular bundles are absent in
both integuments bearing trichomes, inner in- Acarus, which has amphivasal axial bundles. The
tegument longer than outer and forming the 3-dimensional organization of the vascular system
micropyle; stigma minute, sessile; berry oblong- corresponds to the "Rhapis pattern", or Pattern 1
obovoid with thin, leathery pericarp, whitish with of French and Tomlinson (1981). The axial bun-
brownish stigma remnant when fresh, soon drying dles tend to become part of a branched network of
to straw-brown, enclosed by tepals, 1-5( -9)- peripheral vascular bundles after the departure of
seeded; seeds oblong to ellipsoid, with perisperm the leaf traces; this pattern is also found in some
and abundant endosperm, testa light brown, palms and Araceae. The pattern of insertion of the
foveolate or smooth; embryo axile, cylindric. bud vascular traces is unusual and differs from the
A family consisting of one genus with 2( -4) patterns observed in Araceae. A complete central
species, distributed throughout the northern cylinder of bud traces is formed which migrates
hemisphere and occurring in temperate to tropi- through the cortex and then separates into a
cal latitudes; introduced and naturalized in number of bundles that are inserted over the
Europe. surface of the central cylinder of the main axis.
The leaves have a central region oflarge air cavi-
VEGETATIVE MORPHOLOGY. The leaves are disti- ties beneath a chlorenchyma of 3-5 layers of iso-
chous, equitant and unifacial (Kaplan 1970, 1973). diametric cells. A chlorophyll-less hypodermis is
The leaf form arises in a way similar to that of the present. The leaf venation is parallel, with minor
phyllodes of Acacia, by the development of a sec- transverse veins joining the main, longitudinally
ondary, adaxial meristem in the leaf primordium oriented system. This corresponds to the typical
and the suppression of the marginal meristems. monocot venation type (Ertl 1932). Leaf variega-
Intravaginal squamules are present in Acarus tion due to periclinal chimeras occurs in Acarus
(Irmisch 1874; Engler 1877) as in a few genera of and has been studied by Stewart and Derman
Araceae, Spirodela in Lemnaceae, and almost (1979).
all Helobiae (French 1997). Axillary branches Calcium oxalate raphides do not occur in
develop from proleptic buds and begin with a Acarus, but other forms of calcium oxalate have
cataphyllary, 2-keeled prophyll, followed by been reported, e.g. prisms (French 1997). Latici-
8 Acoraceae
f12j
~K
Fig. lA-M. Acoraceae. Acarus calamus. A Habit. B Base of
plant and rhizome. C Detail of leaf venation. D Inflorescence.
E Detail of culm. F Detail of spadix. G Flower. H Flower,
perianth removed. I Pistil, longitudinal section. J Berry. K
Berry, transverse section. L Immature seed. (Orig.)
fers are absent (Engler 1905; French 1988). The
ethereal oils and tannins occur in idioblasts, which
have been studied ultrastructurally (Amelunxen
and Gronau 1969).
FLOWER ANATOMY. Unicellular trichomes are
present on both ovule integuments but most
prominently so on the outer integument in A.
gramineus (French 1987). The inner integument
forms the micropyle and is much longer than the
outer integument. The endothecial thickenings
have a unique, stellate pattern (French 1985). The
ovary walls lack vascular bundles and the vascular
supply of the gynoecium is restricted to an axile
system which supplies the placentas.
 Acoraceae
EMBRYOLOGY. Both perisperm and endosperm
are present. The perisperm consists of a single cell
layer. The anther tapetal cells are 2-4-nucleate.
The anther tapetum is secretory. The endosperm
formation is cellular and, unlike in the Araceae,
there is no chalaza! basal apparatus.
SEEDLING MoRPHOLOGY. The cotyledon is nar-
rowly cylindric and unifacial and the distal part is
haustoria! as in other monocots. The cotyledonary
sheath is short with very narrow, membranous
margins. The hypocotyl is not visible externally.
The root collar bears fairly long, dense rhizoids.
The primary root elongates to approximately the
same length as the cotyledon and then aborts
without branching. The first shoot born roots arise
on each side of the cotyledonary sheath, each with
a short coleorrhiza. The primary leaf is ensiform
and unifacial, closely resembling the adult form.
The seedling morphology resembles that of the
Melanthiaceae, Juncaceae, and Typha more than
that of Araceae (Tillich 1985, 1995).
PALYNOLOGY. The pollen grains are sulcate to
subulcerate, ellipsoid, small [(15-) 18( -20 !liD)];
the exine is shallowly and remotely or more
densely foveolate, otherwise psilate; the apertural
exine is subpsilate (Grayum 1984).
KARYOLOGY (based on Petersen 1989). Acarus
gramineus is known only in diploid populations
(2n = 22, 2n = 24). In Acarus calamus, the dip-
loids (2n = 22 and 2n = 24) are known to occur
naturally only in N America. Triploids (2n = 36)
are characteristic of the European, Near East and
some Indian populations and also occur in eastern
N America. The N American triploids are prob-
ably the result of introductions from Europe by
early settlers. Tetraploids (2n = 44, 2n = 48) are
known only from Asia (also named A. triqueter or
A. calamus var. angustatus).
POLLINATION BIOLOGY. The pollinators or polli-
nating agency of Acarus are unknown; both ento-
mophily and anemophily have been suggested,
but entomophily appears more likely.
FRUITS AND SEED. The fruit a berry and never
capsular, as has been previously reported
(Bochenska and Kozlowski 1974). The seed con-
tains both endosperm and perisperm, and the
testa is foveolate (A. calamus) or smooth (A.
gramineus ). In A. gramineus the trichomes present
on both integuments persist in the seed as bristles.
9
DISPERSAL. Pieces of rhizome are easily dis-
persed by water along rivers. In particular, the
sterile triploid cytotype of Acarus calamus has
been dispersed by this means. The seeds are also
dispersed by water along stream margins. Acarus
calamus was widely dispersed in Europe and parts
of N America by human agency due to its medici-
nal importance, and later became naturalised in
many regions.
PHYTOCHEMISTRY. The dried rhizomes of A. ca-
lamus have been used for centuries as a source of
medicinal compounds. Substances extracted from
the rhizome have been shown to have sedative,
analgetic, insecticidal, and antibacterial proper-
ties. Compounds which have been identified from
the rhizomes include acorin and polyphenols
localized in specialised idioblasts, and varying
concentrations of ethereal oils. The precise com-
position of the ethereal oils varies according to the
geographical source and cytotype of the plant
material (Wulff 1954; Wulff and Stahl 1960);
phenylpropanoids (asarone, methyleugenol),
monoterpenes and sesquiterpenes have been
identified from the ethereal oil. The composition
of ethereal oils in the leaves of A. calamus differs
from that of the rhizome, but also shows a clear
correlation with the cytotype. The rhizomes of
Acarus gramineus also contain ethereal oils, in
which asarone is the main component (Hegnauer
1963, 1986).
AFFINITIES. Acarus has long been considered a
member of Araceae, but Grayum (1987, 1990) has
presented a convincing case for its removal to a
separate family. The long list of significant charac-
ters by which Acoraceae and Araceae are distin-
guished also strongly suggests that they are not
even sister taxa. The presence of ethereal oil cells,
absence of raphides, presence of secretory anther
tapetum, presence of perisperm and dicotlike cel-
lular endosperm development suggest a link to
monocotlike dicot families such as Piperaceae and
Aristolochiaceae. The molecular studies of Duvall
et al. (1993) and Chase et al. (1995) now indicate
that Acarus is sister to all other monocots and
could be considered the most primitive living
monocot taxon.
DISTRIBUTION AND HABITATS. Acarus is found
mainly in the northern hemisphere, from the tem-
perate zone to the tropics; Grayum (1987) reports
records from south of the Equator in Celebes and
New Guinea.
10
Acorus is a herbaceous helophyte which forms
large dense stands due to its extensively branched
rhizomatous stems. It grows along the margins of
rivers, streams, ponds, lakes, ditches, standing or
slowly running water or in swampy places. It is
found from sea level to llOOm in central Europe
and up to 2600 m in China.
PARASITES. The common occurrence of the rust
Uromyces sparganii (at the same stage of its life
cycle) in Acorus and Sparganium (Parmelee and
Savile 1954) has, for some authors, suggested an
affinity between these two genera. (However,
serological studies failed to show a close affinity
between Typhales and other monocot groups
including Acorus, see Lee and Fairbrothers 1972).
FossiL RECORD. The genus Acorus is documented
from the Eocene by the fossil A. brachystachys
(Heer 1870) with parts of the shoot, leaf and spadix
but doubtful (Z. Kvacek, pers. comm.). Acorites
heeri (Berry) Crepet (syn. Acorus heeri Berry) from
the Eocene of N America is also close to Acorus
(Crepet 1977). Acoropsis eximia (syn. Acoropsis
minor) does not belong to the Acoraceae; this
infructescence from the Baltic Amber (Eocene) is a
member of the tribe Monstereae of the Araceae
subfamily Monsteroideae (Bogner 1976).
CoNSERVATION. The species are apparently not
threatened. Both are easy to grow, and A. calamus
is especially prone to naturalise and under
favourable conditions becomes quite weedy.
EcoNOMIC UsEs. Dried rhizomes ("rhizoma
calami") have been used for centuries and were
known to the ancient Egyptians, Greeks and
Romans. Calamus oil ("tinctura calami",
"extractum calami") was used for medicaments
and is still today of some importance. The
chopped rhizomes are still used as a tea to treat
stomach ailments. The rhizomes were also for-
merly used for perfume and today are still used to
make a liqueur. The bitter rhizomes of Acorus
gramineus have also been used medicinally. Both
species are cultivated as plants suitable for wet
sites (bog gardens, ponds, aquatic gardens, etc.)
and cultivars with variegated leaves are popular.
Only one genus:
Acorus L.
Acarus L., Sp. Pl.: 324 (1753); Engler in Pflanzenreich IV.23B:
308-313 (1905); Huttleston, N. Am. Arac.: 16-30 (1953);
Grayum, Taxon 36: 723-729 (1987).
Acoraceae
Characters as for family.
Selected Bibliography
Amelunxen, V.F., Gronau, G. 1969. Untersuchungen an den
Gerbstoffzellen der Niederblatter von Acarus calamus L.
Cytobiologie 1: 58-69.
Bochenska, !., Kozlowski, J. 1974. Por6wnawcze badania nad
budowa owoc6w gatunk6w z rodziny obrazkowatych
(Araceae) wystepujacych w Polsce. Herba Pol. 20: 3-10.
Bogner, J. 1976. Die systematische Stellung von Acoropsis
Conwentz, einer fossilen Aracee aus dem Bernstein. Mitt.
Bayer. Staatssamml. Palaontol. Hist. Geol. 16: 95-98.
Carlquist, S., Schneider, E.L. 1997. Origins and nature of
vessels in Monocotyledons. I. Acorus. Int. ]. Plant Sci.
158(1): 51-56.
Chase, M.W. et al. 1995. See general references.
Crepet, W.L. 1977. Investigations of angiosperms from the
Eocene of North America: an aroid inflorescence. Rev.
Palaeobot. Palynol. 25: 241-252.
Duvall, M.R., Learn G.H., Jr., Eguiarte, L.E., Clegg, M.T. 1993.
Phylogenetic analysis of rbcL sequences identifies Acarus
calamus as the primal extant monocotyledon. Proc. Nat!.
Acad. Sci. USA 90: 4641-4644.
Engler, A. 1877. Vergleichende Untersuchungen tiber die
morphologischen Verhaltnisse der Araceae. II. Theil. Ueber
Blattstellung und Sprossverhaltnisse der Araceae. Nova
Acta Ksl.Leop.-Carol.-Dtsch. Akad. Naturforsch. 39(4): 170,
t.3 fig. 11.
Engler, A. 1905. Araceae-Pothoideae. In: Engler, A. (ed.),
Pflanzenreich IV.23B: 308-313. Leipzig: W. Engelmann.
Ertl, P.O. 1932. Vergleichende Untersuchungen tiber die
Entwicklung der Blattnervatur der Araceen. Flora 126: 118-
120.
French, J.C. 1985. Patterns of endothecial wall thickenings in
Araceae: subfamilies Pothoideae and Monsteroideae. Am. J.
Bot. 72: 472-486.
French, J.C. 1987. Structure of ovular and placental trichomes
of Araceae. Bot. Gaz. 148: 198-208.
French, J.C. 1997. Vegetative anatomy. In: Mayo, S.J., Bogner,
]., Boyce, P.C. (eds.) The genera of Araceae. Royal Botanic
Gardens, Kew.
French, J.C., Tomlinson, P.B. 1981. Vascular patterns in stems
of Araceae: subfamily Pothoideae. Am. J. Bot. 68: 713-
729.
Grayum, M.H. 1984. Palynology and phylogeny of the Araceae.
Ph. D. dissertation. Amherst: Univ. Massachusetts.
Grayum, M.H. 1987. A summary of evidence and arguments
supporting the removal of Acarus from the Araceae. Taxon
36: 723-729.
Grayum, M.H. 1990. Evolution and phylogeny of the Araceae.
Ann. Mo. Bot. Gard. 77: 628-697.
Heer, 0. 1870. Diemiocene Flora und Fauna Spitzbergens.
Kong!. Svenska Vetensk.-Akad. Hand!. 8(7): 1-98.
Hegnauer, R. 1963, 1986. See general references.
Irmisch, T. 1874. Beitriige zur vergleichenden Morphologie
der Pflanzen. 5. Abtheilung. Ueber einige Aroideen. Abh.
Naturforsch. Ges. 13: 1-48, tt. xiv-xix.
Kaplan, D.R. 1970. Comparative foliar histogenesis of Acarus
calamus and its bearing on the phyllode theory of mono-
cotyledonous leaves. Am. J. Bot. 57: 331-336.
Kaplan, D.R. 1973. Comparative developmental analysis of
heteroblastic leaf series of axillary shoots of Acarus calamus
L. (Araceae). Cellule 69: 253-290.
 Acoraceae/ Alismataceae
Lee, D.W., Fairbrothers, D.E. 1972. Taxonomic placement of
the Typhales within the monocotyledons: preliminary sero-
logical investigation. Taxon 21: 39-44.
Mayo, S.j., Bogner,)., Boyce, P.C. 1997. The genera of Araceae.
I-XII, 1-370. Royal Botanic Gardens, Kew.
Parmelee, J.A., Savile, D.B.O. 1954. Life history and relation-
ships of the rusts of Sparganium and Acorus. Mycologia 46:
823-836.
Petersen, G. 1989. Cytology and systematics of Araceae. Nord.
J. Bot. 9: 119-166.
Ray, T.S. 1988. Survey of shoot organization in the Araceae.
Am. J. Bot. 75: 56-84.
Stewart, R.N., Derman, H. 1979. Ontogeny in monocotyledons
as revealed by studies of developmental anatomy of peri-
dina! chimeras. Am. J. Bot. 66: 47-58.
Tillich, H.-J. 1985. Keimlingsbau und verwandtschaftliche
Beziehungen der Araceae. Gleditschia 13: 63-73.
Tillich, H.-J. 1995. Seedlings and systematics in monocotyle-
dons. In: Rudall, P.J., Cribb, P.J., Cutler, D.F., Humphries,
C.j. (eds.) Monocotyledons: systematics and evolution, Vol.
1. Royal Botanic Gardens Kew, pp 303-352.
Wulff, H.D. 1954. Zur Zytologie, geographischen Verbreitung
und Morphologie des Kalmus. Arch. Pharm. 287: 529-541.
Wulff, H.D., Stahl, E. 1960. "Chemische Rassen" bei Acorus
calamus. Naturwissenschaften 47: 114.
11
Alismataceae
R.R. HAYNES, D.H. LEs, and
L.B. HOLM-NIELSEN
Alismataceae Ventenat, Tab!. Regne Veg. 2: 157 (1799), nom.
cons.
Monoecious, polygamous, or dioecious annual
or perennial herbs with milky juice, glabrous to
stellate pubescent, with floating leaves or emer-
gent, in fresh or brackish waters. Roots fibrous,
septate or aseptate, at base of stem or lower
nodes. Stems short, erect, cormlike, often with
rhizomes, the rhizomes occasionally terminated
by tubers, the apices without turions. Leaves
basal, sessile, or petiolate; petioles terete to trian-
gular, with sheathing base, the sheath without
auricles; blade linear, lanceolate, ovate to rhom-
boid, with or without pellucid dots or lines, the
margins entire or undulate, the apex obtuse,
acute, or acuminate, the base either without
basal lobes and attenuate, or otherwise truncate,
cordate, sagittate, or hastate, with parallel
primary veins and reticulate secondary veins.
Inflorescences scapose, mostly erect, rarely
floating, forming racemes or panicles by ver-
ticillate branching, rarely umbellate, without
a subtending spathe, bracteate, the bracts
whorled, linear, entire, obtuse to acute. Flowers
hypogynous, perfect or imperfect, subsessile to
long-pedicellate; perianth actinomorphic, of 6
separate segments in 2 whorls, the outer 3
sepal-like, green, persistent, erect, and enclos-
ing flower and fruit or spreading to reflexed, the
inner 3 petal-like, delicate, caducous; stamens
free, 6, 9, or numerous; if 6, then in pairs alter-
nating with the petals; if 9, then in 2 whorls
with outer whorl of 6 and inner whorl of 3; if
many, densely but irregularly spaced; anthers
tetrasporangiate, extrorse, basifixed or versatile,
dehiscing by longitudinal slits; carpels 6-many,
free, in 1 whorl or irregularly spaced, each
with 1( -2) anatropous basal ovules, rarely
(Damasonium) 2-many on marginal placentae;
styluli terminal or lateral; stigma linear. Fruits
achenes or rarely follicles, mostly numerous.
Seeds 1-few, U-shaped; endosperm absent in
mature seeds.
A family of 12 genera and ca. 80 species of
subcosmopolitan distribution.
12 Alisrnataceae

VEGETATIVE MORPHOLOGY. The Alismataceae

are all marsh plants or aquatic herbs with erect or
floating leaves and grow as helophytes or hydro-
phytes in fresh water. The plants may be terrestrial
with the axis buried in water-logged swamp or
marsh soils (Figs. 2,3).
The nonchlorophyllous axes are short and erect
(corms) or prolonged (rhizomes) and sometimes
creeping and elongate (stolons). They are often
covered by the remains of vascular bundles from
withered petioles. The roots are branched and
rarely develop distal tubers. Stem branching is
sympodial. The sympodium continues from the
axil of the last leaf of the rosette, or the terminal
inflorescence or side buds are sometimes modified
into pseudostolons.
The leaves are distichous, spirodistichous, or
spirally arranged, normally in basal leaf rosettes.
The leaves are usually long-petiolate and emer-
gent with a broad lanceolate to cordate or hastate
blade, but linear or ribbonlike submerged leaves
occur. The blades are mostly erect, but floating
blades occur, e.g., in Luronium and Echinodorus.
The petioles are sheathing and triangular or terete.
The few primary veins are convergent, and a
closed reticulum is formed by the secondary and
tertiary veins. The terminal inflorescence arises
from a bifurcation of the vegetative meristem. The
inflorescences are always emergent, erect panicles,
with rather distant whorls of sometimes long-
pedicellate flowers; the panicles are often reduced
to spikes, pseudospikes, or even umbel-like inflo-
rescences. Squamules are present as a palisadelike
series of scales (Tomlinson 1982).
Starch is commonly found in the rhizome and
along larger vascular bundles in the petiole.

VEGETATIVE ANATOMY. The anatomy of all 11
genera has been studied by several authors and
summarized by Tomlinson (1982). Hairs are gen-
erally absent, but simple unicellular hairs, hair
bases, or stellate hairs are found in a few species of
Echinodorus, Sagittaria, and Limnophyton. The
epidermis of submerged leaves is often chloro-
phyllous, and paracytic stomata are common on
the leaf surfaces. Hydropoten are frequent on sub-
merged or floating leaves. The roots, stems, and
petioles are extensively aerenchymatous, and
the air spaces are traversed by well-developed
transverse diaphragms, which are absent in
Echinodorus and Alisma. Secretory ducts (latici-
fers) containing a milky juice are common in the
leaf and stem, but tanniniferous deposits are rare.
Crystals are frequent in the leaf mesophyll. The
vascular bundles of the peduncle and petioles in-
'
K
Fig. 2A-K. Alisrnataceae. A-D Ranalisma rostra tum. A Habit.
B One-flowered pseudo-umbel. C Fruit in longitudinal
section. D Achene, longitudinal section. E R. humile, habit.
F-1 Damasonium polyspermum. F Habit. G Flower. H Follicles,
two of which in longitudinal section. I Seed. J, K Luronium
natans. J Habit. K Achene. (Takhtajan 1982)
elude a well-developed protoxylem lacuna, with a
U-shaped arrangement of metaxylem. Vessels are
restricted to the roots, where they occur as single
central strands, mostly with simple perforations.
INFLORESCENCE STRUCTURE. The inflorescences
are either paniculate with !-several branches or
 Alismataceae
Fig. 3A-K. Alismataceae. A-E Sagittaria sagittifolia. A Plant
with submersed and floating leaves. B Plant with aerial leaves.
C Fruiting aggregate. D Achene. E Stamen. F-H S. teres.
F Habit. G Achene. H Stamen I, J S. guayanensis. I Habit.
J Achene. (Takhtajan 1982)
spicate and have a low basal internode and 1 or
more successive pseudowhorls. The pseudowhorls
are separated by further long internodes and nor-
mally have 3 bracts subtending on either side
branches or flowers. The meristems of the inflo-
rescences are often (under water stress) changed
into vegetative meristems, producing richly
13
proliferating, occasionally even stoloniferous,
pseudoinflorescences. Ranalisma has a sympodial
inflorescence type (Charlton and Ahmed 1973).
FLORAL STRUCTURE. The flowers are perfect, or
imperfect by abortion of either stamens or carpels
in Sagittaria, Limnophyton, and Burnatia. Plants
of these latter genera are either monoecious
(Sagittaria), polygamous (Limnophyton, Sagit-
taria), or dioecious (Burnatia). The pedicel often
expands into a receptacle. The flowers are trimer-
ous with 2 alternate perianth whorls differentiated
into sepals and petals. The petals are delicate,
white, pink, or purple. In Burnatia the petals are
reduced or absent.
The androecium has 3, 6, or many stamens. The
gynoecium consists of free carpels and is, in
general, connate basally. The carpels are often
laterally compressed, with short styluli, and the
stigmas are apical or slightly decurrent and of the
Dry type. Nectar secretion occurs from the base of
petals, stamens, and carpels.
The development of multistaminate and mul-
ticarpellate flowers is based on a strictly trimerous
pattern, with the lateral and centrifugal apposition
of additional stamen and carpel primordia; there
is no spiral sequence (Singh and Sattler 1972,
1973).
EMBRYOLOGY. Anther wall formation is of the
monocotyledonous type. The endothecial thicken-
ings are of the Girdle type, the tapetal cells of the
Plasmodial and Amoeboid type and uninucleate.
Microsporogenesis is successive and the pollen
grains are 3-celled when shed. The archespore
cells function directly as megaspore mother cells,
and the nucellar epidermis becomes irregularly 2-
layered through periclinal divisions of its apical
cells (Davis 1966).
The embryo sac is of the Allium type. In Alisma,
Damasonium, and Luronium endosperm forma-
tion is Nuclear; in Echinodorus, Limnophyton, and
Sagittaria, however, Helobial endosperm forma-
tion occurs (Johri et al. 1992). The latter may rep-
resent the plesiomorphic state. The formation of
the strongly curved embryo follows the Caryo-
phyllad type.
The ripe seeds are curved and exalbuminous
and have horseshoe-shaped embryos that lack
chlorophyll.
PoLLEN MoRPHOLOGY. The pollen grains are
globose to polyhedral and polyforaminate with 9-
29 apertures (Fig. 4). Two minor types can be dis-
tinguished according to sculpture and shape; their
14
Fig. 4. Alismataceae. Caldesia parnassifolia, polyforate spi-
nulose pollen, SEM X2700. (Palynological Laboratory,
Stockholm)
distribution correlates with other traits, as dem-
onstrated by Argue (1976) (see section Subdivi-
sions and Relationships; below). The pollen of 2
out of the 4 species of Caldesia is quite aberrant in
being lense-shaped and (0- )2( -3) porate.
KARYOLOGY. Base numbers of x = 5-13 have
been reported, with 7, 8, and 11 as the most
common (Fedorov 1969). See also section Subdivi-
sions and Relationships, below.
PoLLINATION. Nectar secretion occurs at the
bases of the floral organs, particularly of the
staminodes in the pistillate flowers. In Alisma
plantago-aquatica Daumann (1965) observed
nectar secretion from the flanks of the carpels; the
nectar later accumulates at the base of the fila-
ments. The flowers open between 09.00 and 11.00
h and close in the afternoon, and they are visited
by small Syrphidae and small Diptera. The flowers
are homogamous and self-compatible. The pollen
is mealy and is also dispersed by the wind.
FRUIT AND DISPERSAL. Each carpel develops
into an achene or into a follicle (Damasonium
polyspermum).
The achenes are often keeled or winged, more or
less glandular nutlets, which are dispersed indi-
vidually. The stylar beak is often peristent and is
likely to play a role in dispersal. In some cases, the
individual achenes tend to stick together, but in
Alismataceae
most cases they fall apart at dispersal time. They
are water- or animal-dispersed, but the dispersal
systems are incompletely known.
PHYTOCHEMISTRY. Condensed tannins are re-
corded from leaves of Luronium and Wiesneria
and the root of Echinodorus. C-glycosylflavones
are common in the leaves of Sagittaria.
SUBDIVISIONS AND RELATIONSHIPS WITHIN THE
FAMILY. The 11 genera of Alismataceae have not
recently been investigated in relation to generic
delimitation. A systematic study will, in fact, prob-
ably reveal that several of the presently accepted
genera are congeneric, as Baldellia, Caldesia,
Luronium, Ranalisma, and Echinodorus have a
high degree of similarity. A significant correlation
among the mode of endosperm formation
(Nuclear vs. Helobial), pollen morphology (granu-
lar vs. spinose), and chromsomal base number
(x = ± 7 vs. ± 11) unites Alisma, Luronium,
Damasonia, and Baldellia against the remaining
genera.
AFFINITIES. The generic name Ranalisma recalls
an earlier taxonomic opinion that the Alis-
mataceae evolved from the dicotyledon family
Ranunculaceae. Other than superficial resem-
blances (e.g., apocarpy, numerous stamens), there
has been no support for this hypothesis from any
contemporary phylogenetic study. Vegetative
organization of Ranalisma is typical of other
Alismataceae and also similar to the genus
Hydrocleys of Limnocharitaceae (Charlton and
Ahmed 1973). An appraisal of the systematic posi-
tion of the Alismataceae is presented in several
recent papers (Les et al. 1993; Les and Haynes
1995; Les and Schneider 1995). Phylogenetically,
the family is placed solidly among the relatively
primitive monocots compnsmg superorder
Alismatanae, whose closest sister group may be
the family Araceae.
The Alismataceae, along with Aponogetonaceae,
Butomaceae, and Limnocharitaceae, have been
grouped to comprise the Alismatales (Dahlgren et
al. 1985), one of two orders commonly recognized
in the superorder Alismatanae. The Limno-
charitaceae are believed to represent the family
closest phylogenetically to the Alismataceae
(Dahlgren et al. 1985; Haynes and Holm-Nielsen
1992) and these families have often been merged.
Butomaceae (which have occasionally been
merged with Limnocharitaceae) are believed
to be the next most closely related family to
Alismataceae, with Hydrocharitaceae as the most
 Alismataceae
distant relative in the order (Haynes and Holm-
Nielsen 1992).
Recent work based on DNA sequence data (Les
et al. 1993, 1997; Les and Haynes 1995) confirms
that the Alismataceae and related families within
the superorder Alismatanae have not even a
remote relationship to the Ranunculaceae. The
Limnocharitaceae are indeed the most closely
related group to Alismataceae, to the extent that
molecular data depict the Limnocharitaceae as
derived from within a paraphyletic Alismataceae.
This result explains the high degree of similarity
observed between Alismataceae and Limno-
charitaceae which led Dahlgren et al. (1985) to
accept their distinctness "with great hesitation".
Our reluctance to combine these families is
based on several observations. Presently, only 8 of
11 genera of Alismataceae and two of three genera
of Limnocharitaceae have been surveyed in the
molecular study. Given the present sample of
genera available for analysis, the constraint of
both Alismataceae and Limnocharitaceae as
monophyletic groups adds only three steps to the
molecular-based cladograms (D.H. Les et al.
1997). Furthermore, several characters (laminar
placentation, numerous ovules per carpel, dehis-
cent fruits) effectively separate Limnocharitaceae
from Alismataceae (with one to few ovules per
carpel, basal placentation, and indehiscent fruits).
Thus, the paraphyly of Alismataceae will not be
convincingly established until additional genera
have been surveyed and more comprehensive
phylogenetic analyses have been completed.
DISTRIBUTION AND HABITATS. The distribution
of Alisma is worldwide, while the large genera
Echinodorus and Sagitta ria have their main center
in the western hemisphere. The other genera are
preferably distributed in the Old World tropics or
temperate zones. Notably disjunct is the distribu-
tion of Damasonia (California/Europe/ Australia).
The Alismataceae occur in aquatic habitats, es-
pecially those with stagnant or flowing freshwater.
The plants grow as amphiphytes or helophytes in
marshes or along the shores of lakes and streams;
some species are hydrophytes with floating leaves.
A few species of Echinodorus are forest floor herbs
in "igap6 forest" along neotropical blackwater
streams. Most species are rooted in muddy and
water-logged substrates, and several species occur
as members of floating island associations
growing directly in water-suspended substrate.
PALEOBOTANY. The Alismataceae have been
shown to have occurred in the Tertiary from the
15
Oligocene and Miocene. Fossils from the Lower
and Upper Cretaceous and the Paleocene that
have tentatively been referred to the Alismata-
ceae are inadaequate for proper identification
(Daghlian 1981; Friis 1985).
ECONOMIC IMPORTANCE. The family includes
some tropical water weeds although these are not
nearly so pernicious as some genera of other fami-
lies, e.g., Eichhornia. Sagittaria sagittifolia is
cultivated in Asia for its edible corm. Roots of S.
latifolia were used as a food source by N American
Indians (Cook 1978). Many species of Echinodorus
and Sagitta ria are cultivated commercially for use
as aquarium plants and several species of various
genera are used regularly as pond ornamentals.
Most species are important as food sources to
water animals such as manatee and birds.
KEY TO THE GENERA
1. Perianth not differentiated into calyx and corolla, or
corolla reduced; plants from the Old World 2
- Perianth differentiated into calyx and corolla; plants
w~~re~ 3
2. Plants dioecious; flowers pedicellate; inflorescences
branched; staminate flowers with 6-9 stamens; carpellate
flowers with many carpels and 0-2 staminodia. Africa
12. Burnatia
- Plants monoecious; flowers sessile; inflorescences spicate;
staminate flowers with 3 stamens; carpellate flowers with
3-6 carpels and 3 staminodia. Paleotropicalll. Wiesneria
3. Flowers imperfect or pistillate flowers with a ring of
stamens 4
- Flowers all perfect 7
4. Receptacle convex; achenes laterally flattened; staminate
flowers with many stamens; cosmopolitan 5
- Receptacle flattened; achenes rounded; staminate flowers
with 6-9 stamens. Paleotropical 6
5. Pistillate flowers without stamens. Mostly north temperate
lOa. Sagittaria subgen. Sagittaria
- Pistillate flowers with a ring of functional or reduced
stamens. Mostly tropical
lOb. Sagittaria subgen. Lophotocarpus
6. Achenes unarmed, with lateral air canals; bracts green
8. Limnophyton
- Achenes with spines to 5 mm long, without lateral air
canals; bracts maroon 9. Astonia
7. Carpels on an elongated receptacle. Paleotropical
5. Ranalisma
- Carpels on a convex or flattened receptacle; widespread
8. Fruits follicles, fused together basally into a starlike con-
gregation; seeds 2-many per carpel 1. Damasonium
- Fruit achenes, separate, forming loose echinate heads;
seeds 1 per carpel 9
9. Plants with floating and emersed leaves; achenes few,
inwardly convex. Europe 4. Luronium
- Plants with erect or floating, emersed, or submersed
leaves; achenes mostly many, outwardly convex 10
10. Carpels arranged in a circle around margin of flattened
receptacle 11
- Carpels spread evenly over convex receptacle 12
16 Alismataceae
11. Carpels many; pericarp coriaceous 3. Alisma
- Carpels few; endocarp hard 7. Caldesia
12. Leaves without pellucid markings; inflorescence a few-
flowered umbel. Paleotemperate 2. Baldellia
- Leaves with or without pellucid markings; inflorescence
many-flowered panicles, or rarely umbels. neotropical
6. Echinodorus
I. Damasonium P. Miller. Fig. 2F-I
Damasonium P. Miller, Gard. Diet. Abr. Ed. 4. 1: 28 (1754).
Machaerocarpus Small (1909).
Monoclinous, emergent. Stems short or tuberi-
ferous, rhizomatous. Leaves basal, emerged, erect,
long-petiolate; blade narrow-lanceolate to ovate,
apex acute, base attenuate-cordate. Inflorescence
scapose, verticillate, simple to paniculate. Flowers
long-pedicellate; sepals persistent; petals longer
than sepals, delicate, white; stamens 6; carpels
6-9, free or basally connate; ovules 2-many,
apotropous. Fruits follicular, triangular, adhering
basally, forming a starlike structure; follicles
beaked; seeds sculptured. Three spp., W and S
Europe, temperate regions of Australia and
Tasmania, one in California.
2. Baldellia Parl.
Baldellia Pari., Nuovi Gen. Sp.: 57 (1854).
Monoclinous, emergent. Stems short, erect, te-
rete, rhizomatous. Leaves basal, submersed or
emersed, erect, long-petiolate; petiole aerenchy-
matous; blade linear to narrow-lanceolate, apex
acute, base attenuate. Inflorescence few-flowered,
obscurely paniculate, umbel-like. Flowers long-
pedicellate, sepals short, recurved, persistent;
petals much longer than sepals, orbicular, white to
pink; stamens 6 in pairs opposite petals; carpels
many, free; achenes forming an echinate head,
achenes oblong, elliptic, with 5 costae. 2n = 14, 16,
18, 22. Two spp., Europe, N Africa, and the Canary
Islands.
3. Alisma L.
Alisma L., Sp. Pl.: 343 (1753); Bjorkqvist, Opera Bot. 19: 1-138
(1968), rev.
Monoclinous, emergent. Stems short, rhizoma-
tous. Leaves basal, emersed, erect; petiole aeren-
chymatous; blade oblong-lanceolate-elliptic, apex
acute, base attenuate to cordate. Inflorescence sea-
pose, emergent, verticillate, paniculate. Flowers
long-pedicellate, sepals persistent; petals larger
than sepals, pink or white; stamens 6( -9); carpels
many, free. Achenes forming an echinate head; the
achenes strongly laterally compressed, keeled, the
keels 2-3. 2n = 10, 14, 26, 28. Nine closely related
spp., worldwide.
4. Luronium Raf. Fig. 2K,L
Luronium Raf., Aut. Bot.: 63 (1840).
Monoclinous. Stems short, rhizomatous. Leaves
basal, long-petiolate, floating; blade ovate to
oblong, apex rounded, base rounded. Inflores-
cence scapose, obscurely paniculate, verticillate.
Flowers long-pedicellate; sepals persistent; petals
much longer than sepals, broad-obovate, delicate,
white to pinkish; stamens 6, grouped in pairs op-
posite petals; carpels 6-9(-10). Achenes ovate,
nearly terete, sulcate; seeds not or weakly sculp-
tured. 2n = 42. Only one sp., L. natans (L.) Raf.,
Temperate Europe.
5. Ranalisma Stapf Fig. 2A-D
Ranalisma Stapf in Hook., Ic. Pl.: Tab. 2652 (1900).
Monoclinous, emersed. Stems short, rhizomatous;
stolons occasional. Leaves basal, submersed or
emersed; petiole flattened; blade ovate to linear-
lanceolate, the apex acute, the base acute to at-
tenuate. Inflorescence a 1-3-flowered umbel, the
flowers occasionally replaced by bulbils; sepals
ovate, membranaceous reflexed after anthesis;
petals delicate, white or pinkish; stamens (6-)9-
12; carpels free on an elongated receptacle, to 70.
Achenes laterally compressed, with a dorsal wing,
nearly semicircular, without sculpturing. Two
spp., tropics of Old World.
6. Echinodorus Rich. ex Engelm.
Echinodorus Rich. ex Engelm. in A. Gray, Manual: 460 (1848);
Rataj, Rev. gen. Echinodorus. Prague (1975), rev.; Haynes &
Holm-Nielsen, Fl. Neotrop. 64: 1-112 (1914).
Monoclinous. Annual or perennial, emersed.
Leaves erect, with pellucid markings as dots or
lines or not; emersed leaves petiolate, blade linear
to broadly ovate, margins entire or undulate, apex
acute to acuminate, base truncate to cordate; sub-
mersed leaves mostly sessile phyllodes, margins
entire or undulate, apex acute to acuminate. Inflo-
rescence erect, emersed, racemose or paniculate,
rarely umbelliform. Flowers perfect; pedicels often
elongating after an thesis; sepals reflexed to spread-
 Alismataceae
ing; petals white, larger than sepals; stamens 9-
many; carpels many, achenes terete, often longitu-
dinally costate and glandular. Twenty-seven spp.,
all but one neotropical, three extending to N
America the other N American. Two subgenera,
subgenus Echinodorus, with versatile anthers and
many more than 20 carpels; subgenus Helanthium,
with basifixed anthers and 20 or fewer carpels.
7. Caldesia Parl.
Caldesia Pari., Fl. Ita!. 3: 598 (1858).
Monoclinous, emersed. Stems short, rhizomatous.
Leaves basal, submersed or floating; petiole
aerenchymatous, septate; blade ovate, sub-
orbiculate or subreniform, base truncate to
cordate, apex rounded to acute. Plants producing
hibernacules. Inflorescence scapose, emergent,
paniculate, verticillate. Flowers long-pedicellate,
sepals broad-ovate; petals equal to sepals, broad-
ovate, rounded, white to bluish white; stamens 6-
9( -11); receptacle flattened; carpels 2-9( -20), free.
Achenes obovoid, 1-seeded, with obscure dorsal
keels or sculptured. 2n = 22. Four spp., Europe
through Africa and Asia to Australia.
8. Limnophyton Miq.
Limnophyton Miq., Fl. Ind. Bat. 3: 242 (1856).
Monoecious, emersed. Stems short, rhizomatous.
Leaves basal, emersed; petiole aerenchymatous,
angled; blade sagittate or linear-lanceolate; apex
acute to acuminate; base hastate or attenuate.
Inflorescence scapose, emergent, paniculate, ver-
ticillate, the upper verticil(s) with staminate, the
lower with perfect flowers, bracts green; sepals
broad, refl.exed after anthesis; petals delicate,
larger than sepals, narrow, white; stamens 6,
green; receptacle flattened; carpels ( 10-) 15-
20( -30), free; style beaklike ventral, caducous, or
only the basal part persistent; achenes obovoid,
unarmed, with lateral air canals. Two spp., pre-
dominantly tropical Africa, L. obtusifolium (L.)
Miq. extending to Madagascar and S and SE Asia.
9. Astonia S.W.L. Jacobs
Astonia S.W.L. Jacobs, Telopea 7: 140 (1997).
Monoecious, emersed. Juvenile leaves linear, sub-
mersed, mature leaves sagittate. Inflorescence
with up to 8 verticils, the lower perfect or mixed
perfect and staminate, the upper staminate; bracts
17
maroon; lower verticils maturing first, peduncle
then bending downwards until the nutlets contact
the water surface, the tip of the inflorescence then
bending upward; sepals green, petals cream or
green, constricted at the base; stamens 6, maroon;
carpels 5-15; achenes crowded, spinescent when
mature, lateral air chambers absent. Only one
sp., A. australiensis (Aston) S.W.L. Jacobs, NE
Queensland.
10. Sagitta ria L. Fig. 3
Sagittaria L., Sp. Pl. 2:993 (1753); Bogin, Mem. N.Y. Bot. Gard.
9: 179-233 (1955), rev.; Rataj, Annot. Zoo!. Bot. Slov. Narod.
Muz. Bratislava 76: 1-31, 78: 1-61 (1972), rev.; Haynes &
Holm-Nielsen, Fl. Neotrop. 64: 1-112 (1994).
Monoecious or rarely dioecious, perennial or
rarely annual, submersed, floating-leaved, or
emersed. Roots septate. Stems often with rhi-
zomes. Leaves submersed, floating, or emersed,
entire, sessile or petiolate, petioles terete to trian-
gular, blades present or absent, without pellucid
markings. Inflorescence erect, floating or sub-
mersed, racemose or paniculate, rarely umbel-
liform, the staminate flowers above, the carpellate
below. Flowers mostly imperfect, rarely the lower
perfect, pedicellate; sepals reflexed in staminate
flowers, reflexed to appressed in carpellate
flowers; petals white or rarely with a pink spot or
tinge; stamens 7-many; carpels many; achenes
compressed, often laterally winged, glandular,
dorsally with a conspicuous wing. 2n = 16, 20, 22.
About 25 spp., predominantly western hemi-
sphere genus of two subgenera, subgenus
Sagittaria, with ascending to reflexed sepals and
mostly spreading to ascending pedicels in fruit,
and all flowers imperfect; subgenus Lophoto-
carpus, with appressed sepals and pedicels rc-
curved in fruit, and the upper flowers perfect.
11. Wiesneria M. Micheli
Wiesneria M. Micheli in A. & C. DC., Monogr. Phan. 3: 82
(1881).
Monoecious, emergent. Stems short, rhizomatous.
Leaves basal, submerged or floating; petiole terete;
blade linear-lanceolate or not expanded, apex
acute, base attenuate. Inflorescence scapose,
emergent, racemose, the lower whorls of carpel-
late flowers, the upper whorls of staminate
flowers. Flowers short-pedicellate or sessile; sepals
ovate, persistent; petals white to pinkish, reduced
or smaller than sepals; staminate flowers with 3
18 Alismataceae

stamens, carpels reduced to nectaries; carpellate
flowers with receptacle slightly curved, with 3
staminodes and 3-6 free carpels. Achenes
obovoid, rounded, weakly sculptured, with 3
dorsal keels. Three spp., W and Central Africa,
Madagascar, and India.
12. Burnatia M. Micheli
Burnatia M. Micheli in A. & C. DC., Monogr. Phan. 3: 81
(1881}.
Rautanenia Buchenau (1897).
Dioecious, emersed. Rhizome tuberous. Leaves
basal; petiole terete; blade linear-lanceolate to
ovate, apex acuminate, the base rounded to
attenuate. Inflorescence scapose, paniculate,
branched, verticillate. Flowers small, sepals per-
sistent; petals white or pinkish, reduced or smaller
than sepals; staminate flowers with (6- )9 stamens
and ca. 12 aborted carpels; carpellate flowers with
flattened receptacle, occasionally with 2 stami-
nodes, and 8-20 free carpels. Achenes obovoid,
laterally compressed, forming a small head. Only
one sp., B. enneandra M. Micheli, Tropical Africa.
Haynes, R.R., Holm-Nielsen, L.B. 1994. Alismataceae. Flora
Neotropica 64: 1-112.
Johri, B.M. et al. 1992. See general references.
Les, D.H., Haynes, R.R. 1995. Systematics of subclass
Alismatidae: a synthesis of approaches. In: Rudall, P.J.,
Cribb, P.J., Cutler, D.F., Humphries, C.J. (eds.) Monocotyle-
dons: systematics and evolution. London: Royal Botanic
Gardens, Kew, pp. 353-377.
Les, D.H., Schneider E.L. 1995. The Nymphaeales, Alis-
matidae, and the theory of an aquatic monocotyledon
origin. In: Rudall, P.J., Cribb, P.J., Cutler, D.F., Humphries,
C.J. (eds.) Monocotyledons: systematics and evolution.
London: Royal Botanic Gardens, Kew, pp. 23-42.
Les, D.H., Garvin, D.K., Wimpee, C.F. 1993. Phylogenetic
studies in the monocot subclass Alismatidae: evidence for a
reappraisal of the aquatic order Najadales. Mol. Phylogenet.
Evol. 2: 304-314.
Les, D.H., Cleland, M.A., Waycott, M. 1997. Phylogenetic
studies in Alismatidae II: evolution of marine angiosperms
(seagrasses) and hydrophily. Syst. Bot. 22: 443-463.
Muller, J. 1981. Fossil pollen records of extant angiosperms.
Bot. Rev. (Lancaster) 47: 1-142.
Rogers, G.K. 1983. The Alismataceae in the Southeastern
United States. J. Arnold Arbor 64: 383-420.
Singh, V., Sattler, R. 1972. Floral development of Alisma
trivia/e. Can. J. Bot. 50: 619-627.
Singh, V., Sattler, R. 1973. Nonspiral androecium and gyno-
ecium of Sagittaria latifolia. Can. J. Bot. 51: 1093-1095.
Takhtajan, A.L. 1982. See general references.
Tomlinson, P.B. 1982. Helobiae (Alismatidae). In: Metcalfe
C.R. (ed.) Anatomy of the monocotyledons, Vol. VII.
Oxford: Clarendon Press.

Selected Bibliography

Argue, C.L. 1976. Pollen studies in the Alismataceae with

special reference to taxonomy. Pollen Spores 18: 15 7-201.
Bjiirkqvist, I. 1968. Studies in Alisma L. II. Chromosome
studies, crossing experiments and taxonomy. Opera Bot. 19:
1-138.
Bogin, C. 1955. Revision of the genus Sagittaria. Mem. N.Y.
Bot. Gard. 9: 179-233.
Charlton, W.A., Ahmed, A. 1973. Studies in Alismataceae. III.
Floral anatomy of Ranalisma humile. Can. J. Bot. 51: 891-
897.
Cook, C.D.K. 1978. Alismataceae. In: Heywood, V. (ed.) Flow-
ering plants of the world. Oxford: Oxford University Press.
Cronquist, A. 1981. See general references.
Daghlian, C.P. 1981. A review of the fossil record of monocoty-
ledons. Bot. Rev. (Lancaster) 47: 517-555.
Dahlgren, R.M.T. eta!. 1985. See general references.
Daumann, E. 1965. Insekten- und Windbestaubung bei Alisma
plantago-aquatica L. Ein Beitrag zur experimentellen
Bliiteniikologie. Osterr. Bot. Z. 112: 295-310.
Davis, G.L. 1966. See general references.
Fassett, N.C. 1955. Echinodorus of the American tropics.
Rhodora 57: 133-212.
Fedorov, A.A. 1969. See general references.
Friis, E.M. 1985. Angiosperm fruits and seeds from the Middle
Miocene of Jutland (Denmark). Alismataceae. K. Dan.
Vidensk. Selsk. Bioi. Skr. 24(3): 71-73.
Haynes, R.R., Holm-Nielsen, L.B. 1985. A generic treatment of
Alismatidae in the neotropics. Acta Amazonica 15(1-2):
153-193.
Haynes, R.R., Holm-Nielsen, L.B. 1992. Limnocharitaceae.
Flora Neotropica 56: 1-34.
 Anarthriaceae 19

Anarthriaceae of two sizes, and are arranged in a single ring

inside the chlorenchyma; the sclerenchyma sheath
H.P. LINDER, B.G. BRIGGS, of each bundle is somewhat enlarged and extends
and L.A.S. JoHNSON 1 to the epidermis, thus interrupting the chloren-
chyma. Silica is absent, but calcium oxalate crys-
tals occur in some species.

INFLORESCENCE AND FLOWER STRUCTURE. The

inflorescence structure shows various degrees of
Anarthriaceae Cutler & Airy Shaw, Kew Bull. 19: 489 (1965). branching, with each branch subtended by a bract,
and the flowers pedicellate or sessile. The uni-
Evergreen dioecious plants, caespitose, shortly sexual flowers are relatively unspecialised (Linder
rhizomatous or with branching aerial stems. 1992). The three anthers are bilocular, and the
Culms simple or branching, terete or flattened,
glabrous. Leaves all basal or arising from aerial
stems, with laterally flattened, equitant blades,
with a small ligule at the transition to the sheath,
sheath split to the base. Inflorescence racemose,
with few or many flowers, sometimes with elon-
gated inflorescence branches, the 2 sexes alike.
Flowers not in spikelets, sessile or pedicellate, each
with 1 or 2 short bracts. Perianth of 6 rigid
or scarious-membranous, identical, glumaceous,
glabrous tepals in 2 whorls. Male flowers with 3
stamens opposite the inner tepals, filaments free,
slender; anthers bilocular, exserted, dorsifixed, 2-
lobed, each lobe dehiscing by a longitudinal slit.
Female flowers usually with staminodes (some-
times minute); ovary superior, free, 3-locular with
a single pendulous atropous ovule in each locule,
with 3 elongated papillose styluli, free to the base
or shortly connate to form a short style. Fruit a 3-
locular, bluntly angular capsule opening at the
angles. Seeds ovoid, white, asymmetrical, the
surface colliculate with lines of large angular or
deeply lobed cells imprinted with a striate pattern.
A monogeneric West Australian family with
seven species, largely restricted to oligotrophic
soils.

VEGETATIVE MORPHOLOGY AND ANATOMY. The

Anarthriaceae are evergreen, rushlike plants (Fig.
5), growing to a maximum height of 1m, forming
either tangled clumps or caespitose tussocks,
or occasionally with stilt roots (Pate 1989). The
culms are photosynthetic and green, terete or
flattened. The leaves are variously developed,
equitant and usually laterally flattened. The leaves
and culms of Anarthriaceae are anatomically simi-
lar (Cutler 1969) but the culms with some vascular
bundles in the central tissues. Directly below the
epidermis is a chlorenchymatous layer of peg cells
one to several cells thick. The vascular bundles are Fig. SA-D. Anarthriaceae. Anarthria scabra. A Plant with
staminate inflorescence. B Staminate flower. C Pistillate
1 Deceased. flower. D Dehiscing capsule. (D. Mackay)
20 Anarthriaceae

ovules are pendulous from the upper axile corner

of the three dehiscent ovary loculi (Linder and
Rudall 1993). Kircher {1986) interpreted the in-
florescence as consisting of 1-flowered spikelets,
but we see no evidence for a preceding spikelet
condition. Both the !-flowered units in Anarthria-
ceae and 1-flowered spikelets as found in some
Restionaceae can, however, be considered as re-
duced from plurinodate conditions.

EMBRYOLOGY. The endothecium of the anthers

contains spiral thickenings but often forming U-
shapes rather than complete helices (Manning and
Linder 1990). The development of the male game-
tophyte is not known. The tenuinucellate ovule is
bitegmic, with the inner integument tanniferous.
Megagametophyte development is of the Poly-
gonum type. There are three antipodals, and the
micropylar nucellar epidermis is uniseriate, with
isodiametric cells. There are no large starch grains
in the mature megagametophyte. The develop-
ment of the embryo is not known (Linder and
Rudall1993).
POLLEN MoRPHOLOGY. Pollen grains (Fig. 6) are
shed as moderately sized monads (A. gracilis: ca.
26-38 X 34-45mm, Chanda 1966; Ladd 1977), and
are ulcerate and more or less spherical. The pore is
surrounded by a annulus formed by the thickened
foot-layer: thus the grain is "graminoid" (Ladd
1977). Scrobiculi penetrate the interapertural
pollen wall (Chanda 1966; Ladd 1977; Linder and
Ferguson 1985).
FRUIT AND SEED. The ovary is a loculicidal cap-
sule, and the seeds are dispersed individually.
REPRODUCTIVE BIOLOGY. The plants are wind-
pollinated. Most species resprout after fire but A.
polyphylla Nees is an obligate seeder (Meney and
Pate 1998).
KARYOLOGY. The chromosomes are larger than
those of Restionaceae, with 2n = 22 or 44 (Briggs
1966).
PHYTOCHEMISTRY. Anarthriaceae resemble
many Restionaceae in the presence of flavonols
but lack flavones. Common flavonols including
myricetin were reported in various species;
proanthocyanidins and flavonoid sulphates are
absent (Williams et al. 1997).
AFFINITIES. Anarthriaceae were included within
Restionaceae until 1965. Anarthria resembles the
Fig. 6. Anarthriaceae. Anarthria prolifera, pollen with scro-
biculate ornamentation and annulate ("graminoid") pore,
SEM X 1800. (Photo H.P. Linder)
latter family in the trilocular ovaries with a single
ovule per locule, the tenuinucellate ovules, and the
graminoid, scrobiculate pollen grains (Linder and
Rudall1993; Kellogg and Linder 1995). However,
they differ from Restionaceae in the different culm
anatomy, which lacks parenchyma and scleren-
chyma rings inside the chlorenchyma, the absence
of starch in the megagametophyte, and the isodia-
metric micropylar nucellar epidermis cells; they
also differ from Restionaceae in the bilocular
anthers. Anarthriaceae appear to lack the 28-kb
inversion of the chloroplast DNA found in Restion-
aceae, Poaceae, Joinvilleaceae and Ecdeiocoleaceae
(Doyle et al. 1992). From such lines of evidence,
Kellogg and Linder (1995) suggest a position closer
to Restionaceae than to Poaceae, but note that the
morphological evidence is not compelling. DNA
sequence data for the rbcL gene (S. Gilmore, A.
Marchant and B. Briggs, pers. comm.) indicates a
relatively close relationship with Lyginia and
Hopkinsia but does not suggest a closer relation-
ship to Restionaceae than to Poaceae.
Only one genus:
Anarthria R.Br. Figs. 5, 6
Anarthria R.Br., Prodr.: 248 (1810).
Characters as for family; seven spp., endemic to
the SW of Western Australia, in well-drained or
seasonally wet sites in areas of moderate or rela-
tively low rainfall; on oligotrophic soils.
 Anarthriaceae/ Aponogetonaceae
Selected Biobliography
Briggs, B.G. 1966. Chromosome numbers of some Australian
monocotyledons. Contrib. N.S.W. Nat!. Herb. 4: 24-34.
Chanda, S. 1966. On the pollen morphology of the Centrol-
epidaceae, Restionaceae and Flagellariaceae, with special
reference to taxonomy. Grana 6: 355-415.
Cutler, D.F. 1969. Juncales. In: Metcalfe, C.R. (ed.) Anatomy of
the monocotyledons, Vol. 4. Oxford: Clarendon Press.
Doyle, J.J., Davis, J.I., Soreng, R.J., Garvin, D., Anderson, M.J.
1992. Chloroplast DNA inversions and the origin of the
grass family (Gramineae). Proc. Nat!. Acad. Sci. U.S.A. 89:
7722-7726.
Kellogg, E.A., Linder, H.P. 1995. Phylogeny of Poales. In:
Rudall, P.J., Cribb, P.J., Cutler, D.F., Humphries, C.J. (eds.)
Monocotyledons: systematics and evolution. London: Royal
Botanic Gardens, Kew, pp. 511-542.
Kircher, P. 1986. Untersuchungen zur Bliiten- und Inflo-
reszenzmorphologie, Embryologie and Systematik der
Restionaceen im Vergleich mit Gramineen and verwandten
Familien. Diss. Bot. 94: 1-219.
Ladd, P.G. 1977. Pollen morphology of some members of the
Restionaceae and related families, with notes on the fossil
record. Grana 16: 1-14.
Linder, H.P. 1992. The gynoecia of Australian Restionaceae:
morphology, anatomy and systematic implications. Aust.
Syst. Bot. 5: 227-245.
Linder, H.P., Ferguson, I.K. 1985. On the pollen morphology
and phylogeny of the Restionales and Poales. Grana 24: 65-
76.
Linder, H.P., Rudall, P.J. 1993. The megagametophyte in
Anarthria (Anarthriaceae, Poales) and its implications for
the phylogeny of the Poales. Am. J. Bot. 80: 1455-1464.
Manning, J.C., Linder, H.P. 1990. A cladistic analysis of pat-
terns of endothecial thickenings in the Poales/Restionales.
Am. J. Bot. 77: 196-210.
Meney, K.A., Pate, J.S. (eds.) 1998. Australian rushes-
biology, identification and conservation of Restionaceae
and allied families. Perth: University of Western Australia
Press.
Pate, J.S. 1989. Australian micro stilt plants. Trends Ecol. Evol.
4: 45-49.
Williams, C.A., Harborne, J.B., Greenham, J., Briggs, B.G.,
Johnson, L.A.S. 1997. Flavonoid evidence and the classifica-
tion of the Anarthriaceae within the Poales. Phytochemistry
45: 1189-1196.
21
Aponogetonaceae
H.W.E. VAN BRUGGEN
Aponogetonaceae J. Agardh, Theoria Syst. Pl.: 44 (1858), nom.
cons.
Perennial water plants with a rootstock or rhi-
zome. Leaves in a rosette, submerged and/or float-
ing, very seldom emergent. Inflorescence on a long
scape emerging above the water surface, in bud
enveloped by a caducous or (rarely) persistent
spathe, and composed of 1 or 2-15 spikes. Plants
usually monoecious, seldom dioecious or apoga-
mous. Flowers hypogynous, sessile, more or
less spirally arranged on the inflorescence axis, or
secund. Tepals (1- )2-6, absent in female speci-
mens of dioecious species. Stamens usually 6, ab-
sent in female specimens of dioecious species and
often in apomictic specimens. Anthers 2-thecate,
4-sporangiate, with extrorse longitudinal dehis-
cence. Pollen probably always yellow. Carpels
usually 3, free, sessile, 1-locular, pear-shaped,
narrowed into the stylulus with a stigmatic ridge
on the ventral side, rudimentary and sterile in
male specimens of dioecious specimens, and
sometimes more numerous in apomictic speci-
mens. Ovules 2-12 per ovary, basal or marginal.
Fruit a follicle with a mostly distinct, lateral or
terminal, often curved beak. Seeds without endo-
sperm.
Only one genus of about 45 spp., confined to
the Old World, predominantly the southern
hemisphere.
VEGETATIVE MORPHOLOGY. The starchy tuber is
elongate, obovoid or globular, seldom long, thin,
creeping and branching (A. rigidifolius). The
leaves are usually distinctly petiolate with a nar-
row sheath. Submerged leaves are band-shaped or
narrowly lanceolate, or more seldom oval, ovate,
obovate, threadlike or spindle-shaped. Often the
blades are undulate, sometimes almost contorted
(A. ulvaceus), and in some species strongly
bullate. In A. madagascariensis the leaves are
fenestrate by loss of the mesophyll between the
veins (Fig. 7A,B). Floating leaves are usually oval
or lanceolate and flat; their adaxial surface is
glossy and water-repellent. Usually the blades
have a distinct midrib, and 1 or more pairs of
22 Aponogetonaceae
Fig. 7A-I. Aponogetonaceae. A-E Aponogeton madagas-
cariensis. A Flowering plant. B Portion of fenestrate leaf. C
Flowers. D Flower. E Stamen. F, G, A. undulatus. F Flowering
spike (Note that tepals in fact are spreading as in G). G Flower.
H, I A. distachyos. H Branched spike. I Flower with bract.
(Takhtajan 1982)
parallel, confluent nerves connected by numerous
cross-veins.
VEGETATIVE ANATOMY. Most of the literature on
the anatomy of Aponogeton has been summarised
by Tomlinson (1982), and some salient features
are as follows. Hydropoten are present on the
leaves and petioles as far as these are exposed to
the water. The leaves are glabrous but squamules
are present in leaf axils. The floating leaves are
dorsiventral, the submerged ones isobilateral. The
peripheral periderm of the corm is interrupted by
root and leaf traces. The roots are provided with
branched or unbranched root hairs. Articulated
laticifers are universally present in leaves and the
inflorescence axis but seem to be absent from the
rhizome and the roots. Crystals are abundant in
the mesophyll of floating leaves as coarse aggrega-
tions of rhombohedral types. Tannin is present in
isolated cells common next to vascular tissue or
in partition cells of the lacunose petiole.
INFLORESCENCE STRUCTURE. The inflorescences
are borne on long peduncles, which may taper
towards the tip (Fig. 7). They are then often rigid
and erect, and so lift the inflorescence above the
water surface. In some species they are thickened,
sometimes even strongly inflated below the
inflorescence and in these cases they contain
aerenchymatous tissue which enables them to
float and thus lift the inflorescence.
FLOWER STRUCTURE. The flowers are usually
small and in bisexual specimens consist of 2
tepals, 6 free stamens, and 3 free carpels, but there
are some exceptions to this structure, e.g. in A.
distachyos, the flowers have only 1 large tepal,
while in A. hexatepalus they have 6. The tepals are
petaloid, free scales, which in species with 1 or 2
tepals point obliquely downwards or laterally;
they vary in colour from white to pink or violet,
yellow or green. In about 5 species the tepals are
caducous. The stamens are arranged in 2 whorls,
but in A. distachyos there are 8-16 stamens in
several whorls.
EMBRYOLOGY (mainly from Bouman in van
Bruggen 1985). Anther wall development con-
forms to the monocotyledonous type with an
ephemeral middle layer and an endothecium with
thickenings. The tapetum cells are uninuclear and
amoeboidal. Microsporogenesis is simultaneous
or successive and the pollen is 3-nucleate when
shed.
The ovules are anatropous to hemianatropous
and without appendages. The ovule is mostly
bitegmic, sometimes with 2 partly fused integu-
ments and rarely unitegmic (A. distachyos). The
outer integument is mostly 3-layered, but addi-
tional cell layers may appear during development;
the inner integument is 2-layered. The nucellus is
 Aponogetonaceae
crassinucellate and the nucellar epidermis may
divide periclinally to form a small nucellar cap
and multilayered flanks. The megaspore tetrad is
linear and its chalazal megaspore develops into a
Polygonum type of embryo sac.
Endosperm development is Helobial, the cha-
laza! chamber with one hypertrophied nucleus
and the large micropylar chamber coenocytic. The
first division of the zygote is transverse, the large
basal cell develops into a long-persisting vesicular
cell with a hypertrophied nucleus. The apical cell
divides longitudinally. Embryogenesis according
to the Caryophyllad type or to the Sagittaria vari-
ant of the Caryophyllad type; the embryo lacks a
suspensor.
POLLEN MORPHOLOGY. The pollen grains are sul-
cate and more or less ellipsoid. The tectum is
perreticulate and bears fine supratectal spines
which are always present and visible in the light
microscope (Fig. 8). In some species the lumina
contain free spinules. The aperture membrane,
which is easily damaged during acetolysis, may
bear small spinules or granules, either separately
or in combination. Exine stratification is distinct
only in electron micrographs. Total exine thick-
ness ranges from 0.8-1.5 ftm. The nexine is thin
(ca. 0.2ftm in A. distachyos) and probably com-
pletely ectexinous. The infratectallayer, which is
not visible in the light microscope, is irregularly
granular and up to ca. 0.4ftm thick in A.
distachyos. The intine is thickened at the apertural
region. The long equatorial axis of the grains
ranges from 16-40ftm (data from the comprehen-
sive survey by Thanikaimoni in van Bruggen 1985,
in which all species were covered).
KARYOLOGY. Somatic chromosome numbers are
known from about a 1/3 of the total number of spe-
cies, in most cases counted only in 1 or a few
specimens. Some numbers are not completely reli-
A B
Fig. 8A,B. Aponogetonaceae. Aponogeton elongatus, sulcate
pollen, SEM X900. A Distal pole. B Proximal pole. (Orig.)
23
able. Even the best metaphase plates are very diffi-
cult to analyse because the chromosomes (0.5-
2.5 ftm long) have a strong tendency to adhere to
one another. However, the numbers 2n = 16, 24,
32, 40 and 56 suggest a basic number of x = 8.
Aneuploidy and infraspecific polyploidy seem to
be common phenomenona, in some cases com-
bined with each other.
It remains to be established whether there is a
connection between high chromosome numbers
and apomixis or dioecism.
PoLLINATION. The flowers are probably polli-
nated by insects, but this has not yet been
established.
FRUIT AND SEED. The ripening infructescence
gradually becomes submerged. On the basis of
seed-coat structure, 3 seed types have been
recognised by Bouman (in van Bruggen 1985): (1)
Double-coated seeds developing from bitegmic
ovules. Testa separable from the tegmen. Testa 3-
4-layered, with ridges or small wings. Exotesta
with bandlike, annular thickenings on their
anticlinal walls. Middle layer of the testa
parenchymatic with small intercellular spaces.
Inner layer not differentiated. Inner layer of the
tegmen tanniniferous and walls impregnated, re-
sponsible for the non-transparent character of the
seed coat. Outer layer of the tegmen collapsing
during development. Embryo achlorophyllous
(Fig. 9). (2) Single-coated seeds developing from
ovules with 1 or with 2 fused integuments. Seed
coat 5-10 cells thick around the embryo sac, with-
out thickenings or a tanniniferous layer and thus
transparent. Middle layers parenchymatic and
rich in intercellular spaces. Embryo chlorophyl-
lous. (3) Single-coated seeds developing from
bitegmic ovules. Testa 3- 5- and tegmen 2-layered.
Seed coat without distinct thickened or tan-
niniferous layers. This group is more or less
intermediate between the first and second.
The seeds have no endosperm. Seeds with a
single coat are glossy and the seed coat is thick and
fleshy; the embryo may have a plumule or not. In
seeds with a double coat the testa is loose, trans-
parent and reticulately veined; the tegmen is
brown and closely encloses the embryo, which is
completely undeveloped and seems not to have an
externally visible plumule.
DISPERSAL. At maturity the pericarp suddenly
decays and the seeds are released. Seeds with a
single coat float on the water surface for a short
period (a few hours), and may reach other locali-
24 Aponogetonaceae
Fig. 9. Aponogetonaceae. Seed of Aponogeton ulvaceus, with
transparent coat and central part with tegmen and embryo.
X40. (Orig.)
ties. Then the testa will also decay, and the embryo
sinks to the bottom. These seeds have no resting
period.
Seeds with a double coat have no floating tissue,
but due to surface tension they may float for some
time and thus can be dispersed. They will germi-
nate after a resting period of 1 to several weeks.
In this way, the plants can easily reach new sites in
the same stream or pool (they always grow in
widely scattered batches), but as the seeds cannot
tolerate desiccation, it is not clear how they reach
new streams or pools.
REPRODUCTIVE BIOLOGY. Most species are her-
maphroditic, but there are five dioecious species
and others, which to a high degree are apomictic.
In some of the dioecious species, male specimens
predominate. The ratio between male and female
specimens seems to be 10: 1, but it is not known
whether this applies to all dioecious species.
Apomixis occurs in many species. Sometimes all
specimens may be apomictic and lack stamens,
but other species are both apomictic and sexual.
In still other species the flowers are perfect, but the
fruit produces many fertile seeds without pollina-
tion (e.g. A. natans). Of the dioecious A. decaryi,
one apomictic population is known without any
male specimens.
Fruit development is usually very rapid, taking
from 1 to about 4 weeks. Species living in desert
areas set fruit in a very short time. Because the
seeds cannot survive dessication, they must ger-
minate, and the seedling must develop a small
tuber before the renewal of drought conditions
following rain.
PHYTOCHEMISTRY. Aponogeton differs from most
Potamogetonales in the presence of leuco-
anthocyanins and flavonols and the absence of
C-glycosyl flavones, probably the plesiomorphic
state in Alismatales (see compilation in Les and
Haynes 1995). Cell wall-bound ferulate was found
to be absent (Harris and Hartley 1980).
AFFINITIES. On the basis of the apocarpous
gynoecium and anatomical evidence (Tomlinson
1982), a position of Aponogeton in the Helobiae is
supported. Like all other Alismatales, the ripe seed
has no endosperm and the embryo is supplied
with storage tissue. The presence of laticifers has
been seen as pointing to affinities with Alismata-
ceae and Butomaceae. The molecular (rbcL) data,
however, that have been reported by Les and
Haynes (1995), indicate for Aponogetonaceae a
position basal to Potamogetonales.
DISTRIBUTION. The 45 species of the single genus
are confined to the Old World, except Europe and
temperate Asia. The centre of diversity is in the
tropical and subtropical areas of Africa and Mada-
gascar. Aponogeton reaches the Cape region of
Africa and occurs also in India and Australia. All
species have fairly restricted ranges. On the whole,
the species do not seem to be endangered, but A.
bullosus from the Atherton Tableland (Australia)
has become very rare through overcollection for
aquarium purposes.
EcoLOGY. All members of the family are true
aquatics. Some species occur in stagnant water,
others in more swiftly flowing streams and rivers,
some even in torrents (rheophytes). One or two
species are found on banks and shores. Some
species grow in permanent, others in temporary,
water.
PALAEOBOTANY. In recent years three species
from the Upper Oligocene have been described,
all from the region of the Aral Lake, almost
3000 km from the nearest recent locality in India.
For one of them, A. tertiarius, definite proof that it
is an Aponogeton (and not a Potamogeton) is still
lacking, because reproductive structures are un-
known. From the other two species (A. zhilinii and
A. czokusianus) only fragments were collected,
 Aponogetonaceae 25

and it is difficult to judge whether they really rep-
resent different species.
A fossil Aponogeton from the Cretaceous of
Patagonia was described by Selling (1947). The
former occurrence of the genus in S. America
seems, however, highly improbable in view of its
recent absence from this continent.
EcoNOMIC IMPORTANCE. About a dozen species
are of some economic importance. In some re-
gions of the world the tubers and in others the
leaves are eaten. In Africa the inflorescences of A.
distachyos are used as an ingredient in soups and
stews; the plant is cultivated for this purpose. Dur-
ing the flowering season the inflorescences are
sold, and they are then even canned.
About five spp. from Sri Lanka and Madagascar
are imported on a large scale to Europe and the
United States for aquarium purposes. A few other
species are imported only occasionally.
A. distachyos is a popular plant for ponds. When
there is no ice on the water it may even flower in
midwinter.
Pneva, G.P. 1988. Eine neue tertiiire Art aus der Gattung
Aponogeton (Aponogetonaceae) aus Kazakhstan und
Karakalpakiya, Bot. Zh. 73: 1597-1597 (in Russian).
Selling, O.H. 1947. Aponogetonaceae in the Cretaceous of
South America. Sven. Bot. Tidskr. 41: 182.
Takhtajan, A.L. 1982. See general references.
Tomlinson, P.B. 1982. Helobiae (Alismatidae). In: Metcalfe,
C.R. (ed.) Anatomy ofthe monocotyledons, Vol. 7. Oxford:
Clarendon Press, pp. 198-225.
Yadav, S.R. 1993. Mechanism of apomixis in Aponogeton
decaryi Jumelle. Phytomorphology 43: 201-207.
Yadav, S.R. 1995. Die Hybride zwischen Aponogeton decaryi
Jumelle und Aponogeton satarensis Raghavan, Kulkarni &
Yadav. Aqua-Planta 20: 71-80.
Zhilin, S.G. 1974. Die erste tertiiire Art aus der Gattung
Aponogeton (Aponogetonaceae). Bot. Zh. 59: 1203-1206 (in
Russian).

Only one genus:

Aponogeton L. fil. Figs. 7-9

Aponogeton L. fil., Suppl. Pl.: 214 (1781); van Bruggen, Bib!.
Bot. 137 (1985) (monogr.).

Characters as for family.

Selected Bibliography

A more comprehensive bibliography can be found in van

Bruggen (1985).
Andrejev, A. 1989. Aponogeton czokusianus A. Andrejev sp.
nov. In: Zhilin, S.G. (ed.) History of the development of the
temperate forest flora in Kazakhstan, USSR from the Oli-
gocene to Early Miocene. Bot. Rev. 55: 292.
Bruggen, H.W.E. van 1985. Monograph of the genus Apono-
geton (Aponogetonaceae). Bib!. Bot. 137. 76 pp., 23 pl.
Bruggen, H.W.E. van 1987. Aponogetonaceae. In:
Dassanayake, M.D. (ed.), Rev. Handb. Flora Ceylon Vol. 6:
3-16. New Delhi: Amerind.
Bruggen, H.W.E. van 1990. Die Gattung Aponogeton
(Aponogetonaceae). Aqua-Planta Sonderheft 2.
Bruggen, H.W.E. van 1991. Neue Erkentnisse tiber die
Aponogetonaceae. Aqua-Planta 16: 46-54.
Harris, P.J., Hartley, R.D. 1980. See general references.
Les, D.H., Haynes, R.R. 1995. Systematics of subclass
Alismatidae: a synthesis of approaches. In: Rudall, P.J.,
Cribb, P.J., Cutler, D.F., Humphries, C.J. (eds.) Monocotyle-
dons: systematics and evolution, Vol. 2. London: Royal
Botanic Gardens, Kew, pp. 353-377.
Lye, K.A. 1986. The geographical variation in Aponogeton
abyssinicus A. Rich. Lydia 1: 67-80.
26 Araceae
Araceae
S.J. MAYO, J. BoGNER, and P.C. BoYcE
Araceae Juss., Gen. Pl.: 23 (1789), nom. cons.
Herbs, diminutive to gigantic, terrestrial, geo-
phytic, epiphytic, or climbing, rarely aquatic;
with true or false stems; tissues with raphides;
laticifers commonly present; mature shoots com-
posed of articles forming a sympodium (except
Pothoideae, Heteropsis ); leaf bifacial, usually dif-
ferentiated into petiole and expanded blade of
very variable shape; midrib compound; primary
venation usually pinnate but sometimes pedate
or arcuate from the base or strictly parallel, fine
venation ± reticulated or parallel-pinnate (i.e.
parallel to primary lateral veins); inflorescences
1-many, each inflorescence a pseudanthium
(especially so when flowers unisexual) consisting
of a spadix (a dense spike of minute, ebracteate
flowers) subtended by a spathe (last leaf of article,
usually specialised in form and colour, incon-
spicuous in Gymnostachys, Orontium); flowers
sessile (except Pedicellarum), bisexual or un-
isexual (unisexual inflorescences in Arisaema -
paradioecy), when unisexual the female flowers
occupying the proximal part and males the me-
dian or distal part of the spadix; perigone present
or absent, tepals free, partly or completely con-
nate; stamens usually 3 + 3 or 2 + 2, or sometimes
less or more, opposite tepals in bisexual flowers,
free to connate, anthers extrorse (except
Zamioculcas, Pedicellarum); gynoecium syncar-
pous, ovary usually 1-3 locular ( 1-8 locular in
Spathicarpeae, 2-47-locular in Philodendron),
ovules bitegmic, atropous, anatropous, ana-
campylotropous or of intermediate form; style
usually inconspicuous, stigma wet at anthesis,
sometimes distinctly lobed; fruit a berry (basally
dehiscent in Lagenandra), usually free and borne
in ± cylindric, dense, fleshy spike, rarely connate
into syncarp; seeds small to very large, testa very
thin to very thick, rarely absent, endosperm
present or absent, embryo minute to large, both
with or without starch, chlorophyllous or not.
A family comprising 104 genera and over 3300
species, with a subcosmopolitan distribution but
most diverse in tropical regions.
VEGETATIVE MoRPHOLOGY. The stem varies
from an elongated aerial axis with extended
internodes, as in the many climbers and climb-
ing hemiepiphytes, to a hypogeal rhizome or
depressed-globose (rarely elongate to napiform)
tuber. Genera with rhizomes or tubers are com-
mon in Lasioideae and Aroideae. The climbing
habit is characteristic of subfamilies Pothoideae
and Monsteroideae. Abbreviated aerial stems
are also frequent, producing rosulate plants, as in
certain epiphytic species of Philodendron and
Anthurium. Erect stems with relatively short
internodes occur in some tribes of Aroideae
(e.g. Aglaonemateae, Dieffenbachieae ). Some,
generally larger, species have an arborescent habit
in which the main axis is a fleshy (Alocasia,
Montrichardia, Xanthosoma sect. Xanthosoma)
or fibrous (Philodendron) stem or a false stem
of petiole sheaths (Arisaema, Dracunculus,
Typhonodorum). Shoots specialised for vege-
tative reproduction include flagelliform shoots
(e.g. Amydrium, Cercestis, Philodendron, Rhodo-
spatha, Syngonium), bulbils adapted for animal
dispersal (Remusatia), cormlets (Dracontium,
Dracontioides) and hypogeal stolons (Ari-
saema, Colocasia, Cryptocoryne, Lasimorpha,
Spathiphyllum ).
In virtually all genera the mature stem is a
sympodium composed of articles ending with
an inflorescence or aborted inflorescence. Foliage
leaves and cataphylls occur in an often predictable
sequence within each article; the constancy of the
sequence varies from genus to genus. With the
termination of the article by an inflorescence,
growth of the stem is continued in most genera by
a branch axillary to the foliage leaf or cataphyll
situated at the second node below spathe
insertion. In the Orontioideae this continuation
shoot arises at the first node below the spathe.
"Enrichment" of the flowering shoot to form
a synflorescence takes place commonly by the
development of a sequence of short articles,
each consisting at least of prophyll, spathe and
inflorescence, beginning in the axil of the leaf
immediately below the spathe. The resulting
synflorescences exhibit a range of structural varia-
tion which may be quite complex; extreme forms
are found in Gymnostachys and Homalo-
mena. Anomalous shoot arcliitecture occurs in
Gymnostachys, Potheae and Heteropsis, in which
flowering is confined to lateral short shoots from
the monopodia! main vegetative axis.
Roots in Araceae are always shoot born; the
primary root withers soon after germination or is
 Araceae 27

absent (Pistia). In root climbers and hemi- be arcuate-parallel (i.e. acrodromous, campylo-
epiphytes, root dimorphism is frequent in; non- dromous, convergate or curvipalmate, e.g.
geotropic anchor roots, adapted for clinging on Anthurium sect. Digitinervium), pedate (e.g.
trees, are anatomically or morphologically distin- Sauromatum) or radiate (e.g. many Arisaema
guishable from larger feeding roots which grow spp.) but are most commonly pinnately organised.
down to the forest floor, as in Monstera, Philoden- Even in pedately and radiately lobed leaves, the
dron, Rhaphidophora. In some geophytic genera, primary veins of the lobes are generally pinnate.
e.g. Arisarum, Arum, Biarum, and in Crypto- In pinnately veined leaves the major branches of
coryne, which is rhizomatous, contractile roots the midrib are thus usually primary veins, but are
occur. referred to as primary lateral veins. Except possi-
In virtually all genera the leaf is clearly differen- bly in some compound leaves, the primary veins
tiated into a blade, petiole and petiole sheath; ex- always run throughout the leaf blade, ultimately
ceptions are Gymnostachys, some Biarum spp. joining together at the leaf apex. In many genera
and Pistia. The sheath normally clasps the sub- either one or several of the primary veins form
tended internode, at least basally, and has an an- a submarginal collective vein on each side in ad-
nular insertion (except many Potheae and most dition to the marginal vein. The finer venation
Heteropsis ). The foliage leaves of mature shoot ar- may be reticulated (e.g. Anthurium) or parallel-
ticles often have short or very reduced sheaths. pinnate (i.e. more or less parallel to the major
Leaf blade size may range from diminutive lateral veins); the latter is normally referred to as
(e.g. Ambrosina) to gigantic (e.g. Amorphophallus, "parallel" or "striate" venation in the taxonomic
Colocasia, Cyrtosperma, Xanthosoma). Shape literature on aroids. A third, intermediate type
varies from linear (Biarum, Typhonium) to of fine venation is recognised for the tribes
dracontioid (Anchomanes, Dracontium, Pseudo- Colocasieae and Caladieae ("colocasioid vena-
hydrosme, Pycnospatha, Taccarum, Thom- tion"). Here the fine veins arising from major lat-
sonieae), through cordate, sagittate, hastate, eral veins arch towards the leaf margin, and may
trilobed or trisect, pedately, pinnately and radi- run virtually without reticulation (Colocasia) or
ately lobed or -sect. Bipinnatifid, tripinnatifid fuse to form sinuose sympodial collective veins
and combined pedatisect-pinnatifid leaves also ("interprimary veins") between the major lateral
occur. Dracontioid leaves are generally trisect veins (most Alocasia spp., Xanthosoma). Strictly
at the petiole insertion and thereafter more or parallel venation of the typical monocot type is
less dichotomously subdivided. Zamioculcas, found only in Gymnostachys. Intermediate forms
one species of Anaphyllum and one species of occur between virtually all recognised patterns
Anaphyllopsis have truly pinnate compound and this makes venation a difficult character to
leaves. use. Ertl's study (1932) is the only large-scale
Heteroblasty is a striking feature and defines a survey yet made.
number of climbing genera [Cercestis, Monstera,
Philodendron, Pathos (except subgen. Pathos), VEGETATIVE ANATOMY. 1 The rhizodermis is 1-,
Rhaphidophora, Rhodospatha, Syngonium]. It oc- less commonly several-layered. True multiple root
curs both in ontogeny from seedling to mature epidermis resembling a velamen occurs in various
plant and in association with the development Anthurium spp. The cells are not or at most
of flagelliform shoots. Fenestrate or perforated vaguely differentiated into long and short cells.
leaves are another peculiarity well shown in gen- A sclerotic root hypodermis occurs in Culcasieae,
era such as Amydrium, Cercestis, Dracontioides, Montrichardia, Philodendron, Homalomeneae
Monstera and Rhaphidophora. In genera with di- and Anubias. Lobing of the central vascular cylin-
vided leaves the first foliage leaf is normally entire, der occurs in the roots of some Philodendron and
although in Gonatopus and Thomsonieae it is Cercestis species.
already divided. A number of genera have spe- In the stem, a cortical vascular system, distinct
cies with peltate leaves (Ariopsis, Caladium, from the central cylinder, is present in some gen-
Colocasieae and a few species of Anthurium and era (Pathos, Pothoidium, Heteropsis, Anthurium,
Homalomena).
Leaf venation in Araceae is complex and varied
in detail but little studied and the terminology
used is confusing in practice. The primary veins 'This account is based on the treatment by J.C. French in
(i.e. the major veins including the midrib) may Mayo eta!. (1997).
28 Araceae
Monstereae, Philodendron, Dieffenbachia, Syn-
gonium, Caladium, Chlorospatha, Xanthosoma),
which in Colocasieae often forms a character-
istic anastomosing system. An unusual type of
endodermis, which surrounds each axial vas-
cular bundle individually but not the central
cylinder or leaf traces, is widespread in the
Schismatoglottideae.
Bud trace insertion patterns are also variable.
Three patterns are recognisable in Araceae. Two
of them are found mainly in climbing genera and
a third is characteristic of genera with rhizomes or
tubers (French and Tomlinson 1981a-d, 1983,
1984).
In the leaf, trichomes and larger epidermal
scalelike processes are rare (occurring in Bognera,
Callopsis, Pedicellarum, Pistia and certain species
of Anubias, Cryptocoryne, Homalomena, Philo-
dendron, Schismatoglottis, Stylochaeton, Syn-
gonium, Xanthosoma). Prickles are found on
stems, petioles and peduncles in various species
of Lasioideae, Nephthytideae, Homalomena,
Anubias, and Montrichardia. Warty petioles and
peduncles occur in some genera of Lasioideae and
in Amorphophallus and Arisaema. Stomata tend
to be concentrated on the abaxial leaf surface. The
number of subsidiary cells varies from none
(anomocytic, e.g. in Lysichiton, Orontium, Pistia,
Symplocarpus) to 2 (paracytic, e.g. in Amor-
phophallus, Arisaema, Arum, Rhaphidophora,
Typhonium) to 4-8 (tetracytic, e.g. in Dieffen-
bachia, Lagenandra, Philodendron and Rhaphido-
phora); paracytic is the commonest type. Leaf
mesophyll is predominantly bifacial and in
many species of Homalomena, Philodendron,
Piptospatha, Schismatoglottis and Typhonium is
distinctively chambered. Petiole anatomy has so
far been little studied. The occurrence of a
geniculum (pulvinus), usually at the petiole apex,
is a useful taxonomic character and commonest in
genera with bisexual flowers. In Gonatopus,
genicula also occur at the base of the major leaf
pinnae, and in G. boivinii and in at least one
species of Anthurium (A. oerstedianum) the
geniculum is located in the middle of the petiole
instead of the apex or nearly so.
Variegation of the leaf blade, petiole and pe-
duncle is a striking feature of many genera, in-
cluding several which are well known in
horticulture (Aglaonema, Dieffenbachia, Cala-
dium). Variegated leaves also occur in the juvenile
plants of some species of Monstera, Philodendron,
Scindapsus and Syngonium and in general seem to
be more common in plants growing in tropical
rainforests on or near the ground. In Caladium
and Zantedeschia, necrotic areas provide another
mode of attactive leaf variegation.
Trichosclereids are characteristic of the Spathi-
phylleae and Monstereae (sparsely present in
Amydrium).
Laticifers, normally associated with the phloem
of the vascular bundles, occur in the majority of
Araceae, although absent from bisexual-flowered
genera except Calla and Orontium. Anastomos-
ing laticifers occur in tribes Colocasieae (in
Ariopsis the laticifers are simple), Caladieae, and
Zomicarpeae. Latex composition varies, being
clear or milky with a variety of constituent latex
particles and chemical constituents. Large quanti-
ties of tannin have also been detected in aroid
latex.
Resin canals containing terpenoids are known
from the leaves, stems, roots and inflorescen-
ces of Cercestis, Culcasia, Homalomena and
Philodendron.
Extrafloral nectaries occur on the leaves, peti-
oles and spathes of Philodendron and Culcasia,
wax glands in the Alocasia species previously
recognised as Xenophya and intravaginal
squamules, of unknown function, occur in the leaf
axils or at the nodes in Cryptocoryne, Lagenandra
and Philodendron.
Calcium oxalate has been recorded from almost
all genera and is found in various crystalline
forms, including raphides, druses and specialised
idioblasts (biforines). The latter have a "blowgun"
explosive discharge and contain large, often
grooved, raphides neatly arranged in bundles. For
illustration of the various crystal forms see
Seubert (1993).
Vessels have been found in the roots of many
genera and in the stems of a few species of
Epipremnum, Philodendron, Pathos, Rhaphido-
phora and Scindapsus.
INFLORESCENCE AND FLOWER STRUCTURE. The
inflorescence of almost all Araceae is an un-
branched, fleshy spike of numerous, very small
flowers, and is known as a spadix. The family is
characterized by the absence of floral bracts
and the lack of pedicels in all genera except
Pedicellarum. Occasionally the spadix is not
fleshy or only few-flowered. The flowers are
usually spirally arranged and tightly packed, al-
though in some species of Amorphophallus (male
and female flowers), most species of Arisaema
and Arisarum (male flowers), Dieffenbachieae
(female flowers), Pedicellarum and Pathos series
Goniuri, they may be somewhat distant from one
another.
 Araceae
The last leaf of each flowering article is called the
spathe and is part of the inflorescence. The spathe
is usually distinctive, although in Gymnostachys
and Orontium it remains an inconspicuous bract.
The internode between spathe and spadix (termed
the spadix stipe) is usually rudimentary, whereas
that between spathe and uppermost foliage leaf
(termed peduncle) is much longer. In some con-
servative genera, however, this arrangement is re-
versed (Gymnostachys, Orontioideae, some Pathos
spp.). Gymnostachys, Orontioideae and Poth-
oideae seem to represent earlier, less uniform
phases of organisation. The typical araceous
inflorescence has given rise to a wide range
of variant forms in different genera, which are
seen as representing an overall evolutionary
trend of increasing integration towards a pseu-
danthium. The major phyletic modifications are
(1) loss of perigone in the flowers; (2) differentia-
tion of flowers on the spadix into a lower female
zone, upper male zone and, often, one or several
sterile zones of sterile flowers or axial regions
lacking flowers - terminal appendices are a well-
known feature of more advanced genera (e.g.
Areae); (3) differentiation of the spathe into a
lower, convolute tube and an upper, expanded
blade.
Spathe and spadix modifications are closely re-
lated, so that in a phyletic sense the spathe has
become increasingly integrated into the pseu-
danthium. In extreme cases (e.g. Cryptocoryneae,
Ambrosina, some Typhonium spp., Pinellia,
Pistia, Spathantheum and Spathicarpa), fusion
and still more elaborate modifications have
brought about division of the spathe into separate
chambers. In some genera (e.g. Philodendron) the
entire spathe persists until fruit, but in most
Caladieae, Colocasieae and Schismatoglottideae
the blade withers or drops off immediately after
anthesis and only the tube persists until fruiting.
In Monstereae the entire spathe withers or drops
off soon after flowering, a behaviour which seems
to be correlated with the presence in this tribe of
numerous protective trichosclereids in the style
tissue. Terminal appendices of the spadix are
found in some genera of the Colocasieae and
Schismatoglottideae, all genera of the Zomi-
carpeae, Thomsonieae, Areae and Arisaemateae
and sporadically elsewhere in the family. The
function of the appendix, where studied, is to pro-
duce odours which attract pollinators. Structur-
ally the appendix varies from being covered by
staminodes (e.g. Pseudodracontium) to entirely
smooth with no vestige of floral organs (e.g. most
Areae).
29
Flowers in Araceae may be 2- or 3-merous. In
perigoniate flowers the tepals, when free, are
organised in 2 whorls. The tepals are usually ±
fleshy and fornicate apically (except Potheae)
and in some genera or sections (Anadendrum,
Holochlamys, Pedicellarum, Spathiphyllum sect.
Massowia, Stylochaeton) they are fused so that the
perigone is a single, cuplike structure. Stamens
in perigoniate flowers and in the naked bisexual
flowers of most Monsteroideae have essentially
the typical structure of a distinct (usually
flattened) filament, terminal basifixed anther
and slender, inconspicuous connective. In the
unisexual flowers of Philodendron, Anubias and
Homalomeneae, however, the stamens typically
lack filaments and have a thick, fleshy connective
which is probably osmophoric. Stamens of the
Colocasieae and Caladieae are similar but always
fused into synandria. In Arophyteae and other
Aroideae the stamens may be fused or not but
generally lack conspicuous filaments. Large, fused
connectives also occur in the Spathicarpeae but
differ from those of the Caladieae and Coloca-
sieae. Well-developed connectives also occur in
Zomicarpeae, especially in Filarum, where they
are elongated and thread-like. Anthers are
almost always extrorse (introrse in Zamioculcas,
latrorse in Pedicellarum). The thecae dehisce by
longitudinal slits (most genera with bisexual
flowers and some unisexual flowered genera:
Anubias, some Areae, Arisaema, Arisarum,
Stylochaeton) or by apical or subapical pores or
short slits. In Philodendron, Anubias, Homalo-
meneae, Caladieae and Colocasieae, the dehis-
cence of each theca is usually by a subapical pore,
which seems to be correlated with the extrusion of
the pollen in strands; similar structures occur in
Amorphophallus and Dracunculus.
The number of locules in the ovary normally
varies between 1 and 3, and, when unilocular, the
gynoecium often shows traces of 2- or 3-merous
origin through the presence of a several-lobed
stigma (e.g. Typhonodorum) or more than 1 pla-
centa (e.g. most Schismatoglottis spp.). Gynoecia
with more than 3 locules are less common but are
found in the Spathicarpeae (1-8-locular) and are
normal in Philodendron (2-47-locular). Placenta-
tion varies from axile to parietal or basal or apical
or both basal and apical, with many intermediates.
Ovules may be anatropous, amphitropous,
anacampylotropous, atropous or intermediate be-
tween these types. Ovular trichomes are usually
present and secrete a clear, mucilaginous sub-
stance which in many genera (e.g. Monstereae,
Philodendron) entirely fills the ovary locules; this
30 Araceae

secretion may play a role in pollen-tube growth. form a parietal cell or not), most Araceae so far
The style may be narrow and elongated (e.g. investigated, except Symplocarpus and Calla, have
Dracontium) but in most genera it is superficially tenuinucellate ovules.
inconspicuous, even when a thick stylar region Linear megaspore tetrads are the commonest
is developed above the ovary locules. Stigmas type, but T-shaped tetrads are also found and
are wet in Araceae and in some genera (Anthu- sometimes both types occur in the same species.
rium, Arum, several Lasieae) produce con- The mature embryo sac is usually of the 8-nucleate
spicuous nectar droplets at anthesis. In Alo- type (10-12-nucleate in Nephthytis). Embryogeny
casia, Amorphophallus, Dieffenbachia and is clearly understood in only nine genera. Onagrad
some Spathicarpeae the lobing of the stigma is and asterad embryogeny are known in unisexual-
pronounced or the stigma is relatively massive. flowered genera and caryophyllad and solanad
types are known in bisexual-flowered genera.
INFLORESCENCE AND FLOWER ANATOMY. Eyde et Cellular and free-nuclear types of endosperm
al. (1967) established that in floral anatomy, as in development both occur but, according to
other characters, Acorus differed markedly from Grayum (1991), endosperm development in
the Araceae and confirmed the absence of floral Araceae is best interpreted as a form of the
bracts in Araceae. Studies of inflorescence and flo- Helobial type. Chalaza! and micropylar cells
ral anatomy in Philodendron have demonstrated ("chambers") are formed by the first division of
the existence of spathe tissues adapted for opening the endosperm nucleus, followed by cellular
and closing movements, resin secretion of various or free nuclear divisions in the micropylar
types from the spathe and a wide variety of chamber. The chalazal cell remains undivided,
gynoecial and androecial structure (e.g. Mayo usually with a single, hypertrophied nucleus,
1986). Barabe and coworkers (e.g. Barabe and forming a "haustorium" or "basal apparatus". An
Forget 1988) have studied the floral anatomy of endothelium, derived from the inner surface
representatives of a range of tribes with par- of the inner integument, is reported for most
ticular emphasis on floral vasculature. Family- genera investigated.
wide surveys by French of endothecial thickenings
(French 1985a,b, 1986a), stamen vasculature PALYNOLOGY. Major contributions: Thanikai-
(French 1986b), ovule vasculature (French 1986c) moni (1969), Grayum (1984, 1985, 1986a, b, 1992).
and ovular trichomes (French 1987) have further The pollen grains may be sulcate, extensive-
confirmed the distinctness of Acorus and revealed sulcate, meridionosulcate (zonate), disulcate
taxonomically useful character variation. (diaperturate), forate (periporate) or inaperturate
(omniaperturate) (Fig. 10).
EMBRYOLOGY. The embryology of Araceae has Sulcate pollen grains are commonest in the
been reviewed by Jiissen (1928) and Grayum more conservative genera, i.e. those with bisexual,
(1984, 1991). Thorough treatments of all embryo- perigoniate flowers. Spathiphyllum, in spite of
logical aspects have been published for Peltan- several primitive characters, has inaperturate
dra virginica, Synandrospadix vermitoxicus and pollen grains as do all monoecious taxa except
Theriophonum minutum; other genera are known Zamioculcadeae.
less completely. Generalisations about the family Exine ornamentation may be smooth (psilate),
are therefore based on rather fragmentary scabrate, foveolate, reticulate, spinulose, spinose,
coverage. fossulate, rarely gemmate, verrucate, areolate,
Araceae have a periplasmodial anther tapetum, rugulate or striate. Foveolate or reticulate exines,
thus differing from Acorus, which has the secre- widespread among monocots, are considered to
tory type. Pollen mother cell division is probably be primitive in Araceae.
always successive. Both 2-nucleate and 3-nucleate Pollen-grain shape is globose to ellipsoid (boat-
pollen is widespread in the family, but pollen shed shaped) or hamburger-shaped (Gonatopus, some
at the 2-nuclear stage is the condition normally Heteropsis spp., most Monstereae, Zamioculcas).
found in the less advanced genera (Grayum The pollen grains are usually shed as monads, but
1986a). in two relatively advanced genera (Chlorospatha,
The nucellar epidermis of virtually all Araceae Xanthosoma) they are release as tetrades; the tet-
forms a "nucellar cap", usually 2-3 cell layers rads are arranged tetrahedrally or in a serial se-
thick (1 in Pistia), and for this reason the Araceae quence (Chlorospatha longipoda).
have been termed crassinucellate. However, in a The size of the pollen grains varies from small
strict sense (division of the archesporia! cell to (12 f!m in Homalomena versteegii) to large (ca.
 Araceae 31

Fig. lOA-E. Araceae. Pollen SEM micrographs. A Gymno-

stachys anceps, sulcate, exine foveolate. X3800. B Gonatopus
angustus, zonasulcate, exine foveolate. X 1200. C
Sauromatum venosum, inaperturate, exine spinose. XS60. D
Spathiphyllum cochlearispathum, inaperturate, exine striate.
X2200. E Pistia stratiotes, inaperturate, exine plicate with
undulate frills. X900. (Photo W. Barthlott)

60 ~-tm) or very large (106 ~-tm in Pseudohydrosme The pollen of most genera contains starch (73%
gabunensis), but in the majority of genera (68%) of the examined species), the remainder being
the grains are medium-sized (between 25 and starchless; in Schismatoglottis both types have
50 ~-tm diam., mean 37~-tm). been found in different species.
32 Araceae

It seems probable that exine ornamentation is The size and shape of the chromosomes are also
correlated with pollinator type. Smooth pollen quite variable. The exceptionally large chromo-
(and probably pollen extruded in strands) is usu- somes in Anchomanes and Nephthytis (Marchant
ally associated with beetle pollination according to 1971a) is an important reason for placing these
existing observations, while spinose pollen is usu- genera in the same tribe.
ally found in fly-pollinated genera. Exceptions
have, however, been observed. In Cryptocoryne, PHYTOCHEMISTRY. The Araceae are well known
smooth pollen grains are associated with pollina- as a family which includes poisonous plants. Many
tion by minute flies; in bee-pollinated Peltandra ornamentals (e.g. species of Alocasia, Caladium,
sagittifolia pollen is smooth; Amorphophallus Dieffenbachia, Philodendron, Zantedeschia) are
maculatus, with somewhat scabrous pollen, known to cause poisoning, particularly wheu veg-
appears to be pollinated by both beetles and etative plant parts are accidentally eaten by chil-
blowflies. dren and pets. The red berries of Arum are known
to be poisonous and the tubers of all e :lible
KARYOLOGY. Petersen (1989) has provided a criti- Araceae must be cooked or treated before the'r can
cal and comprehensive review. Chromosome be eaten. The presence of active compounds i~ a: so
numbers in Araceae vary from 2n = 14 (Ulearum) evident from the use of Araceae for medicinal
up to 2n = 168 (Arisaema). The number may vary purposes by people in many parts of the world;
considerably within one genus, e.g. in Crypto- e.g. see Plowman (1969); Knecht (1983); !:own
coryne (2n = 20 to 132) or in Arisaema (2n = 20 (1988). The chemical basis of the toxicity of
to 168). Other genera have very stable numbers, Araceae is not completely understood, but it is
as in Anthurium, the largest genus of the family generally assumed that calcium oxalate raphicles
(2n = 30 in the great majority of spp.), Dracon- can cause minute lesions in the mouth, thus fac:Ji-
tium (2n = 26), and Zantedeschia (2n = 32). tating the access of toxic chemical compounds to
The genera with bisexual, perigoniate flowers the blood system.
tend to have less variation in chromosome num- Ethereal oils are present in Homalor.1ena
ber and neither very high nor very low numbers; but do not occur in specialised idioblast1:. In
for example, Pothos has 2n = 24 and 2n = 36 and Araceae, major groups of compounds include
Spathiphyllum 2n = 30 and 2n = 60. The highest saponins, proanthocyanidins, cinnamic acids,
numbers appear in tribes with advanced floral and flavonoids, cyanogenic glucosides, alkaloids a1d
inflorescence morphology, e.g. the Areae, the amines; cyanogenesis is widespread. Irritant
Arisaemateae and the Cryptocoryneae. In certain compounds so far identified include protocate-
groups with advanced floral characters (e.g. chuic aldehyde, homogentisic acid and :h·~ir
Zomicarpeae), there has been a reduction in chro- glycosides (Alocasia, Colocasia, Pinellia) nd
mosome number. allergenic 5-alkyl- and 5-alkenylresorci nols
Although a primary base number of x = 7 has (Philodendron). The flavonoids and antho-
been proposed by other authors, Petersen (1989) cyanins have been surveyed in Araceae (Will ams
considered that the base numbers x = 14 or x = 12 et al. 1981). C-glycoflavones, flavonols and pro-
may have been the starting point for the deriva- cyanidins are the main flavonoids, while flav,)nes
tion of all the chromosome numbers in extant are of limited ocurrence. Broadly speaking, tlle
genera of Araceae. These numbers may represent chemistry of the Araceae is comparable to th 1t of
secondary base numbers. the Liliiflorae (Hegnauer 1987, 1997 in Mayo et :tl.
High and diverse chromosome numbers in 1997).
Araceae are thought to have been derived by aneu-
ploidy and polyploidy. Thus the aneuploid series PoLLINATION. Araceae inflorescences are almost
x = 11, 10, 9, 8 (and probably 13, 12) could have always insect pollinated, although "wind-tunnel"
been derived by reduction, and x = 15, 16, 17, 18, pollination has been proposed for Pinellia (Uhlarz
19, 20, 21 by an increase in chromosome number 1985). Pollinators so far reported for Araceae
from a primitive (but possibly secondary) base (Grayum 1984) include trigonid bees (Mons,'era,
number of x = 14. Spathiphyllum), euglossine bees (Anthur,'um,
Diploids, triploids, tetraploids and still higher Spathiphyllum), beetles of the families Asilidae
levels of ploidy can be found within one species, as (Amorphophallus ), Cetoniidae (Amorphophallu.; ),
in Cryptocoryne ciliata with diploids and triploids Curculionidae (Anthurium, ?Pistia), Dermestidae
(2n = 22, 33), and Cryptocoryne cordata with (Dracunculus), Nitidulidae (Alocasia, Amcr-
2n = 34,68, 85,102. phophallus, Anchomanes, Anubias, Aridarum,
 Araceae 33

Cercestis, Culcasia, Cyrtosperma, Nephthytis, and many Spathiphyllum species have pure white
Typhonium), Scaphidiidae (Pseudohydrosme), spathes.
Scarabaeidae (Alocasia, Amorphophallus, Odour is evidently a prime factor in attracting
Anubias, Caladium, Dieffenbachia, neotropical pollinators, and Araceae are famous for their foul
Homalomena, Philodendron, Syngonium), Scyd- blossom odours. Many genera, however, are
maenidae (Typhonium), Silphidae (Amorpho- less offensive. Spathiphyllum, Philodendron and
phallus), Staphylinidae (Alocasia, Anthurium, Xanthosoma have heavy, spicy odours and in An-
Chlorospatha, Dracunculus, Lysichiton, Pseu- thurium, odours range from spicy to the smell of
dohydrosme, Typhonium), and flies of the fam- decaying apples. Amorphophallus galbra is known
ilies Anthomyiidae (Alocasia), Calliphoridae to have a pleasant smell. Few studies of odour
(Dracunculus, Helicodiceros), Ceratopogonidae chemistry have been made, and this field is still in
(Arum, Cryptocoryne), Chloropidae (Peltandra), its beginnings.
Choridae (Pseudohydrosme), Drosophilidae Inflorescence heating (thermogenesis) in con-
(Alocasia, Colocasia, Culcasia, Homalomena, nection with flowering has been examined mainly
Nephthytis), Ephydridae (Cryptocoryne), Myceto- in Alocasia, Arum, Philodendron, Sauromatum
philidae (Arisaema, Arisarum), Neurochaetidae and Symplocarpus. This phenomenon appears to
(Alocasia), Phoridae (Cryptocoryne), Psychodidae be fairly widespread in the family, although by no
(Arum), Sciaridae (Arisaema, Arum), Simuli- means universal. Its function is generally agreed
idae (Arum), Sphaeroceridae (Arum, Pseudo- to be volatolisation of odour compounds. Attrac-
hydrosme) and Syrphidae (Peltandra). tion to a warm site is also thought to have a role,
The kettle trap mechanisms (in German litera- at least in cases involving carrion beetles and flies.
ture referred to as Kesselfallenblumen) of genera In many genera, however, odours are produced
with unisexual flowers and complex spathes apparently without heating.
(e.g. Arisaema, Arisarum, Arum, Cryptocoryne, Odour production and thermogenesis (when
Lagenandra, Philodendron, etc.) have attracted present) take place mainly in terminal appendices
most attention. Much less is known about pollina- (e.g. Arum) or in the male or sterile male zones of
tion mechanisms in genera with bisexual flowers the spadix (e.g. Philodendron). Odour production
and simpler or spreading spathes (e.g. Anthurium, from the spathe is suspected in Dracontium and
Monstereae, Lasioideae). While kettle "trap" probably occurs in other genera. The thickened
seems an adequate description for Arum and connectives of stamens, staminodes, synandria
Arisaema, in other genera (e.g. Amorphophallus, or synandrodes in Philodendron, Anubias,
Philodendron) it is unclear whether the pollinators Homalomeneae, Caladieae and Colocasieae prob-
are unable or merely unwilling to leave the inflo- ably represent adaptations for osmophore func-
rescence once they have entered it, kept there tion. In Cryptocoryneae, the so-called olfactory
perhaps by attractants such as stigma secretions, bodies are probably osmophoric, although Vogel
food bodies, pollen, odour compounds (male (1963) found osmophores on the spathe limb
euglossine bees) or sites for reproduction. in Cryptocoryne. In genera with well-developed
The colour of the spathe, and to a lesser extent of terminal spadix appendices (e.g. Areae, Arisae-
the spadix, varies considerably within the family, mateae, Thomsonieae), thickened connectives are
ranging from inconspicuous greens (e.g. Anthu- generally absent. Odour production in genera
rium, Nephthytis) to elaborate patterns (e.g. with bisexual flowers is poorly understood. In
Colletogyne, Sauromatum) or striking "flags" Spathiphyllum the stigma is thought to play this
(e.g. Anthurium andraeanum). Fleshlike colours role and in Anthurium the thickened tepal apices
play an important role in attracting pollinators may be involved.
(myiophilous colours) in some genera. Differenti- Araceae are always protogynous and the female
ated colour zones are frequent. In Philodendron (stigma receptivity) and male (anther dehiscence)
many species have purple zones inside the spathe phases usually do not overlap, so that obligate
tube, while the blade is white or pale green. In outcrossing is the general rule. Some cases of self-
Arisaema and Arisarum, the reverse situation is pollination or apomixis are known or suspected
found, with white spathe tubes and dark purple (Anthurium gracile, triploid Amorphophallus
blades which are often striped. The foul odours of paeoniifolius, Arum idaeum, A. hygrophilum,
such species as Amorphophallus konjac are very Pinellia).
often associated with flesh-coloured or livid Manipulation of pollinator behaviour within the
spathes, resembling carrion. By contrast, the per- inflorescence may be the basis for many of the
fumed inflorescences of Zantedeschia aethiopica specialised features of the spadix, particularly in
34 Araceae

unisexual-flowered genera. Spathe constnctwns the inner layer of the pericarp may also be
may act as "skid zones" (Arum) or, in conjunction mucilaginous. This makes the seeds sticky and
with closing movements, as "brooms" to eject aids their dispersal by birds or mammals.
pollinators from the female chamber after pollina- The seeds are usually straight, but in Lasieae,
tion (Philodendron). The various types of hairs, Monstereae and Pothoideae they may be curved
scales and warts found on the inner surface of the or even strongly horseshoe- or kidney-shaped
spathe (e.g. Amorphophallus) or the wide range of (anacampylotropous). Endosperm may be abun-
staminodial or pistillodial structures on the spadixdant or absent and all intermediate states exist.
(e.g. Areae, Bucephalandra) have less obvious Sometimes the endosperm is reduced at seed ma-
functions; in Arum the filamentous pistillodes and turity to a single cell layer. The embryo is usually
staminodes are thought to exclude inappropri- hardly differentiated. Rarely, the plumule is highly
ately large insect visitors and in Dieffenbachia, the
developed, e.g. as in Peltandra and Typhono-
staminodes are food bodies (Young 1986). In dorum. In Cryptocoryne ciliata, which grows
Helicodiceros the spathe and spadix hairs may act in freshwater tidal zones, between 20 and 40
to give the inflorescence the appearance of a mam- cataphylls are formed in the embryo and these
malian corpse. probably serve to fix the seed to the substrate,
Observations have been made of various sec- preventing it from being swept away with the tide.
ondary associations between insects and aroid in- Seeds with well-developed plumules, except
florescences, which do not involve pollination. Cryptocoryne, usually lack endosperm and have
Some species of drosophilid flies are known to only a very thin, papery testa. The testa may be
breed on the inflorescences of Alocasia, Colocasia smooth or variously sculptured, thin to thick or
and Homalomena (Okada 1986). They also exhibit occasionally absent (Gymnostachys, Nephthytis,
specialisation in their behaviour on the spadix: Orontium). In some Lasieae the testa is strongly
stamenicolous species lay their eggs in the male warty, hard and thickened other genera (Areae)
zone, pistilicolous species in the female zone. Fur-have strophiolate seeds, while in Pistia the seed
thermore, several different pairs of fly species, one
has a true operculum.
stamenicolous and the other pistilicolous, are The Araceae have the most varied endosperm
known to breed in association on one aroid spe- of any family. Genera conservative in this re-
cies (synhospitality). Hymenopteran parasitism spect, such as Epipremnum and Spathiphyllum,
in Philodendron inflorescences has also been possess abundant starch-free endosperm rich
observed (Mayo 1989). in lipids and aleurone. Accumulation of starch
is a feature of many of the specialised genera,
FRUIT AND SEED (see Seubert 1993). The fruits of beginning with Anthurium, where the endo-
the Araceae are usually juicy berries (more rarely sperm cells contain both lipids and starch grains,
rather dry and firmly carnose or leathery), densely leading to pure starch endosperms (e.g.
packed into a cylindric (sometimes subspherical) Xanthosoma), and finally to the "cereal type
infructescence. The berries are normally free, but endosperm" differentiated into a peripheral
in Syngonium they form an indehiscent syncarp, aleurone layer sharply delimited from the lipid-
and in Cryptocoryne a syncarp which opens free starchy remainder, as in Areae and Pistia.
apically, releasing the seeds. The berries are usu- Another trend leads to reduction and finally
ally indehiscent and may decay at maturity, but loss of endosperm, as in Lasia and Lysichiton,
the ripe berries of Lagenandra actively open at the where the endosperm is reduced to a single cell
base to release the seeds. The colour may be vivid layer, or Dieffenbachia, Orontium and Pothos,
orange (Nephthytis) to red (Arum) or purple-red where no trace of endosperm is left in the ripe
(Sauromatum) or white (many Philodendron spp., seeds. In a few genera (e.g. Anthurium, Gymno-
most Stenospermation spp., some Biarum spp.), stachys, Zomicarpa) the endosperm is chloro-
sometimes green (Arophyton buchetii, Lysichi- phyllous, the embryo more frequently so.
ton, Orontium, Peltandra virginica, Pinellia) or Chlorophyllous embryos occur both in genera
yellow (Typhonodorum), rarely blue (Gymno- with storage embryos (i.e. seeds lacking en-
stachys, Amorphophallus yunnanensis) or brown- dosperm, as in Anaphyllum and Pothos) and in
ish (!asarum). genera with a well-developed endosperm (e.g.
The seeds are often embedded in a mucilaginous Ambrosina, Arisarum, Calla, Dracunculus).
pulp secreted by the ovular and placental Gymnostachys is unique in having both green
trichomes. In many epiphytic Anthurium species endosperm and an embryo.
 Araceae 35

DISPERSAL. Dispersal of the genera with red- or west to E Africa and east to N America.
orange-coloured berries is probably carried out Amorphophallus, Remusatia, Rhaphidophora and
mainly by birds, though data are scarce. Mature Sauromatum are shared between tropical and
berries have been observed to stay fresh for a subtropical Africa and Asia. Amorphophallus
relatively long time while awaiting dispersal and Remusatia also occur in Madagascar.
(Arum, Nephthytis). In many genera (Cala- Pathos is found in Madagascar as well as in
dium, Colocasia, Philodendron, Syngonium), tropical Asia and NE Australia. Lysichiton and
the infructescences are markedly ephemeral, Symplocarpus occur in NE Asia as well as in tem-
disappearing almost as soon as they are ex- perate N America. Calla and Lysichiton extend
posed. Dispersal by ants (myrmecochory) has northwards to the subarctic zone. An unusual
been observed in Biarum and some species of geographical pattern is represented by the tribe
Amazonian Araceae (e.g. Anthurium ernestii, Peltandreae, which consists of Peltandra (eastern
Philodendron megalophyllum) are reported to N America) and Typhonodorum (Madagascar).
be associated with ant-gardens. Dispersal by Perhaps the most interesting feature of aroid
water (hydrochory) is also known (Cryp- geography is the distribution of three genera
tocoryne, Lagenandra, Montrichardia, Pistia, Homalomena, Schismatoglottis and Spathiphyl-
Typhonodorum). Dispersal by mammals, includ- lum which are disjunct between tropical SE Asia or
ing primates, has been observed (e.g. Anchomanes Melanesia on the one hand and tropical America
difformis - chimpanzee; Philodendron bipin- on the other. These genera are rainforest herbs
natijidum monkeys, bats; Xanthosoma whose long-distance dispersal by water or far-
robustum - bats). The seeds of Araceae, particu- ranging animal vectors is implausible. It is also
larly those with large chlorophyllous embryos, difficult to imagine them being rafted by the
do not remain viable for a long time. Bulbils southern Gondwanic route via Antarctica or via
with hooks or hairlike processes are found in the Bering Straits because of their intolerance of
Remusatia, which must be dispersed by birds or subtropical conditions, much less temperate cli-
mammals. mates. No really plausible historical explanation
A number of species which are important as has yet been proposed for these disjunctions. It is,
human food (Alocasia macrorrhizos, Colocasia nevertheless, tempting to speculate that these pat-
esculenta, Cyrtosperma merkusii, Xanthosoma terns are the relicts of a once-continuous distribu-
sagittifolium complex) or ornamentals (Epiprem- tion during the Cretaceous.
num pinnatum 'Aureum', Monstera spp., Philo-
dendron spp., Syngonium spp., Zantedeschia FOSSIL RECORD. Gregor and Bogner (1984, 1989),
aethiopica) have been widely dispersed through- Herendeen and Crane (1995) and Mayo et al.
out the tropics by man and have become (1997) have reviewed the literature on Araceae
naturalised as well. Some Typhonium species, fossils. Fossil leaves, spadices and infructescences,
possibly dispersed widely because of their medici- fruits and seeds are well documented from the
nal value, have become weedy in many tropical Eocene onwards; Spathiphyllum-like pollen has
regions. been described from the Palaeocene.

DISTRIBUTION AND HABITATS. The genera of
Araceae are concentrated in the tropics of
America and SE Asia. Tropical and southern
Africa is the next richest region, followed
in order of decreasing diversity by temperate
Eurasia, Madagascar and the Seychelles, and N
America (including northern Mexico). Australia
has only one endemic genus (Gymnostachys).
The other native Australian Araceae represent
an extension of the tropical Asian araceous
flora.
The majority of genera are endemic to the major
continental regions, but some extend further;
Pistia is pantropical and Calla is circumboreal.
Arisaema is most diverse in SW China and extends
EcoLOGY AND LIFE FoRMS. The Araceae include
a wide range of life forms: (1) terrestrial plants
with aerial stems; (2) geophytic plants with sub-
terranean stems modified into subglobose tubers
(sometimes cormlike) or thickened rhizomes; (3)
root climbers, epiphytes, hemiepiphytes and litho-
phytes; and (4) aquatic and semiaquatic plants.
The least specialised life form appears to be that
shown by terrestrial rainforest genera such as
Aglaonema, Chlorospatha, Dieffenbachia, Homa-
lomena, Schismatoglottis and Steudnera. The stem
of such plants is aerial, decumbent or erect, with
short, green internodes. In some other genera,
certain species may exhibit this habit. e.g. Anthu-
rium, Culcasia.
36 Araceae
About 1/3 of the genera of Araceae prefer wet to
swampy conditions, or grow near open water.
Among the most specialised aroids are the
rheophytes, which are flood-resistant plants grow-
ing up to the flood level in or along swift-running
streams or rivers, a habitat in which competition
from other plants is low. They typically have
narrow, leathery leaves and short stems, cling-
ing tightly to rocks. All species of the genera
Aridarum, Bucephalandra, Hottarum and Pipto-
spatha are true rheophytes while other genera
contain a few species which are true or facultative
rheophytes, as in Homalomena and Schis-
matoglottis.
]asarum steyermarkii and many Cryptocoryne
species are rhizomatous, permanently submerged
hydrophytes. In flower, the inflorescence as a
whole (]asarum) or the tip of the spathe
(Cryptocoryne) is held above the water surface.
Pistia stratiotes is the only free-floating aroid.
Many Araceae are helophytes (emergent rooted
aquatics or swamp plants), which show a great
range of habit, including rhizomatous, creeping,
arborescent or with simple, erect, aerial stems.
Typhonodorum, Lasiomorpha and Montrichardia
form extensive pure stands along rivers and in
freshwater lagoons. Aglaodorum griffithii occurs
in the freshwater tidal zone in both fresh and
brackish water.
Many of the geophytic species grow in decidu-
ous forests or woodlands in seasonally dry cli-
mates, but some also grow in rainforests (e.g.
many Amorphophallus and Dracontium spp.).
Only one species of Araceae is known from true
desert conditions (Eminium spiculatum subsp.
negevense), but several genera are represented in
semiarid areas, e.g. Arum and Eminium inC Asia,
Arum and Biarum in N Africa and the Orient,
Arisaema and Sauromatum venosum in the Ara-
bian Peninsula and E Africa, and a few species of
Stylochaeton from the Sahel zone of Africa. In the
northern temperate region, the tuberous life form
enables genera like Arisaema, Arum and Pinellia
to survive the winter by means of dormancy.
These genera usually grow in woodlands or in
shady places under trees or shrubs. A few tuber-
ous genera are found at high altitudes in tropical
regions where warm temperate climates exist,
as, for example, in the Andes (Gorgonidium to
ca. 3000m) or African mountains (Arisaema
ruwenzoricum up to 3000m in the Ruwenzori).
The highest altitude is reached in the Himalayas,
where Arisaema lobatum and A. jacquemontii
grow up to 4500 m and A. flavum to 4400 m.
Some Araceae grow in association with rocks
(lithophytes, epilithic or saxicolous plants), with
their stems, rhizomes or tubers growing in humus
deposits in holes or crevices of steep outcrops (e.g.
Colocasia gigantea, Typhonium albispathum in
limestone hills in Thailand and the Malay Penin-
sula) or in eroded limestone cavities in Mada-
gascar (Amorphophallus ankarana, Colletogyne),
or on granitic or sandstone mountains in Eastern
Brazil (thick- and brittle-leaved Anthurium
species such as A. erskinei). Limestones, with their
eroded cavities, are especially favourable for the
collection of vegetable debris and leaf litter, and
Araceae often prefer these habitats.
The root climbers and hemiepiphytes normally
grow in humid forests, often very high in the
canopy. When losing connection with the forest
floor, they may send down long roots to the
ground; such plants may be referred to as
secondary hemiepiphytes. Hemiepiphytes and
root climbers cling tightly to the bark of the host
with negatively phototropic roots. Skototropism,
an orientation movement towards a well-defined
dark area, has been observed in Monstera tenuis
and other genera and is an adaptation in juvenile
plants for seeking a suitable host tree, following
germination on the forest floor.
True epiphytes are known in Anthurium, Philo-
dendron and Stenospermation. Here the plants are
always independent of the ground, their seeds ger-
minating directly on the host tree after dispersal,
probably by birds. The bird's nest epiphytes of
Anthurium sect. Pachyneurium and in the genus
Philodendron (e.g. P. insigne) collect leaflitter and
humus in their leaf rosettes and root balls.
Remusatia vivipara is a common epiphyte in the
Old World, apparently dispersed by its hooked
bulbils.
CoNSERVATION. The main threat to the long-
term survival of many Araceae species is the loss
and reduction in quality of their natural habitats,
especially the rainforests of tropical Asia, Africa
and America. Some Araceae particularly highly
adapted to specific habitats cannot survive
the transformation of their environments; e.g.
Chlorospatha and rheophytic species of the tribe
Schismatoglottideae. These taxa have proved diffi-
cult to cultivate and are unlikely to survive in the
long term in botanic gardens. This is also true
of forest aquatics such as ]asarum (found only
in blackwater rivers) and several species of
Cryptocoryne. Currently, no species of Araceae is
listed in CITES (Convention on International
Trade in Endangered Species).
 Araceae
EcoNOMIC IMPORTANCE. Colocasia esculenta
(taro), with edible tubers and leaves, is the major
traditional staple food of the Pacific Islands and an
important subsistence food crop throughout the
tropics. Xanthosoma sagittifolium (cocoyam;
treated here in the broad sense including various
races and possibly species), originally from tropi-
cal America, is grown throughout the tropics
for its tuberous stems which are even more nutri-
tious than those of Colocasia. Amorphophallus
paeoniifolius (elephant yam) is cultivated for its
starch-rich tubers in tropical Asia, particularly
India. Amorphophallus konjac (konjac) is impor-
tant in Japan, where "Konnyaku", a tasteless gel-
ling agent much used in food, is made from the
glucomannan-rich tubers. Cyrtosperma merkusii
is grown from SE Asia to Oceania as a famine
food or for animals, and likewise Alocasia
macrorrhizos. Typhonodorum lindleyanum seeds
are cooked and eaten in Madagascar, and
Montrichardia seeds are roasted and eaten by
the Amerindians of tropical South America.
Many Araceae are poisonous when fresh, and in
virtually all edible species, the stems, rhizomes,
tubers and leaves must be cooked before they
can be consumed (Alocasia, Amorphophallus,
Colocasia, Cyrtosperma, Xanthosoma). In classical
Greece the starchy tubers of Arum italicum
were used as food and in the Middle Ages Arum
maculatum tubers were used as famine food
in England. The infructescences of Monstera
deliciosa are delicious when fresh, with a
pineapple-like flavour.
The frequent use of numerous different Araceae
by indigenous people in folk medicine, as arrow
poisons etc. may indicate the rich occurrence in
the family of biodynamic substances.
Aerial roots of Heteropsis and Philodendron are
or have been a source of fibre for Amerindians in
tropical South America, the former being particu-
larly highly regarded. Even today, roots of
Heteropsis species are used for basket-weaving
in Amazonia.
The Araceae include many very handsome orna-
mental plants and are cultivated everywhere. In
temperate regions, the genera Arisaema and
Arum, and to a lesser extent Biarum, Lysichiton,
Orontium and Symplocarpus, provide much-loved
perennial herbs for woodland, rock and bog
gardens. Arisaema species are particularly valued
for this purpose in China and Japan and are.
increasingly sought after in Europe as woodland
garden plants. Some aroids are ubiquitous
house plants and in the tropics are also grown
outside in gardens, e.g. species of Aglaonema,
37
Anthurium, Caladium, Dieffenbachia, Epiprem-
num, Monstera, Philodendron, Spathiphyllum,
Syngonium and Zantedeschia. Some large and
especially beautiful tropical species have been
important for modern garden design in the trop-
ics, particularly in the work of the famous Brazil-
ian landscape architect, Roberto Burle Marx.
Anthurium andraeanum, Zantedeschia aethiopica
and their hybrids are an important source of cut
flowers, and the former is the basis of an impor-
tant industry in Hawaii, the Netherlands and else-
where. Several species of Cryptocoryne are much
prized as aquarium plants. In general, it may be
said that the Araceae are probably the most im-
portant foliage ornamental plant group in horti-
culture, worldwide. A fuller account of economic
and medicinal aspects of Araceae is given by Bown
{1988).
SUBDIVISION AND RELATIONSHIPS. Gymno-
stachys differs markedly in from all other Araceae.
The Spathiphylleae are conservative morpho-
logically and the Orontioideae and Lasioideae
have many primitive characters, notably bisexual,
perigoniate flowers. Cladistic analyses based
on morphological and molecular data (French
et al. 1995; Mayo et al. 1997) showed that all
genera with unisexual flowers form one mono-
phyletic group, recognised here as subfamily
Aroideae. The new classification of Mayo et
al. {1997), based on these studies, is adopted
here.
AFFINITIES. The affinities of the Araceae have
been considered obscure until recently. Compre-
hensive reviews by Grayum {1984-1992) and the
cladistic studies of French et al. ( 1995) and Mayo
et al. {1997) suggest the following as ancestral
characters for Araceae: laticifers absent, raphides
present; the stem short, upright, unspecialised
either as a climber or a geophyte; leaves pro-
bably simple, bifacial, with an expanded blade
(?distichously arranged) and a geniculate petiole;
finer leaf venation may have been either reticulate
or parallel-pinnate; inflorescence a spike (axis not
necessarily fleshy), subtended by a relatively in-
conspicuous bract (spathe); flowers bractless, bi-
sexual, 3-merous (?2-merous), with 2 whorls of 3
(?2) free tepals, 2 whorls of 3 (?2) free stamens,
anthers terminal, basifixed, extrorse, on ± elon-
gated filaments, thecae dehiscing by longitudinal
slits, tapetum probably periplasmodial, pollen
monosulcate and ellipsoid, starchless, binucleate;
ovary superior, syncarpous, 3-locular, with several
anatropous ovules per locule and axile placenta-
38 Araceae
tion; the seeds have endosperm with cellular
of free-nucleate development in the micropylar
chamber and a 1-celled haustoria! chalazal
chamber, embryo small with caryophyllad or
solanad embrogeny; base chromosome number
x = 7 or 14 (or 12). Following the proposal by
Tillich (1985) and the reviews by Grayum (1987,
1990), Acorus has been removed from Araceae
to its own family. Strong support for this change
has come from the molecular studies of Duvall
et al. (1993) and Chase et al. (1995), who placed
it at the base of the monocot clade. It differs
from Araceae in a number of important char-
acters: unifacial (ensiform) leaves, specialised
idioblasts containing essential oils, 2 separate
vascular systems in the peduncle, introrse anthers
(in Araceae known only in Zamioculcas), stellate
pattern of endothecial thickenings, distinct mor-
phology of ovular trichomes and their location
on the outer and inner integuments, presence of
perisperm, absence of raphides, secretory anther
tapetum, unique bud trace insertion pattern
and absence of dorsal vascular bundles in the
gynoecium.
The Lemnaceae have been considered by most
authors to be closely related to Araceae and to
have evolved from them by neotenous reduction.
This view was based mainly on the similarity
between Pistia seedlings and Spirodela, embryo-
logical characters, the putative homology between
the aroid spathe and the "spathe" of Spirodela
and Lemna, presence of grooved raphides, oper-
culate seeds and a similar pattern of germina-
tion in Pistia and Lemnaceae. More recently, new
lines of evidence confirming the close link be-
tween the two families have emerged. French et
al. (1995), in a detailed cpDNA study, showed that
Lemnaceae are embedded within the Araceae,
while the fossil genus Limnobiophyllum (Stockey
et al. 1997) appears to bridge the gap between
Araceae and Lemnaceae. There is a growing body
of opinion that favours the formal reduction of
Lemnaceae to a subgroup of Araceae (e.g. Mayo
et al. 1997). The pollen morphology differs signifi-
cantly, however; in Lemnaceae the pollen is ulcer-
ate and spinose while in Pistia it is inaperturate
and plicate.
Earlier authorities, such as Engler and Gilg
(1919) and Wettstein (1935), grouped the Araceae
together with the Palmae and Cyclanthaceae. This
view was based on the common occurrence in
these families of a tendency to floral reduction,
"spathe" development, the condensation of the
inflorescence into a spadix or spadices and an as-
sociated tendency to pseudanthial inflorescences.
Clear evidence of the "monocot type" was
recognised in the floral structure, but an inde-
pendent evolution from a primitive monocot form
was regarded as the best explanation. Engler
(1920a) went so far as to state that the most primi-
tive Araceae (his subfamily Pothoideae) differed
from the Liliaceae essentially only in having a
fleshy outer seed integument. In his classification,
however, Engler (e.g. Engler and Gilg 1919) did
not associate Araceae and Liliaceae closely and
it is reasonable to suppose that he saw the simi-
larities between them in terms of shared primi-
tive characters (plesiomorphies). Hutchinson
(1934) visualised a close relationship with the
Liliaceae. He considered that Araceae evolved
from tribe Aspidistreae of Liliaceae (his circum-
scriptions) due to their common possession of
densely spicate inflorescences. Araceae have also
been associated in the past with Pandanaceae
and Typhales.
Dahlgren and coworkers (Dahlgren and Clifford
1982; Dahlgren et al. 1985) argued for a closer
relationship of Arales (Araceae, Lemnaceae) to the
Alismatiflorae than to any other group, pointing
out many important differences from the palms.
The characters that best link the Alismatiflorae to
the more primitive tribes of Araceae appear to
be extrorse anthers, periplasmodial tapetum
and caryophyllad embryogeny. The Alismatiflorae
also differ, however, in important respects:
absence of raphides, usually apocarpous gyno-
ecium, helobial or nuclear endosperm develop-
ment, non-berried fruits and usually absence of
endosperm.
Grayum (1984) agreed with Dahlgren's views.
His re-interpretation of aroid endosperm devel-
opment as helobial further emphasises a relation-
ship with Alismatiflorae. Recent molecular studies
by Chase et al. (1995) have further strengthened
the link between Arales and Alismatiflorae. Their
cladograms show the two taxa as sister groups,
forming a well-supported clade near the base of
the monocotyledons.
CLASS! FICA TION OF THE ARACEAE
a. Flowers bisexual
I. Subfamily Gymnostachydoideae Bogner & Nicolson
(1991).
(genus 1)
II. Subfamily Orontioideae Mayo, Bogner & Boyce (1997).
(genera 2-4)
III. Subfamily Pothoideae Engler (1876).
1. Tribe Potheae Engler (1876).
(genera 5-7)
2. Tribe Anthurieae Engler (1876).
(genus 8)
 Araceae
IV. Subfamily Monsteroideae Engler (1876).
1. Tribe Spathiphylleae Engler (1887).
(genera 9-10)
2. Tribe Anadendreae Bogner & French (1984).
(genus 11)
3. Tribe Heteropsideae Engler (1906).
(genus 12)
4. Tribe Monstereae Engler (1876).
(genera 13-20)
V. Subfamily Lasioideae Engler (1876).
(genera 21-30)
VI. Subfamily Calloideae Endlicher (1837).
(genus 31)
b. Flowers unisexual
VII. Subfamily Aroideae Engler (1887).
1. Tribe Zamioculcadeae Engler (1876).
(genera 32-33)
2. Tribe Stylochaetoneae Schott (1856).
(genus 34)
3. Tribe Dieffenbachieae Engler (1876).
(genera 35-36)
4. Tribe Spathicarpeae Schott (1856).
(genera 37-44)
5. Tribe Philodendreae Schott (1856).
(genus 45)
6. Tribe Homalomeneae (Schott) M. Hotta (1970).
(genera 46-47)
7. Tribe Anubiadeae Engler (1876).
(genus 48)
8. Tribe Schismatoglottideae Nakai ( 1943).
(genera 49-55)
9. Tribe Cryptocoryneae Blume (1836).
(genera 56-57)
10. Tribe Zomicarpeae Schott (1856).
(genera 58-61)
11. Tribe Caladieae Schott (1832).
(genera 62-68)
12. Tribe Nephthytideae Engler (1887).
(genera 69-71)
13. Tribe Aglaonemateae Engler (1876).
(genera 72-73)
14. Tribe Culcasieae Engler (1887).
(genera 74-75)
15. Tribe Montrichardieae Engler (1876).
(genus 76)
16. Tribe Zantedeschieae Engler (1887).
(genus 77)
17. Tribe Callopsideae Engler ( 1906).
(genus 78)
18. Tribe Thomsonieae Blume (1837).
(genera 79-80)
19. Tribe Arophyteae Bogner (1972).
(genera 81-83)
20. Tribe Peltandreae Engler ( 1876).
(genera 84-85)
21. Tribe Arisareae Dumortier (1827).
(genus 86)
22. Tribe Ambrosineae Schott (1832).
(genus 87)
23. Tribe Areae Engler (1876).
(genera 88-96)
24. Tribe Arisaemateae Nakai (1943).
(genera 97-98)
25. Tribe Colocasieae Engler (1876).
(genera 99-104)
39
26. Tribe Pistieae Blume (1836).
(genus 105)
KEY TO THE GENERA OF ARACEAE
AND AcoRACEAE
1. Plants free-floating aquatics; leaves rosulate, hairy;
flowers unisexual, naked; inflorescence with a single
female flower and a few male flowers 104. Pistia
- Plants terrestrial or helophytes, climbers, hemiepiphytes,
epiphytes or lithophytes or other but never floating 2
2. Leaves not differentiated into petiole and blade, primary
venation strictly parallel; inflorescence borne on a
culmlike axis 3
- Leaves with distinct petiole and expanded blade, primary
venation never strictly parallel 4
3. Leaves ensiform, unifacial; spadix solitary, pseudolateral
and overtopped by a single, erect, leaflike spathe; flowers
3-merous, tepals 6 Acorus (Acoraceae)
- Leaves dorsiventrally flattened, bifacial; flowering shoot
with long culmlike axis, bearing numerous spadices dis-
tally, these borne in axillary clusters subtended by elon-
gate bracts; flowers 2-merous, tepals 4 1. Gymnostachys
4. Flowers with obvious perigone of free or fused tepals
(except Pycnospatha which lacks perigone, but has
dracontioid leaf, tuberous stem and boat -shaped, forni-
cate spathe - see lead 22) 5
- Flowers without perigone of free or fused tepals 27
5. Flowers bisexual, spadix uniform in appearance with
flowers of only one type 6
- Flowers unisexual, spadix clearly divided into basal
female zone and apical male zone; tropical Africa 25
6. Higher-order leaf venation parallel to primary lateral
veins; tissues with abundant trichosclereids 7
- Higher-order leaf venation clearly reticulate; tissues
without trichosclereids or trichosclereids very few 8
7. Spathe persistent; tepals free or connate; ovary 2-4-
locular; ovules 2-8 per locule, placenta axile
9. Spathiphyllum
- Spathe deliquescent; tepals connate; ovary 1-locular;
ovules several, placenta basal 10. Holochlamys
8. Stem aerial, not tuberous or rhizomatous, never aculeate;
plant usually a climber, hemiepiphyte or epiphyte, less
often lithophyte or terrestrial, only very rarely helophytic
(some spp. of Anthurium) 9
- Stem typically subterranean, tuberous or rhizomatous,
sometimes aerial and creeping or scrambling, but then
aculeate; plant frequently a helophyte 13
9. Neotropical plants; seeds with copious endosperm;
pollen usually forate, never monosulcate 8. Anthurium
- Palaeotropical plants; seeds without endosperm; pollen
monosulcate or inaperturate 10
10. Stigma transversely oblong; stamens always 4 per flower;
pollen inaperturate; perigone consisting of a single
cuplike structure 11. Anadendrum
- Stigma hemispheric to discoid; stamens usually 6 per
flower; pollen sulcate; perigone usually consisting of free
tepals or when connate and cuplike the flowers are borne
on short pedicels 11
11. Ovary 3-locular; locules 1-ovulate; flowering shoot with
inflorescences always axillary 12
- Ovary 1-locular; flowering shoots terminating in a
branching system of spadices 7. Pothoidium
12. Flowers sessile; tepals free, very rarely basally united
5. Pothos
- Flowers pedicellate; tepals connate 6. Pedicellarum
40 Araceae
13. Plants of temperate regions (N America, NE Asia); leaf
blade always entire, ovate to elliptic 14
- Plants of tropical and subtropical regions; leaf blade
sagittate, pinnatifid, pinnatisect or dracontioid 16
14. Ovary 2-locular; ovules 2 per locule, placentation axile
3. Lysichiton
- Ovary 1-locular; ovule 1, placenta apical or basal 15
15. Placentation basal; spathe inconspicuous; spadix cylin-
dric, stipe very long 2. Orontium
- Placenta apical; spathe thick, ventricose, enclosing
spadix; spadix subglobose, stipe short 4. Symplocarpus
16. Leaf deeply sagittate, anterior division not pinnatifid or
pinnatisect 17
- Leaf blade pinnatifid, pinnatisect, dracontioid or some-
times :±:: pedatifid; anterior division always pinnately
divided, either pinnatifid, pinnatisect or yet more highly
divided 21
17. Ovary many- to 2-ovulate, rarely 1-ovulate; seeds with
endosperm 18
- Ovary 1-ovulate, rarely 2-ovulate; seeds without en-
dosperm or rarely with a little endosperm 19
18. Plants without stolons; petiole spines dispersed; stamen
filaments free; tropical Asia to Oceania 26. Cyrtosperma
- Plants with long stolons; petiole spines in ridges; stamen
filaments free or connate; tropical W Africa
27. Lasimorpha
19. Petiole aculeate, with obvious spines; Malay Archipelago
28. Podolasia
- Petiole smooth to scabrid-verrucose, never aculeate;
tropical America 20
20. Leaf blade never perforated; spathe lanceolate, very long-
acuminate and usually spirally twisted; ovary locules with
(1- )2 to several ovules; neotropics but not east coastal
Brazil 30. Urospatha
- Leaf blade often with a few perforations of irregular size
between primary lateral veins; spathe fornicate; endemic
to east coastal Brazil 22. Dracontioides
21. Stem aculeate, aerial and scrambling to prostrate, intern-
odes distinct, green 29. Lasia
- Stem not aculeate, subterranean, internodes very abbre-
viated, not green 22
22. Leaf blade dracontioid, anterior division bipinnatifid or
yet more highly divided; stem a depressed-globose tuber;
spathe fornicate 23
- Leaf blade pinnatifid, pinnatisect, or sometimes :±::
pedatifid, anterior division pinnatifid to pinnatisect; stem
a vertical or horizontal rhizome; spathe erect, not forni-
cate, blade often spirally twisted apically 24
23. Tropical America; flowers with perigone of 4-8 free
tepals; berries smooth, red 21. Dracontium
- Tropical SE Asia; flowers without perigone; berries ac-
uleate, dark green 24. Pycnospatha
24. Tropical America; testa thick, verrucose; embryo curved
23. Anaphyllopsis
- S India; testa membranous, smooth; embryo straight
25. Anaphyllum
25. Leaf pinnatisect to tri- or quadripinnatifid; tepals free;
spathe margins free 26
- Leaf entire, linear to cordate, sagittate or hastate; tepals
connate into cup; spathe margins connate basally
34. Stylochaeton
26. Leaf blade pinnatisect; stamens free 32. Zamioculcas
- Leaf blade bipinnatifid to quadripinnatifid, at least
in lowest pinnae; stamen filaments connate
33. Gonatopus
27. Flowers bisexual; spadix uniform in appearance with
flowers of only one type (sometimes with sterile flowers
at spadix base) 28
- Flowers unisexual; spadix clearly divided into basal
female zone and apical or intermediate male zone,
flowers very rarely in longitudinal rows (Spathicarpa) 37
28. Helophytes from temperate regions of northern hemi-
sphere; petiole sheath with long apical ligule 31. Calla
- Climbers, hemiepiphytes or sometimes epiphytes from
tropical regions; petiole sheath non-ligulate or ligule only
~~ ~
29. Petiole usually very short with non-annular insertion;
trichosclereids not present in tissues, leaf never per-
forated or lobed; primary lateral veins forming distinct
submarginal vein 8. Heteropsis
- Petiole well developed with annular insertion and usually
conspicuous sheath; trichosclereids present in tissues,
or if absent (or nearly so) then leaf with conspicuously
reticulate higher-order venation and often perforated or
lobed (Amydrium); primary lateral veins usually not
forming distinct submarginal vein 30
30. Trichosclereids rare or nearly absent; higher-order leaf
venation completely reticulate; ovary 1-locular, placenta
1, intrusive-parietal, ovules 2 13. Amydrium
- Trichosclereids abundant; higher-order leaf venation
parallel to primary lateral veins, or only finest venation
reticulate 31
31. Ovary 1-locular or incompletely 2-locular 32
- Ovary 2-5 locular 35
32. Ovules anatropous, more than 1 33
- Ovules amphitropous to anatropous, solitary, basal 34
33. Ovules numerous, superposed on 2 (rarely 3) parietal
placentas; seeds fusiform, straight, 1.3-3.2mm long,
0.6-l.Omm wide 14. Rhaphidophora
- Ovules 2-4 (-6) at base of a single intrusive placenta;
seeds curved, 3-7mm long, 1.5-4.0mm wide
15. Epipremnum
34. Adult leaf blade entire; palaeotropical 16. Scindapsus
- Adult leaf blade pinnatifid; neotropical (Amazonia)
18. Alloschemone
35. Seeds fusiform, claviform or lenticular, less than 3mm
long, endosperm present; ovules (2-)3-many per locule;
leaf blade entire 36
- Seeds globose to oblong, 6-22mm long, the raphe S-
shaped; endosperm absent; ovules 2 per locule; leaf blade
variously shaped, often perforated or pinnatifid or both
17. Monstera
36. Placentation basal; seeds fusiform to claviform; leaf
blade thickly coriaceous 20. Stenospermation
- Placentation axile; seeds lenticular and flattened, strongly
curved; leaf blade mostly membranous
19. Rhodospatha
37. Spadix fused laterally on both sides to spathe and entirely
enclosed by it, forming a septum dividing the spathe into
2 chambers, with a single gynoecium on one side and the
male flowers arranged in 2 rows on the other; very small,
seasonally dormant plants endemic to western
Mediterranean 87. Ambrosina
- Spadix free or fused to spathe in various degrees but
never fused laterally on both sides to spathe to form 2
internal chambers with a single gynoecium on one side
and the male flowers on the other 38
38. Stamens of each male flower free or only the filaments
connate 39
- Stamens of each male flower entirely connate into a dis-
tinct synandrium, synandrium rarely reduced to single
stamen and then endemic to Madagascar ( Colletogyne) or
 Araceae
stamens free and basally connate with remote glo-
bose thecae and then endemic to Andean S America
(Gorgonidium mirabile), or only filaments connate and
then stigma stellate and 5-8-lobed (Spathantheum) 78
39. Spadix never entirely enclosed by spathe in a basal
"kettle" formed of connate spathe margins (or if spathe
margins basally connate, then plant never aquatic) 40
- Spadix entirely enclosed by spathe in a basal "kettle"
formed of connate spathe margins, plants always
helophytic or aquatic 77
40. Higher-order leaf venation parallel-pinnate 41
- Higher-order leaf venation reticulate 53
41. Upper part of spathe persisting as long as lower part;
petiole sheath lacking ligule; ovary !-many-locular;
thecae dehiscing by subapical pores or longitudinal slits;
connective usually conspicuously thickened 42
- Upper part of spathe marcescent or caducous at anthesis,
lower part long-persistent; petiole sheath with long,
marcescent ligule (except most Schismatoglottis spp.);
ovary !-locular; thecae dehiscing by apical pores, connec-
tive not conspicuously thickened 47
42. Spathe variously shaped, never campanulate; plants
tropical American or tropical Asian; peduncle usually
short, if long, then female flowers in single whorl 43
- Spathe obconic to campanulate; plants from southern
Africa (naturalised in America and Asia); peduncle long,
sometimes longer than leaves 77. Zantedeschia
43. Plant always terrestrial, rarely aquatic, never climbing or
epiphytic; inflorescences not secreting resin at anthesis;
endothecium with cell wall thickenings; ovary !-locular
or incompletely 2-5-locular; mostly tropical Asian
(except Homalomena sect. Curmeria) 44
- Plant usually climbers, hemiepiphytes or epiphytes; in-
florescences secreting resin from spathe or spadix at an-
thesis; endothecium nearly always lacking cell-wall
thickenings; ovary completely 2-many locular, placenta
axile to basal; tropical America 45. Philodendron
44. Seed without endosperm, embryo large; ovule 1, placenta
basal or parietal 45
- Seed with copious endosperm, embryo relatively small;
ovules several to many, placenta basal, parietal or axile
46
45. Inflorescence with short peduncle; female flowers in spi-
rals; stem erect to repent; placentation basal; forest plants
72. Aglaonema
- Inflorescence with long peduncle; female flowers in a
single whorl; stem repent; placenta parietal; on tidal
mudflats 73. Aglaodorum
46. Male flower consisting of solitary stamen overtopped by
flask-shaped pistillode; ovary !-locular, placenta basal
46. Furtadoa
- Male flower consisting of ( 1) 2-6 stamens, pistillodes
absent; ovary incompletely 2-5 locular, placentas parietal
and axile or nearly basal 47. Homalomena
47. Placentation parietal; thecae truncate 48
- Placentation basal or basal and apical; thecae truncate or
horned 49
48. Spathe constricted; ovules anatropous to hemi-
anatropous; petiole sheath usually not ligulate; upper
part of spadix usually sterile 49. Schismatoglottis
- Spathe not constricted; ovules hemiatropous to atropous;
petiole sheath with long, marcescent ligule; spadix fertile
almost to apex SO. Piptospatha
49. Thecae truncate; placentation basal 51. Hottarum
- Thecae horned; placentation basal or basal and apical
so
41
50. Stigma smaller than ovary; upper part of spadix sterile
with a distinct appendix of hornless sterile flowers;
spathe constricted or not; stamens never excavated
apically 51
- Stigma as broad as ovary; upper part of spadix mostly
fertile to apex and without a distinct appendix; spathe not
conspicuously constricted; stamens all or mostly exca-
vated apically 52
51. Spathe not constricted; male flowers smooth or verru-
cose; sterile flowers between male and female flowers
flattened 52. Bucephalandra
- Spathe constricted; male flowers densely tuberculate;
sterile flowers between male and female flowers
subcylindric 53. Phymatarum
52. Stamens all excavated; placenta basal 54. Aridarum
- Two lateral stamens of each male flower excavated and
thecae horned, central stamen truncate and thecae horn-
less; placentas basal (fertile ovules) and apical (appar-
ently sterile) 55. Heteroaridarum
53. Leaf blade dracontioid, leaf solitary in each growth
period 54
- Leaf blade shape of various types but never dracontioid;
usually several leaves present 57
54. Petiole usually aculeate; at least some of the ultimate leaf
lobes trapezoid, truncate or shallowly bifid, veins not
forming regular submarginal collective vein on each side
55
- Petiole usually smooth, sometimes rugose but never
aculeate; ultimate leaf lobes usually oblong-elliptic,
acuminate, with primary lateral veins forming regular
submarginal collective veins on each side 56
55. Peduncle long; ovary !-locular 70. Anchomanes
- Peduncle very short; ovary 2-locular
71. Pseudohydrosme
56. Ovary 1-4 locular; terminal appendix smooth, rugose,
rarely verrucose, or staminodial (appendix absent in
Amorphophallus margaritifer and A. coudercii)
79. Amorphophallus
- Ovary always !-locular; terminal appendix staminodial,
separated from male zone by naked axial region
80. Pseudodracontium
57. Spadix fertile to apex, terminal appendix absent 58
- Spadix with :±: smooth terminal appendix 62
58. Helophytes with robust, erect stems and an apical crown
of sagittate to hastate (rarely trisect) leaves; tropical
America 76. Montrichardia
- Terrestrial, climbing or epiphytic plants, leaf blade
variously shaped; tropical Africa 59
59. Leaf blade usually with pellucid resin canals (lines or
points); plants mostly climbers; hemiepiphytes; spathe
boat-shaped, convolute basally; anthers lacking endo-
thecial thickenings 60
- Leaf blade lacking pellucid resin glands; plants terrestrial;
spathe :±: fully expanded, not convolute; anthers with
endothecial thickenings 61
60. Laticifers absent; flagelliform shoots absent; leaf blade
always simple, acute to rounded at base; ovary 1-3-
locular; spadix stipitate or sessile 74. Culcasia
- Laticifers present; flagelliform shoots present; leaf blade
oblong-lanceolate to cordate, sagittate, hastate, trifid or
laciniate to pinnatifid; ovary !-locular; spadix sessile
75. Cercestis
61. Leaf cordate-sagittate or subtriangular, deeply sagittate
or trifid, glabrous; spathe green; spadix entirely free of
spathe 69. Nephthytis
42 Araceae

- Leaf cordate-ovate, minutely hispid abaxially; spathe - Ovary 1-ovulate; female zone of spadix adnate to spathe;
pure white; female zone of spadix adnate to spathe spathe usually with transverse septum between male and
78. Callopsis female zones 96. Pinellia
62. Laticifers anastomosing; tropical S America 63 76. Flowers of both sexes always present in a single inflores-
- Laticifers simple; temperate Eurasia and palaeotropics cence; male flowers 1-androus; lower spathe margins
66 connate; leaf blade ovate or sagittate 86. Arisarum
63. Ovary 6- to 9-ovulate; leaf blade trisect to pedatisect; stem - Flowers of both sexes present in a single inflorescence, or
a subglobose tuber 58. Zomicarpa more often with male and female flowers appearing on
- Ovary 1- to 6-ovulate; leaf blade cordate-sagittate; stem a separate inflorescences; male flowers 2-5-androus; lower
subglobose tuber or rhizome 64 spathe margins convolute; leaf blade normally trisect,
64. Appendix slender 65 pedatisect or radiatisect, rarely simple and ovate
- Appendix relatively thick and subcylindric; stem a creep- 97. Arisaema
ing rhizome; ovary 1-ovulate 60. Ulearum 77. Female flowers spirally arranged (pseudowhorl in
65. Stamen connective much prolonged, threadlike; stem Lagenandra nairii, whorled in L. gomezii) and free;
tuberous; ovary 1-ovulate 61. Filarum spathe tube kettle with connate margins occupying
- Stamen connective not at all prolonged; ovary 1-6- entire spathe tube; spathe blade opening by a straight or
ovulate; stem a creeping rhizome 58. Zomicarpella twisted slit; berries free, opening from base; leaf ptyxis
66. Spadix ± elongated, filiform to clavate with zone of involute 56. Lagenandra
sterile flowers between male and female zones, rarely - Female flowers in a single whorl, connate; spathe tube
with a naked axis between female and male zones of kettle occupying only lower part of spathe tube, remain-
spadix (Arum pictum) or with fertile zones contiguous der also with connate margins (except Cryptocoryne
(Dracunculus) 67 spiral is), blade spreading or twisted; berries connate into
- Spadix usually without sterile flowers (sometimes present a syncarp which opens from the apex; leaf ptyxis
in Arisaema) 75 convolute 57. Cryptocoryne
67. Placenta parietal; leaf blade sagittate or hastate 78. Laticifers simple 79
88.Arum - Laticifers anastomosing 95
- Placentas apical and/or basal; leaf blade variously shaped 79. Synandria connate, thecae of adjacent synandria encir-
68 cling pits in the spadix, each with a somewhat prominent
68. Placentas basal and apical 69 upper margin; leaf peltate; Myanmar to India
- Placenta basal 71 98. Ariopsis
69. Male zone of spadix contiguous with female zone; leaf - Synandria free; leaf not peltate; Africa, Madagascar or
blade pedatifid but lobes not spirally twisted up Americas 80
90. Dracunculus 80. Higher-order leaf venation parallel-pinnate or if reti-
- Male zone of spadix separated from female zone by subu- culate then stem a creeping rhizome and plant from
late to filiform sterile flowers; leaf blade variously shaped tropical S America (Bognera) 81
70 - Higher -order leaf venation reticulate; stem usually a
70. Appendix covered with subulate to setiform sterile flow- subglobose tuber, if rhizomatous then plant Madagascan
ers; leaf blade pedatifid, lobes twisting up on each side 85
91. Helicodiceros 81. Ovules anatropous; primary lateral veins ofleaf forming
- Appendix smooth; leaf blade oblong-lanceolate or a single marginal vein, no submarginal collective vein
sagittate-hastate 92. Theriophonum present; plant from tropical America or continental
71. Lower spathe margins free 72 tropical Africa 82
- Lower spathe margins connate for an appreciable dis- - Ovules atropous to hemiatropous; primary veins of leaf
tance (entirely free in Biarum aleppicum) 73 forming submarginal collective vein and 1-2 marginal
72. Infructescence borne above ground level, berries dark veins; plants from temperate eastern N America or
red to purple, pericarp juicy; sterile zone between male Madagascar 84
and female zones of spadix relatively long, often partially 82. Female zone of spadix free; plant from tropical W and
naked; tropical and subtropical to warm temperate Asia Central Africa 48. Anubias
to Australia 93. Typhonium - Female zone of spadix entirely adnate to spathe; plant
- Infructescence borne at or below ground, berries white to from tropical America 83
pale lilac, pericarp firm, not juicy; sterile zone between 83. Female flowers each with whorl of several staminodes;
male and female zones relatively short and covered en- higher-order leaf venation strictly parallel-pinnate
tirely with subulate sterile flowers; Turkey, western N 35. Dieffenbachia
Africa, Middle East, Central Asia 89. Eminium - Female flowers without staminodes; higher-order leaf
73. Leaf usually solitary, blade pedate; ovary 2-several- venation reticulate 36. Bognera
ovulate 94. Sauromatum 84. Giant herbs (to 4 m) with massive false stem of petiole
- Leaves several; blade linear to ovate, elliptic or obovate; sheaths; staminodes of female flowers free; Madagascar
ovary 1-ovulate 74 (also naturalised in Pemba, Zanzibar and Mascarene Is.)
74. Leaf blade broad elliptic, spathe tube septate 85. Typhonodorum
93. Typhonium pro parte - Relatively small herbs (less than 1m) without false stem;
- Leaf blade linear, ovate, elliptic-oblong or obovate; staminodes of female flowers connate into a cuplike
spathe tube not septate 95. Biarum synandrodium; eastern N America 84. Peltandra
75. Ovary several-ovulate; female zone of spadix free from 85. Madagascan; seed lacking endosperm; ovary 1-locular;
spathe; spathe without a transverse septum separating leaf venation with primary lateral veins forming submar-
male and female zones 76 ginal collective vein and 1-2 marginal veins on each side
86
 Araceae
- S American; seed with abundant endosperm; ovary with
more than 1 locule (except Spathicarpa); leaf venation
with primary lateral veins usually forming single
marginal vein on each side, submarginal collective veins
usually absent 88
86. Stamens either completely connate with marginal thecae
or only partially connate by filaments; bisexual flowers
often present between male and female zones of the
spadix 82. Carlephyton
- Stamens completely connate into truncate synandria or
synandria reduced to 1 stamen 87
87. Synandria reduced to 1 stamen, thecae apical on conical
filament; spadix fertile to apex; leaf blade always cordate
83. Colletogyne
- Thecae apical on a truncate synandrium; spadix appen-
dix present or not; leaf blade cordate, hastate, trifid,
trisect or pedatisect 81. Arophyton
88. Spadix free or only female zone adnate to spathe 89
- Spadix usually entirely adnate to spathe (male zone free
in Spathantheum intermedium) 94
89. Ovules anatropous 90
- Ovules atropous 92
90. Ovules 2 per locule; leaf blade entire, linear to
subsagittate; spadix free, with terminal appendix of
synandrodes 37.~angonia
- Ovules 1 per locule; leaf blade entire, pinnatifid to
subdracontioid; spadix fertile to apex 91
91. Leaf blade pinnatifid to bipinnatifid or subdracontioid;
synandria elongate; stigma capitate or lobed; staminodes
offemale flowers free (connate in Taccarum caudatum)
38. Taccarum
- Leaf blade usually pinnatisect, rarely entire; synandria
short and domed; stigma deeply lobed; staminodes of
female flowers free or connate 39. Asterostigma
92. Styles and synandria elongate 93
- Styles and synandria short and squat; staminodes in
female flowers obovate 42 . .Gearum
93. Staminodes in female flowers filiform to subclavate;
synandria with free filament apices or not; leaf blade
pinnatifid, pinnatisect or bipinnatifid 40. Gorgonidium
- Staminodes in female flowers elongate-triangular;
synandria entirely connate; leaf blade ovate-emarginate
to cordate 41. Synandrospadix
94. Ovary 6-8-locular; female flowers below, male above; leaf
blade entire or pinnately lobed 43. Spathantheum
- Ovary 1-locular; female and male flowers intermixed
(2 central rows of male flowers, 2 outer rows of female
flowers); leaf blade entire 44. Spathicarpa
95. Plants climbers or hemiepiphytes, sometimes creeping
on ground in submature growth, internodes long; berries
connate into a syncarp 67. Syngonium
- Plants terrestrial or geophytic, rarely aquatic, not climb-
ing; internodes very short, berries free from each other
96
96. Spadix without an appendix (present in Hapaline
appendiculata, included here, occasionally absent in
Colocasia esculenta, excluded here) 97
Spadix with an appendix (occasionally absent in
Colocasia esculenta); palaeotropical 104
97. Ovary completely to incompletely 2- to several-locular
with deeply intrusive parietal palcentas (1-locular with
basal placenta in ]asarum, Scaphispatha and a few species
of Caladium and Xanthosoma); neotropical 98
- Ovary clearly 1-locular, placentas not intrusive;
palaeotropical 102
98. Helophytes or terrestrial; leaf blade ovate, sagittate to
43
hastate or pedatifid 99
- Submerged aquatic; leaf blade linear 64. ]asarum
99. Pollen shed in tetrads; style usually laterally thickened
or expanded into a diaphanous mantle; leaf blade entire
or pedatifid 100
- Pollen shed in monads; stylar region not laterally ex-
panded; leaf blade entire or trifid 101
100. Spathe tube subglobose, inflated; female zone of spadix
free; styles normally discoid (laterally swollen) and co-
herent (except Xanthosoma plowmanii); synandrodes
(sterile flowers) between male and female flowers well-
developed, :±:: prismatic 65. Xanthosoma
- Spathe tube narrow, elongate; female zone of spadix
mostly adnate to spathe; stylar region thin, spreading,
diaphanous, mantlelike; synandrodes (sterile flowers)
between male and female flowers usually irregular or
fungiform, not prismatic 66. Chlorospatha
101. Spathe tube always convolute; stylar region as broad as
ovary (Caladium paradoxum has discoid, coherent stylar
regions); synandrodes between male and female flowers
well developed, prismatic; placentas 1-2(-3), parietal;
seeds several (rarely 1-2) 63. Caladium
Spathe tube gaping widely at anthesis; style much nar-
rower than ovary; synandrodes (sterile flowers) lacking,
male and female zones contiguous; placenta 1, basal; seed
solitary 62. Scaphispatha
102. Ovules more than 1; leaf blade pel tate 103
- Ovule solitary; leaf blade not peltate 68. Hapaline
103. Female flowers with staminodes; spathe not constricted;
stem trunk-like or creeping 100. Steudnera
- Female flowers without staminodes; spathe with 1 or 2
constrictions; stem tuberous, producing erect or spread-
ing stolons bearing small tubercles covered in hooked
scales 101. Remusatia
104. Leaf blade pedatisect; female flowers each with several
large staminodes 99. Protarum
- Leaf blade entire or pinnatifid; female flowers without
staminodes (except single small ones in Colocasia
esculenta) 105
105. Placentas parietal; ovules many; leaf blade always entire
102. Colocasia
- Placenta basal; ovules few; leaf blade entire or pinnatifid
103. Alocasia
I. Subfam. Gymnostachydoideae Bogner &
Nicolson (1991)
Laticifers absent; stem a short rhizome; leaves dis-
tichous, linear, not differentiated into blade and
petiole, venation parallel; flowering shoot a com-
plex synflorescence consisting of 3-7 successive,
short, perennating floral sympodia, borne on an
erect peduncular axis, each sympodium axillary
to a bract and composed of numerous inflores-
cences; spathe inconspicuous; spadix long-
stipitate; flowers bisexual, 2-merous, tepals 4,
fornicate, stamens 4, thecae dehiscing by longitu-
dinal slit; ovary 1-locular, ovule 1, atropous, pla-
centa apical; berries deep blue, exserted beyond
tepals; seed obovoid, testa absent, embryo axile,
endosperm copious, starchy apart from peripheral
layer.
44 Araceae

@
1. Gymnostachys R. Br.
Gymnostachys R. Br., Prodr.: 337 (1810}.
~~
E~
Leaves basal, midrib distinct, margins erose-
serrate; spathe short, alternating with prophylls in
each floral sympodium; spadix cylindric, erect in F
flower, pendent in fruit. 2n = 48. Only one sp., G.
anceps R. Br., Australia (Queensland, New South
Wales), in moist hardwood forests.

II. Subfam. Orontioideae Mayo, Bogner &

P.C. Boyce (1997).

1. Tribe Orontieae R. Br. ex Dumort. (1827).

Laticifers absent (except Orontium); swamp
plants, helophytes; stem a stout, erect rhizome,
continuation shoot in axil of last leaf preceding
spathe; leaf blade entire, oblong-elliptic or cor-
date; flowers bisexual, with perigone of fornicate
tepals, stamens with distinct filament, anther ter-
minal, connective inconspicuous, thecae dehisc-
ing by longitudinal slits; ovary ± immersed in
spadix axis; testa smooth, embryo large, en-
dosperm absent in Orontium, reduced to 1 cell
layer in the other genera.

2. Orontium L.
Orontium L., Sp. Pl.: 324 (1753); Huttleston, Temp. N Am.
Arac.: 50-55 (1953) (unpubl. thesis).

Laticifers present, simple; leaf blade aerial or float-

ing; primary veins arcuate-parallel; inflorescences
1-2; peduncle suppressed; spathe inconspicuous,
short, inserted at attachment to rhizome; spadix
stipe very long, swollen distally; spadix emergent,
conical, slender, golden yellow; tepals (4-)6;
stamens 6 or 4; ovary 1-locular, ovule 1,
hemianatropous, placenta basal; berry depressed-
globose, green; testa thin. 2n = 26. Only one sp., 0.
aquaticum L., temperate and warm temperate
eastern N America, partly submersed.
3. Lysichiton Schott Fig. 11A-C
Lysichiton Schott, Oesterr. Bot. Wochenbl. 7: 62 (1857); Hulten
& St. John, Sven. Bot. Tidsk. 25:453-464 (1931); Huttleston,
Temp. N. Am. Arac.: 31-38 (1953) (unpubl. thesis).
Fig. llA-F. Araceae-Orontioideae. A-C Lysichiton camts-
chatcensis. A Habit. B Inflorescence. C Flower, one tepal and
one stamen removed. D-F Symplocarpus foetidus . D Inflores-
cence. E Flower, seen from above. F Flower, longitudinal
section; note single ovule with apical placentation. (Takhtajan
1982)
spadix stipe, distally boat-shaped, gaping; spadix
cylindric, stipe very long; tepals 4; stamens 4;
gynoecium elongate-ovoid, style distinct, atten-
uate, ovary 2-locular or incompletely 2-locular,
ovules atropous, 1-2 per locule, placenta axile,
stigma capitate; seed ellipsoid or ovoid. 2n = 28.
Two spp., ·temperate E Asia and western N
America, in swamps and wet woodlands.

Robust, rosulate; petiole flattened; primary veins

arcuate-parallel to subpinnate; inflorescence 1; 4. Symplocarpus Salish. ex Nuttall Fig. 11D- F
peduncle suppressed; spathe inserted at attach- Symplocarpus Salish. ex Nuttall, Gen. 1: 105 (1818), nom.
ment to rhizome, proximally narrow and clasping cons.; Lee, Ill. Fl. Korea: 183 (1989).
 Araceae
Robust, rosulate; leaf blade subcordate or cor-
date-ovate, primary veins arcuate-parallel to
subpinnate; inflorescence 1(-2); peduncle short;
spathe thick, ventricose or conchiform, cucullate-
convolute below, open above; spadix short,
subglobose, stipe ± short; teplas 4, stamens 4;
ovary 1-locular, ovule 1, subatropous, placenta
apical-parietal, style long-attenuate; infructes-
cence ± globular, berries immersed in spongy
axis; seed globose, testa thin. 2n = 30, 60. Three
spp., temperate E Asia and eastern N America,
swamps and water courses.
III. Subfam. Pothoideae Engler (1876).
Laticifers absent; stem usually aerial; petiole
geniculate apically; higher-order leaf venation re-
ticulate; spathe simple, spreading to reflexed, not
enclosing spadix; flowers bisexual, perigoniate;
stamen filaments distinct, anthers terminal,
thecae dehiscing by longitudinal slits, connective
inconspicuous.
1. Tribe Potheae Engler (1876).
Shrubby climbers with tough woody stems; main
shoot monopodia!, flowering shoots axillary or
infra-axillary; leaves distichous; petiole sheath
long and often broad, flattened and apically au-
riculate (reduced in Pedicellarum and Pothos se-
ries Goniuri); flowers (2-)3-merous; tepals (4-)6,
unthickened and without truncate apices, stamens
6; ovules 1 per locule, anatropous, stigma sessile,
usually umbonate; embryo large, endosperm
absent.
5. Pothos L.
Pothos L., Sp. Pl.: 968 (1753).
Lower branches of stem rooting, the upper
ones free; petiole sheath broad, flattened and
auriculate apically or much narrower and ±
involute, leaf blade linear- to ovate-lanceolate;
flowering articles axillary or infra-axillary,
usually with basal cataphylls; peduncle some-
times reflexed; spathe ovate to linear; spadix
globose, ellipsoid or narrowly cylindric, sessile
to long-stipitate; tepals ± fornicate; ovary
3-locular, placenta axile; berry ellipsoid or abo-
void, red, 1-3-seeded; seed compressed -ellipsoid,
testa thickish, smooth. 2n = 24, 36. About 80
spp., Madagascar and Comores Is., tropical S
and SE Asia, Indonesia, Borneo, Philippines,
45
New Guinea, NE Australia; in tropical humid
forests.
6. Pedicellarum M. Hotta
Pedicellarum M. Hotta, Acta Phytotaxon. Geobot. 27: 61
(1976); Nicolson, Aroideana 7: 56-57 (1984).
Petiole sheath narrow, ± involute; leaf blade
lanceolate to elliptic, venation pinnate, forming
submarginal collecting veins; flowering articles
infra-axillary, with cataphylls basally; spathe
ovate; spadix axis flexuose; flowers sparse, distant,
on short pedicels, swollen below perigone into a
receptacle-like structure, tepals connate; thecae
latrorse; ovary 3-locular, placenta basal; berry
obovoid, red, 1-3(?)-seeded; seed compressed-
ellipsoid. n = ? Only one sp., P. paiei M. Hotta,
Borneo, in humid forests.
7. Pothoidium Schott
Pothoidium Schott, Aroideae 26 (1856-57); Oesterr. Bot.
Wochenbl. 7: 70 (1857).
Petiole broad, flattened, apically rounded; leaf
blade triangular-lanceolate, venation ± arcuate-
parallel; flowering shoot terminating in a branch-
ing system of spadices; spathe lanceolate, short,
or lacking entirely in the most apical inflores-
cences; spadix cylindric, stipitate; ovary 1-locular,
placenta parietal-basal; berry ovoid, acute;
seed oblong, testa smooth. 2n = 24. Only one
sp., P. lobbianum Schott, Philippines, Celebes,
Moluccas, Taiwan; in humid forests.
2. Tribe Anthurieae Engler (1876).
Sympodial articles composed of prophyll, cata-
phyll, foliage leaf, inflorescence (often aborted);
mature petiole with very short sheath, geniculate
apically and basally; primary lateral veins pinnate
(rarely arcuate-parallel), usually forming sub-
marginal collective veins; spathe usually simple,
spreading, reflexed or erect; spadix usually
tapering to subcylindric; flowers bisexual, with
perigone of 4 fornicate tepals, stamens 4, filaments
distinct, anther terminal, connective inconspicu-
ous; ovary 2-locular, locules 1-2-ovulate (rarely
more), ovules anatropous to campylotropous,
placenta axile-subapical; mature berry exserted
and usually dangling; seed often sticky, endo-
sperm copious.
46
8. Anthurium Schott
Anthurium Schott, Wiener Z. Kunst 1829 (3): 828 {1829);
Madison, Selbyana 2: 239-282 {1978); Croat & Bunting,
Aroideana 2: 15-25 {1979); Mayo, Kew Bull. 36: 691-719
{1982); Croat & Sheffer, Aroideana 6: 85-123 {1983); Croat,
Ann. Mo. Bot. Gard. 70: 211-420 {1984) & Monogr. Mo. Bot.
Gard. 14: 1-204 {1986) & Ann. Mo. Bot. Gard. 78: 539-855.
Stem creeping to erect, sometimes root-climbing;
leaf blade entire and linear to cordate, hastate or
sagittate, trilobed or trisect, pedately or radiately
lobed or -sect (primary lobes rarely themselves
pinnately lobed); inflorescence 1; peduncle rela-
tively long, more rarely short; spathe usually per-
sistent, :::t::: rarely brightly coloured; spadix sessile
to long-stipitate, sometimes clavate, rarely glo-
bose; mature berry globose to elongate-fusiform,
variously and often brightly coloured, 2-4-seeded
(rarely more); seed usually :::t::: oblong, testa :::t:::
verruculose, embryo axile, subcylindric. 2n = 30,
60, 90 (20, 24, 28, 40, 48, 56, 84). About 800 spp.
(probably more according toT. Croat), Mexico to
Argentina, from humid to dry tropical lowland
to lower montane forest, normally epiphytic or
climbing hemiepiphytic, also terrestrial, litho-
phytic or rarely rheophytic; A. scherzerianum
Schott and A. andraeanum Linden important in
horticulture; 19 sections recognised (Croat and
Sheffer 1983).
IV. Subfam. Monsteroideae Engler (1876).
Laticifers absent; trichosclereids (H- or T-shaped)
abundant (except Anadendreae, Heteropsideae;
in Amydrium present in inflorescence and very
sparse elsewhere); terrestrial, climbers, hemi-
epiphytes or more rarely epiphytes; petiole
geniculate apically, sheath usually long; leaf blade
never sagittate, sometimes pinnatifid or perfo-
rated; spathe not constricted centrally, expanded
or boat-shaped; spadix fertile to apex; flowers
bisexual, perigone present or absent, 2-merous
(2-3-merous in Spathiphylleae).
1. Tribe Spathiphylleae Engler (1887).
Tissues with small trichosclereids arranged in
bundles; stem short, upright; leaves (sub- )disti-
chous, petiole geniculate apically; leaf blade ob-
long, cuspidate-acuminate, primary leaf venation
pinnate, finer veins parallel to them; inflorescence
solitary; flowers bisexual, 2-3-merous, tepals free
or connate; thecae dehiscing by longitudinal slit;
ovules anatropous to hemianatropous; seed :::t:::
oblong, endosperm copious.
Araceae
9. Spathiphyllum Schott
Spathiphyllum Schott in Schott & Endlicher, Melet. Bot.: 22
{1832); Bunting, Mem. N.Y. Bot. Gard. 10: 1-53 {1960), rev.;
Nicolson, Blumea 16: 119-121 {1968) (rev. Old World spp.).
Peduncle subequal to or longer than leaf; spathe
cuspidate, usually spreading, rarely enclosing spa-
dix, persistent; spadix sessile or stipitate, erect,
shorter than spathe; flowers (2- )3-merous; tepals
4-6, fornicate at apex or if connate, then forming a
truncate cup; stamens as many as tepals; ovary
(2- )3(4)-locular, ovules 2-8 per locule, placenta
axile, style comic or inconspicuous; berry rounded
or :::t::: attenuate apically, 1-8-seeded; embryo axile,
narrowly cylindric. 2n = 30, 60. Forty one spp.,
tropical America, Melanesia, E Malesia, Pacific,
in damp and humid places; Bunting (1960) re-
cognised four sections; S. solomonense Nicolson is
very distinct and recognised as Sect. Chlaeno-
phyllum by Nicolson (1994).
10. Holochlamys Engler
Holochlamys Engler in Beccari, Malesia 1: 265 {1883).
Peduncle shorter than petiole; spathe tightly
clasping the spadix, not spreading, drying after
an thesis and long persistent, gradually decompos-
ing; spadix sessile; tepals 4, connate into a trun-
cate cup; stamens 4; ovary 1-locular, ovules many,
anatropous, placenta basal; berry few- to 1-
seeded. 2n = 60. Only one sp., H. beccarii (Engler)
Engler, Melanesia (New Britain, New Guinea), on
forest floor.
2. Tribe Anadendreae Bogner & French (1984).
Trichosclereids absent; climbers; petiole geni-
culate apically, sheathed almost to apex; finer
venation reticulate; peduncle relatively long;
spathe boat-shaped, longer than spadix; perigone
membranaceous, urceolate, shorter than gyno-
ecium; stamens 4, thecae dehiscing by longi-
tudinal slit; gynoecium truncate apically, ovary
1-locular, ovule 1, hemianatropous, placenta
basal, stigma transversely oblong or subspheroid;
endosperm absent, embryo large.
11. Anadendrum Schott
Anadendrum Schott, Bonplandia 5: 45 (1857).
Leaf blade obliquely ovate-oblong, entire; spathe
oblong-ovate, white; spadix cylindric, stipitate;
berry subglobose, truncate apically; seed sub-
 Araceae
globose, testa thickish. 2n = 60. About seven spp.,
SE Asia, Indonesia, Borneo, hemiepiphytes.
3. Tribe Heteropsideae Engler (1905).
Trichosclereids absent; bushy root climbers or
hemiepiphytes; main shoot monopodia!, flower-
ing articles usually short, axillary; petiole usually
almost entirely adnate to succeeding internode,
leaving only apical geniculum free; primary and
secondary leaf venation parallel-pinnate, veins
fused distally into submarginal collective vein,
finer venation ± reticulate; spathe convolute,
opening at anthesis; perigone absent; thecae
dehiscing by apical slit; gynoecium truncate,
ovary incompletely 2-locular, ovules 2 per locule,
anatropous, axile at base of partial septum;
stigma oblong or rounded; endosperm absent.
12. Heteropsis Kunth
Heteropsis Kunth, Enum. pl. 3: 59 (1841).
Inflorescence 1, axillary or terminal on short pe-
duncle; spathe ovate or ovate-oblong; spadix stipi-
tate, shorter than spathe, cylindric or ellipsoid;
berry shortly obovoid or obpyramidal, 1-4-
seeded; seed obovoid. 2n = 28. About 13 spp.,
Costa Rica to S Brazil.
4. Tribe Monstereae Engler (1876).
Climbing hemiepiphytes (except Stenosperma-
tion); leaves distichous; petiole geniculate
apically, sheath long (except Amydrium), almost
equalling petiole, leaf blade normally oblique; pe-
duncle relatively short (except Stenospermation),
spathe usually erect and boat-shaped (except
Amydrium); spadix equalling or shorter than
spathe; perigone absent, thecae dehiscing by
longitudinal slit not reaching base.
13. Amydrium Schott
Amydrium Schott, Ann. Mus. Bot. Lugd.-Batav. 1: 127 (1863);
Nicolson, Blumea 16: 123-127 (1968); Carvell, Am. J. Bot. 6
(Suppl.): 231 (1989).
Epipremnopsis Engler (1908).
Trichosclereids sparsely present (most abundant
in style according to Carvell 1989); stem some-
times creeping; petiole sheath short; leaf blade
entire or pinnately lobed, usually perforate; pri-
mary lateral veins forming obscure, submarginal
collective veins, finer venation reticulate; inflores-
47
cence usually 1; spathe conchiform to ovate, re-
flexed at anthesis; spadix sessile to long-stipitate;
ovary 1-locular, ovules 2, anatropous, placenta
near base of single, deeply intrusive septum;
berry white-green or orange; seed subglobose,
endosperm present. 2n = 60. Six spp., tropical
SE Asia and Malesia, climbing hemiepiphytes.
14. Rhaphidophora Hasskarl
Rhaphidophora Hasskarl, Flora 25, Beibl. 1: 11 (1842).
Raphidophora Schott (1858), orth. var.
Afrorhaphidophora Engler (1906).
Sometimes repent, often robust; leaf blade entire,
irregularly perforate or pinnately lobed, finer
venation subparallel-pinnate; inflorescences 1-3;
spadix sessile, lowermost flowers often female;
ovary partially 2-locular, septa deeply intrusive,
placentas basal and parietal on septa, ovules
many, anatropous; berry red, many-seeded;
seed oblong, testa thin, smooth, endosperm
copious. 2n = 60, 120 (42, 54, 56). About 100
spp., tropical Africa and from S and SE Asia
to Melanesia, Australia, and the Pacific, climbing
hemiepiphytes, rarely rheophytic (R. beccarii
Engler).
15. Epipremnum Schott
Epipremnum Schott, Bonplandia 5: 45 (1857).
Leaf blade entire, perforate or pinnately lobed;
finer venation subparallel-pinnate; ovary 1-
locular; ovules 2, more rarely 4-8, anatropous,
placenta parietal or near base of parietal partial
septa; berry 1-8-seeded; seed kidney-shaped, testa
thickish, endosperm copious. 2n = 60 (56, 84).
Seventeen spp., tropical S and SE Asia to Australia
and the Pacific, climbing hemiepiphytes.
16. Scindapsus Schott
Scindapsus Schott in Schott & Endlicher, Melet. Bot. 21 (1832).
Sometimes robust; petiole sheath rather broad, in
some spp., decomposing to form a highly irritant,
fibrous mass; leaf blade entire, fine venation
parallel-pinnate; spadix sessile; ovary 1-locular,
ovule 1, anatropous, placenta basal; berry rather
large; seed rounded, subreniform, compressed,
testa thickish, endosperm present. 2n = 60 (42, 56,
58, 64, 70, 112). Thirty spp., from tropical Sand SE
Asia through Melanesia into the Pacific, hemi-
epiphytic climbers or rupicolous.
48 Araceae

17. Monstera Adanson Fig. 12A-C

Monstera Adanson, Fam. 2: 470 (1763), nom. cons.; Madison,
Contrib. Gray Herb. 207: 1-100 (1977), rev.

Leaf blade entire or perforate and/or pinnatifid,

fine venation parallel-pinnate or ± reticulated;
inflorescences !-several; spadix sessile; ovary 2-
locular, ovules 2 per locule, anatropous, placenta
at base of septum; seed obovoid or subcordate, a
little compressed, raphe S-shaped, testa smooth,
thickish, endosperm absent, embryo large. 2n
= 60 (24, 48, 56, 58, 70). About 40 spp., trop-
ical America (Mexico to S Brazil), climbers,
hemiepiphytes in humid forest; the variegated,
juvenile leaves of M. dubia (H.B.K.) Engler &
Krause, adpressed to tree boles, frequently attract
the attention of travellers in Amazonian forests;
the pulp surrounding the seeds of M. deliciosa
Liebm. is delicious and made into ice cream
and soft drinks in Mexico; M. deliciosa is one of
the world's most ubiquitous ornamental foliage
plants; four sections are recognised.

18. Alloschemone Schott

Alloschemone Schott, Gen. Aroid.: 99 (1858); Madison,
Selbyana 1: 325-327 (1976).

Leaves distant; petiole ± terete; leaf blade pin-

nately lobed, subcordate, lobes acute, falcate, 4-6
per side, each with only 1 primary lateral vein,
finer venation parallel-pinnate; spadix sessile;
stamens free to connate; ovary 1-locular, ovule
1, amphitropous, placenta basal; fruit and seed
unknown. 2n = 84. Two spp., Amazonian Brazil,
climbers or hemiepiphytes in forest.

19. Rhodospatha Poeppig

Rhodospatha Poeppig in Poeppig & Endlicher, Nova Gen. Sp.
3: 91 (1845).
Anepsias Schott (1858).
Leaf blade entire, fine venation parallel-pinnate;
spathe sometimes rose-coloured within; spadix ±
long-stipitate, flowers small; pollen extruded in
strands; ovary 2( -3-6)-locular, ovules usually
numerous per locule, amphitropous, placentas
axile; berry small, 10-12-seeded; seeds ::±:: flat-
tened, rounded-reniform, testa hard, verruculose,
endosperm present, embryo rather large. 2n = 28,
56. About 75 spp., tropical America (Mexico to
SE Brazil), climbers, hemiepiphytes in humid
forest.
Fig. 12A-I. Araceae-Monsteroideae. A-C Monstera obliqua.
A Habit. B Spadix. C Stamen. D-1 Stenospermation spru-
ceanum. D Spadix. E Flowers seen from above. F Flower,
longitudinal section, showing two stamens, basal placenta
with anatropous ovules, and broad stylar region with small
stigma. G Ovule, note papillose lower part of funicle. H
Stamen, front view. I Stamen, side view. (Takhtajan 1982)
20. Stenospermation Schott Fig. 12D-I
Stenospermation Schott, Gen. Aroid.: t 70 (1858); Gomez, Rev.
Stenospermation (Araceae) in Central America, Masters
Thesis, St. Louis University, 114 pp. (1983).
 Araceae
Shoots usually short; leaf blade entire; fine vena-
tion parallel-pinnate; inflorescence 1, nodding or
erect; peduncle relatively long; spadix stipitate;
ovary 2-locular, ovules 4 to many per locule, ana-
tropous, placenta basal; berry obovoid, white,
rarely orange, several- to many-seeded; seed
clavate-cylindric, slender, testa thickish, smooth,
endosperm copious. 2n = 28. About 36 spp., tro-
pical America (Guatemala to SE Brazil), in humid
forest, epiphytic or on the ground.
V. Subfam. Lasioideae Engler (1876).
Laticifers absent; terrestrial or rooted aquatics;
stem tuberous or rhizomatous, usually geophytic
(except Lasia, Podolasia); petiole usually aculeate
or warty or with striking coloration, long, usually
± geniculate apically; primary lateral veins of ma-
jor leaf blade divisions pinnate to arcuate-parallel,
higher-order venation reticulate; spadix flowering
and fruiting in basipetal sequence; flowers bi-
sexual, perigoniate (except Pycnospatha); tepals
fornicate, ± truncate, free, in 2 or more whorls;
stamens free (except Lasimorpha), anther termi-
nal; embryo large, endosperm present (except
Anaphyllum), forming a thin but distinct layer.
21. Dracontium L.
Dracontium L., Sp. Pl.: 967 (1753).
Echidnium Schott (1857).
Sometimes gigantic, tuber depressed-subglobose;
leaf usually 1; petiole sometimes geniculate api-
cally; leaf blade dracontioid, ultimate lobes with
acuminate tips, leaf dissection attained at least
partly by perforation; inflorescence 1; spathe ±
boat-shaped, persistent; spadix stipitate, much
shorter than spathe; tepals 4-8; stamens 4-6(-12);
pollen extruded in masses; ovary incompletely 1-
6-locular, locules 1(2)-ovulate, ovules anatropous
to subcampylotropous, placenta axile to subbasal,
style long, attenuate; berry obpyramidal to obo-
void, red to purplish red; seed campylotropous,
testa thick, crested and warty. 2n = 26. About 23
spp., tropical America (Mexico to Paraguay and
Bolivia), in forest and savanna, ground-dwelling,
seasonally dormant or evergreen.
22. Dracontioides Engler
Dracontioides Engler, Pfianzenreich IV.23C: 36 (1911); Mayo,
Aroideana 1: 4-10 (1978).
Dracontium sect. Urospathopsis Engler (1884).
49
Rhizome hypogeal, erect; leaves several; petiole
geniculate apically, sheath long; leaf blade stron-
gly sagittate, often perforated; inflorescence 1;
peduncle long; spathe convolute, basally forming
distinct tube, blade strongly fornicate; spadix
sessile, much shorter than spathe; tepals 4; sta-
mens 4; ovary 2-locular, locules 1-ovulate, ovules
anatropous, placenta axile, style conical; berry
obovoid, purplish red; seed campylotropous, testa
thick, crested and warty. 2n = 26. Only one sp., D.
desciscens (Schott) Engler, E Brazil, in humid
forest and wet, shaded places.
23. Anaphyllopsis A. Hay
Anaphyllopsis A. Hay, Aroideana 11: 25-31 (1989, "1988").
Rhizome hypogeal, erect; leaf 1(-2); petiole
geniculate apically; leaf blade broadly hastate-
sagittate in outline, pinnately lobed or pinnate,
often irregularly perforated, ultimate lobes often
bifidtruncate; inflorescence 1(-2); peduncle long;
spathe marcescent, convolute basally, spirally
twisted apically; spadix stipitate, much shorter
than spathe; tepals 4; stamens 4; ovary 1-locular,
ovules 1-2, anatropous, placenta basal; seed
campylotropous. 2n = 26. Three spp., northern S
America, in humid forest and wet places.
24. Pycnospatha Thorel ex Gagnep.
Pycnospatha Thorel ex Gagnep., Bull. Soc. Bot. Fr. 88: 511
(1941); Bogner, Oesterr. Bot. Z. 122: 202 (1973).
Tuber depressed-subglobose; leaves 1-2; petiole
smooth, rough or aculeate, strikingly patterned,
not geniculate apically; leaf blade dracontioid, tri-
sect; inflorescence 1, appearing before or with leaf;
peduncle short; spathe thick, ± convolute below
but margins not overlapping; spadix stipitate,
much shorter than spathe; flowers without
perigone; stamens ca. 6 or more, crowded densely
together; ovary 1-locular, ovule 1, anatropous to
hemianatropous, placenta basal to subparietal;
berry spiny, globose, dark green; seed ellipsoid,
testa thick, verrucose, embryo large, endosperm
sparse. 2n = 26. Two spp., Laos, Thailand,
Vietnam on forest floor, seasonally dormant.
25. Anaphyllum Schott
Anaphyllum Schott, Bonplandia 5: 126 (1857); Sivadasan,
Taxonomic study of Araceae of S India, unpubl. doctoral
thesis, University of Calicut, Kerala (1982).
50 Araceae
Clump- or colony-forming, rhizome creeping;
leaves few; petiole geniculate apically; leaf blade
pinnately lobed or pinnate; inflorescence 1; pe-
duncle long; spathe marcescent, either convolute
basally, spirally twisted and long-acuminate
above, or oblong-ovate and fully expanded; spadix
stipitate, rather short; tepals 4; stamens 4; ovary
1-locular, ovule 1, anatropous, placenta parietal,
intrusive; berry ovoid, red; seed ovoid, testa
unlignified, membranous, smooth, embryo large,
endosperm absent. 2n = 26. Two spp., southern
India, in evergreen forest, swamp forest
undergrowth.
26. Cyrtosperma Griffith
Cyrtosperma Griffith, Notul. Pl. Asiat. (Posthum. Pap.) 3: 149
(1851) & Icon. 3, t.169 (1851); Thompson, J. Agric. Tradit.
Bot. -Appl. 29: 185-203 (1982); Hay, Blumea 33: 427-457
(1988).
Sometimes gigantic, rhizome thick, condensed,
creeping; leaves several; petiole aculeate, geni-
culate apically; leaf blade sagittate to hastate;
inflorescence 1(-2); peduncle long; spathe erect,
upper part sometimes long-acuminate and spiral-
ling, rarely fornicate; spadix usually much shorter
than spathe; tepals 4-6; stamens 4-6; ovary 1-
locular, ovules 1-many, ovule campylotropous
to subamphitropous, placenta basal to parietal,
style short or inconspicuous; berry obovoid, 1-7-
seeded, usually red; seed reniform, orbicular or
helically twisted, crested, warty or smooth. 2n =
26. About 12 spp., tropical SE Asia to Melanesia
and the Pacific, helophytic, alongside streams,
pond margins and other wet places; the rhizomes
of C. merkusii (Hassk.) Schott are an important
subsistence food in the Pacific (Thompson 1982);
four informal groups recognised by Hay (1988).
27. Lasimorpha Schott
Lasimorpha Schott, Bonplandia 5: 127 (1857); Hay, Blumea 33:
465-468 (1988).
Sometimes gigantic, forming large colonies, rhi-
zome short, thick, hypogeal, stoloniferous; leaves
several; petiole 4-6-angled, aculeate along angles,
weakly geniculate apically, sheath short; inflores-
cence 1; peduncle long; spathe ovate, convolute
basally, persistent; spadix sessile to shortly stipi-
tate, obtuse; tepals 4-6; stamens 4-6, filaments
free or ± connate; ovary 1-locular, ovules 4-6,
campylotropous, placenta basal and intrusive-
parietal; berry ± globose, red, 1-4-seeded; seed
campylotropous, testa warty and spiny, en-
dosperm present, sparse. 2n = 26. Only one sp., L.
senegalensis Schott, Senegal to Angola, helophytic.
28. Podolasia N.E. Brown
Podolasia N.E. Brown, Gard. Chron. II, 18: 70 (1882); Hay,
Blumea 33: 463-465 (1988).
Rhizome short, erect to decumbent, petiole ac-
uleate, spines usually pointing downwards; leaf
blade deeply sagittate to hastate; inflorescence 1;
peduncle long; spathe linear-lanceolate, fully ex-
panded; spadix relatively long, usually stipitate;
tepals 4-6; stamens 4-6; ovary 1-locular, ovule 1,
anatropous, placenta parietal to basal; berry
obovoid, red; seed campylotropous, ± spherical,
testa thin, hard, smooth. 2n = 26. Only one sp., P.
stipitata N.E. Brown, SE Asia.
29. Lasia Loureiro
Lasia Loureiro, Fl. Cochinch. 64, 81 (1790); Hay, Blumea 33:
458-463 ( 1988).
Clump-forming; stem and petiole aculeate; leaf
blade pinnately to pedately lobed, rarely hastate-
sagittate; inflorescence 1; peduncle long; spathe
erect, linear-lanceolate, caducous, lowest part
gaping at anthesis, upper part spirally twisted,
thick, spongy; spadix short, sessile; tepals 4-6, sta-
mens 4-6; ovary 1-locular, ovule 1, anatropous,
placenta apical, style inconspicuous; berry green,
apex often minutely spiny; seed ± pyramidal,
campylotropous, testa thin, hard, sparsely spiny.
2n = 26. Two spp., S and SE Asia, Indonesia,
Borneo, New Guinea, aquatic or swampy sites.
30. Urospatha Schott
Urospatha Schott, Aroideae 3 (1853).
Urospathella Bunting (1988).
Rhizome hypogeal, horizontal to vertical; leaves
several; petiole smooth to scabrid-verrucose; leaf
blade sagittate to hastate, rarely narrowly lan-
ceolate; inflorescence 1; peduncle long; spathe
convolute below, above long acuminate, usually
spirally twisted; spadix sessile to shortly stipitate,
short; tepals 4-6; stamens 4-6; ovary incompletely
1-2-locular, ovules (1-)2 or more per locule,
anatropous, placenta axile or basal-parietal; seed
cat1lpylotropous, testa hard, thickish, smooth to
verrucose. 2n = 52. About ten spp., Guatemala to
SE Brazil, open aquatic habitats, swamps, along
rivers, in brackish water.
 Araceae 51

VI. Subfam. Calloideae Endl. (1837). to basal; berry and seed large, testa smooth, en-
dosperm absent.
Laticifers present, simple, articulated; rooted
aquatic; stem rhizomatous; leaves distichous;
32. Zamioculcas Schott Fig. 13
petiole sheath long; leaf blade cordate; venation
parallel-pinnate, uniformly fine; spathe fully ex- Zamioculcas Schott, Syn. Aroid.: 71 (1856); Mayo, Fl. Trop.
panded, elliptic- to ovate-lanceolate, persistent; East Africa, Araceae: 15-16 (1985).
flowers bisexual, 3-merous, perigone absent; sta-
mens 6 (or more); ovary 1-locular, ovules 6-9, Rhizome short, very thick; leaves few to several,
anatropous, placenta basal; endosperm copious. petiole thickened basally, geniculate distally; leaf
blade pinnately compound, leaflets oblong-
elliptic, thick, deciduous, capable of rooting to
31. Calla L. form new plants; inflorescences 1-2; peduncle
Calla L., Sp. Pl.: 968 (1753); Riedl in Hegi, Ill. Fl. Mitteleuropa, short; male flowers: filaments free, thecae introrse;
ed. 3, 2: 326-327 (1979). sterile flowers: with 4 tepals surrounding clavate
pistillode; female flowers: staminodes absent;
Stem green; petiole sheath with long, free apical berry 1-2-seeded, white; seed ellipsoid. 2n = 34.
ligule; inflorescence usually 1; spathe white within, Only one sp., Z. zamiifolia (Loddiges) Engler,
green without; spadix stipitate, apical flowers tropical and southern subtropical Africa (Kenya
often male; berry several-seeded, red; seed terete- to Natal); in moist forest and savannas, seasonally
oblong, testa thick. 2n = 36, 54, 72. Only one dormant or evergreen.
sp., C. palustris L., circumboreal, aquatic and
helophytic in streams, ponds, swamps and bogs. 33. Gonatopus J.D. Hooker ex Engler
Gonatopus J.D. Hooker ex Engler in A. & C. De Candolle,
VII. Subfam. Aroideae Engler (1876). Monogr. Phan. 2: 208 (1879); Mayo, Fl. Trop. East Africa,
Araceae: 9-15 (1985).
Heterolobium Peter (1930).
Laticifers present, rarely absent, usually simple, Microculcas Peter (1930).
more rarely anastomosing; stem most frequently
hypogeal, tuberous or rhizomatous, less often Rhizome or subglobose tuber; leaf 1; petiole
aerial, rarely hemiepiphytes, climbers or epi- geniculate basally or centrally; leaf blade pinnately
phytes, very rarely floating aquatics (Pistieae); compound, primary lateral pinnae pinnately to
petiole usually not geniculate apically (except tripinnately lobed; inflorescences 1-4, often
Anubiadeae, Bognera, Culcasieae, Zami.ocul- appearing before leaf; peduncle short to long;
cadeae, rarely in Philodendreae and Homalo- stamen filaments connate; female flowers:
meneae); spathe usually differentiated into lower, staminodes absent, rarely 1; berry ovoid-ellipsoid,
convolute tube and upper, gaping blade; flowers orange, red or :±: dirty white, large. 2n = 34, 68.
unisexual, perigone absent (except Stylochaet- Five spp., tropical and southern subtropical Africa
oneae, Zamioculcadeae). (Kenya to Natal), on forest floor, seasonally
dormant.
1. Tribe Zamioculcadeae Engler (1876).
2. Tribe Stylochaetoneae Schott (1856).
Laticifers absent; seasonally dormant, stem hypo-
geal; leaf blade compound; petiole geniculate; Laticifers absent; stem hypogeal, rhizomatous,
spathe convolute basally, blade reflexed at anthe- roots often very thick, fleshy; leaf primary lateral
sis; spadix subequalling spathe, female zone sepa- veins mostly arising basally, arcuate, finer vena-
rated from male by short, constricted zone of tion reticulate; spathe tube margins connate; spa-
sterile flowers, male zone cylindric to clavate, dix mostly hidden within spathe, fertile to apex;
fertile to apex; flowers unisexual with perigone of flowers with :±: urceolate, connate perigone; male
4 free, :±: prismatic tepals; male flowers: stamens 4, flower usually with central peglike pistillode, sta-
surrounding :±: clavate pistillode, filaments dis- men filaments filiform, anthers terminal, dehisc-
tinct (connate in Gonatopus), anther terminal, ing by longitudinal slit, connective slender; ovary
connective inconspicuous, pollen extruded in 1-4-locular, locules 1- to many-ovulate, ovules
strands; female flowers: ovary 2-locular, locules anatropous, placenta basal, parietal or axile; en-
1-ovulate, ovules hemianatropous, placenta axile dosperm copious.
52 Araceae
Fig. 13A-G. Araceae-Aroideae. Zamioculcas zamiifolia. A
Habit; note geniculate leaves. B Inflorescence. C Staminate
flower. D Sterile flower. E Stamen. F Pistillate flower, ovary
wall removed to show locules. G Rooted leaflet with young
tuberous rhizome and shoot. (Takhtajan 1982)
34. Stylochaeton Leprieur
Stylochaeton Leprieur, Ann. Sci. Nat. Bot. II, 2: 184 (1834);
Mayo, Fl. Trop. East Africa, Araceae: 42-54 (1985); Ntepe-
Nyame, Fl. Cameroun 31: 42-46 (1988).
Leaves 1-few; leaf blade lanceolate to cordate,
hastate or sagittate; inflorescences l-4, often sub-
tended by cataphylls, borne at or below ground
level; spathe tube ±ventricose at base, rarely con-
stricted between lower and upper swollen zones
or entirely narrow-cylindric; female flowers borne
spirally or in 1 whorl, sometimes contiguous with
male zone, often with ± naked zone or sterile
flowers in between; male flower 2-7 -androus;
perigone often with fleshy margins; infructescence
borne at or below ground level, berry fleshy, 1-
few-seeded. 2n = 28, 56. Seventeen spp., tropical
and southern subtropical Africa, in humid to dry
forest, woodland, savannas, seasonally dormant
or evergreen geophytes.
3. Tribe Dieffenbachieae Engler (1876).
Internodes distinct; spadix: female zone entirely
adnate to spathe, laxly flowered, sterile male flow-
ers (synandrodes) usually present between female
and male zones; male flowers: stamens connate
into a truncate, prismatic synandrium, thecae lat-
eral, fused connectives strongly thickened; female
flowers: distant, ovary 1-3-locular, locules 1-
ovulate, ovule anatropous, placenta basal or
basal-axile.
35. Dieffenbachia Schott
Dieffenbachia Schott, Wiener Z. Kunst 1829 (3): 803 (1829);
Young, Am. J. Bot. 73: 931-944 (1986).
Maguirea A.D. Hawkes (1948).
Sometimes robust; stem erect or decumbent; leaves
several per article; petiole not geniculate, sheath
relatively long; leaf blade ovate to lanceolate, often
variegated; primary lateral veins pinnate, forming
a marginal vein, finer venation parallel-pinnate;
inflorescences (1-)2-7; peduncle usually shorter
than spathe; spathe slightly constricted, tube usu-
ally much longer than blade, persistent; spadix:
female zone usually separated from male zone by
± naked axis bearing a few scattered sterile
flowers; male flowers: 4-5-androus; female flowers:
gynoecium surrounded by 4-5 clavate staminodes;
ovary subglobose to ovoid, 1-3-locular, placenta
axile-basal or basal; berry 1-3-seeded, globose,
often scarlet; seeds globose to ovoid, testa smooth,
thickish, embryo large, endosperm absent. 2n =
34, 68. About 30 spp., Mexico to northern Argen-
tina, evergreen, terrestrial or helophytic, in humid
forests and damp sites.
36. Bognera Mayo & Nicolson
Bognera Mayo & Nicolson, Taxon 33: 690 (1984); Madison
in Aroideana 3: 101-102 (1980) (as Ulearum recondita
Madison).
 Araceae
Stem creeping, epigeal, rhizomatous; foliage leaf
1 per article, alternating with cataphylls; petiole
indistinctly geniculate at base and apex, shortly-
sheathed; leaf blade ovate-lanceolate; venation
pinnate, secondary veins arcuate, ± parallel to
primaries, finer veins distinctly reticulated; inflo-
rescence 1; peduncle shorter than petiole; spathe
boat-shaped, unconstricted; spadix: female zone
separated from male zone by ± naked zone bear-
ing a few globose sterile flowers; male flowers: 3-
4-androus; female flowers: staminodes absent,
ovary 1-locular, placenta basal; berry and seed
unknown. 2n = 34. Only one sp., B. recondita
(Madison) Mayo & Nicolson, Brazil (western
Amazonia), on forest floor.
4. Tribe Spathicarpeae Schott (1856)
Seasonally dormant, stem tuberous, rarely
rhizomatous (Gearum), hypogeal; primary lat-
eral veins of leaf or leaf lobes forming single
marginal vein (except Spathantheum and Spa-
thicarpa), finer venation reticulate; spathe usually
unconstricted; spadix fertile to apex (except
Mangonia); male flower a synandrium of ± con-
nate stamens (some partly or completely free in
Gorgonidium); gynoecium surrounded by free
staminodes or urceolate synandrode; ovary
locules !-ovulate (except Mangonia); embryo
usually relatively small, axile, endosperm copious,
rarely absent (Gearum).
37. Mangonia Schott
Mangonia Schott, Oesterr. Bot. Wochenbl. 7: 77 (1857);
Bogner, Darwiniana 18: 70-79 (1973).
Felipponia Hicken (1917), non Felipponea Brotherus (1912).
Felipponiella Hicken (1928).
Small; leaves several; leaf blade narrowly elliptic
to oblong-sagittate; inflorescence 1, appearing
before leaves, subtended by cataphylls; spathe:
tube subcylindric, blade lanceolate to oblong,
erect, gaping; spadix: female zone short, contigu-
ous with male zone or separated by short ± naked
zone, male zone more laxly flowered, terminal
sterile appendix of sterile male flowers;
synandrium 2-5-androus; ovary 2-3-locular,
ovules 2 per locule, anatropous, placenta axile;
testa smooth. 2n = ? Two spp., S Brazil, Uruguay,
in seasonally wet places.
38. Taccarum Brongn. ex Schott
Taccarum Brongn. ex Schott, Oesterr. Bot. Wochenbl. 7: 221
(1857); Bogner, Willdenowia 19: 191-198 (1989).
53
Often robust; leaf 1; leaf blade bipinnatifid to
tripinnatifid, ± dracontioid, rarely pinnatifid; in-
florescence 1(-2), subtended by cataphylls; pe-
duncle shorter than petiole; spathe boat-shaped,
tube convolute, blade gaping, marcescent and
later deciduous; spadix sometimes much longer
than spathe; female zone free, usually contiguous
with male, rarely with a few bisexual flowers in
between; thecae oblong to broadly ellipsoid; ovary
3-6(?-7)-locular, ovules anatropous, placenta
axile; berry depressed-globose, slightly furrowed,
scarlet, 3-5-seeded; seed ellipsoid, testa smooth or
granulate. 2n = 34. Five spp., tropical and sub-
tropical S America.
39. Asterostigma F.E.L. Fischer
& C.A. Meyer Fig. 14
Asterostigma F.E.L. Fischer & C.A. Meyer, Bull. Cl. Phys.-
Math. Acad. Imp. Sci. Saint Petersbourg II, 3: 148 (1845);
Bogner, Adansonia II, 9: 125-130 (1969).
Andromycia A. Rich. (1850).
Leaf usually 1; leaf blade pinnatisect or rarely
entire; inflorescences 1-3, appearing before or
with leaf; peduncle relatively long; spathe ±
unconstricted, persistent, tube narrow, blade
gaping at anthesis; spadix: female zone free or
partly adnate to spathe, laxly flowered, contig-
uous with male zone; synandrium mushroom-
shaped, umbonate; gynoecial staminodes 3-5,
free or connate, ovary 3-5-locular, ovules anatro-
pous, placenta axile-basal; berry ± depressed-
globose; testa thin, smooth. 2n 34.
About seven spp., southern tropical S America,
terrestrial.
40. Gorgonidium Schott
Gorgonidium Schott, Ann. Mus. Bot. Lugd.-Bat. 1: 282 (1864);
Bogner & Nicolson, Bot. Jahrb. 109: 529-554 (1988), rev.
Leaf 1; leaf blade pinnatifid, pinnatisect or
bipinnatifid; inflorescence 1(-2), appearing before
leaf; peduncle usually shorter than spathe;
spathe unconstricted, boat-shaped; spadix: female
zone continguous with male; male flowers: 3-
7-androus, stamens free or ± connate; central
pistillode often present and composed of 3-4
stylodia and stigmatoids (G. vargasii Bogner &
Nicolson); gynoecial staminodes 6-8, ovary
(2-)4-5( -7)-locular, ovules atropous, placenta
axile-basal; berry globose to depressed-globose;
testa rough. 2n = 34. Three spp., N Argentina,
Bolivia, Peru, 900-3000 m alt.
54
~ male by sterile zone bearing shallow, truncate
~~ c
©
' stigma subsessile, weakly 3-4-lobed; berry
l
D
Fig. 14A-F. Araceae-Aroideae. Asterostigma luschnathia-
num. A Habit. B Spadix. C, D Synandrium seen from the side
and from above. E Pistil. F Same, longitudinal section.
(Takhtajan 1982)
41. Synandrospadix Engler
Synandrospadix Engler, Bot. Jahrb. Syst. 4: 61 (1883); Crisci,
Rev. Mus. La Plata, Bot. 11:255-260 (1971), Croat & Mount,
Fl. Paraguay, Araceae: 34-37 (1988).
Lilloa Spegazzini (1897).
Leaves several; petiole sheath long; leaf blade en-
tire, cordate-sagittate; inflorescences 1-2, appear-
Araceae
ing before or with leaves; peduncle short; spathe
ovate-lanceolate, unconstricted; spadix shorter
than spathe, female zone partly adnate to spathe,
laxly flowered, separated from male by few bi-
sexual or sterile flowers; synandrium long-
stipitate, elongate-conoid, apical whorl of anthers
sometimes shortly overtopped by central
pistillode; gynoecial staminodes 3-5, ovary 3-5-
locular, ovules atropous, placenta axile-basal;
berry 3-5-seeded; seed rather large, testa thick,
succose. 2n = 34. Only one sp., S. vermitoxicus
(Griseb.) Engler, northern Argentina, Bolivia,
Paraguay, in dry thorn forest.
42. Gearum N.E. Brown
Gearum N.E. Brown, J. Bot. 20: 196 (1882); Bogner & Nicolson,
Bot. Jahrb. 109: 531 (1988); Mayo eta!., Kew Bull. 49: 785-
788 (1994).
Rhizome vertical; leaves 1-2, appearing soon
after inflorescence; petiole sheath about half the
petiole length; leaf blade pedatisect, coriaceous,
venation reticulate; inflorescence 1; peduncle
short; spathe constricted; spadix shorter than
spathe; female zone of spadix separated from
synandrodes; ovary depressed-globose, 3-4-
locular, ovules atropous, placenta axile-basal,
depressed-globose; seed broadly ellipsoid; testa
thin; embryo large, endosperm absent. 2n = 34
and ca. 68. Only one sp., G. brasiliense N.E. Brown,
C Brazil.
43. Spathantheum Schott
Spathantheum Schott, Bonplandia 7: 164 (1859); Crisci, Rev.
Mus. La Plata, Bot. 11:266-268 (1971).
Leaf 1, petiole sheath very short; leaf blade entire
to pinnately lobed, emarginate to subcordate at
base, primary lateral veins forming submarginal
collective vein; inflorescence 1-2; spathe oblong-
elliptic, unconstricted; spadix entirely adnate to
spathe or male zone free (S. intermedium Bogner),
female zone either contiguous with male or with
an intermediate zone of mixed male and female
flowers; stamens completely connate, anthers lat-
eral; gynoecial staminodes 5-8, ovary 5-8-locular,
ovules atropous, placenta axile-basal; berry 5-8-
seeded; testa smooth to slightly roughened.
2n = 34. Two spp., northern Argentina, Bolivia,
Peru.
 Araceae
44. Spathicarpa W.J. Hooker
Spathicarpa W.J. Hooker, Bot. Misc. 2: 146 (1831); Crisci, Rev.
Mus. La Plata, Bot. 11: 260-266 (1971); Uhlarz, Beitr. Bioi.
Pflanz. 57: 389-429 (1982); Croat & Mount, Fl. Paraguay,
Araceae: 29-34 (1988).
Aropsis Rojas Acosta (1918).
Small; tuber horizontal; leaves several; petiole
sheath rather long; leaf blade lanceolate to hastate,
sagittate; primary lateral veins mostly arising
basally, forming 1-2 submarginal collective veins;
inflorescence 1, appearing with leaves; peduncle
long; spathe unconstricted, persistent; spadix en-
tirely adnate to spathe, often recurved at an thesis,
laxly flowered, female flowers forming 2 outer lon-
gitudinal rows, enclosing 2 parallel rows of male
flowers; synandrium stipitate, peltate; gynoecial
staminodes 3, situated only on one side of gyno-
ecium, ovary 1-locular, ovule atropous, placenta
basal; berry ovoid; testa succose. 2n = 34. About
five spp., NE Brazil to northern Argentina, in
semiarid and humid forests.
5. Tribe Philodendreae Schott (1856)
Laticifers present, simple; resin canals present in
roots, stems, leaves and inflorescences; sclerotic
hypodermis present in roots; terrestrial, climbing
or epiphytic, stem usually epigeal, intravaginal
squamules present; petiole only rarely geniculate
apically; primary lateral veins pinnate (rarely
pedate) forming 1 marginal vein, finer venation
parallel-pinnate; spathe closing after anthesis and
persisting until fruit maturity, then deciduous at
base; flowers unisexual, perigone absent; anther
thecae lacking cell wall thickenings in endo-
thecium (except P. goeldii, P. leal-costae); en-
dosperm copious.
45. Philodendron Schott
Philodendron Schott, Wiener Z. Kunst 1829 (3): 780 (1829),
nom. et orth. cons. ("Philodendrum"); Bunting, Acta Bot.
Venezuel. 10: 289-318 (1977, "1975") & Rev. Fac. Agron.
(Maracay) 10: 209-253 (1980); Mayo & Barroso, Aroideana
2: 82-94 (1979); Mayo, Systematics of Philodendron Schott
(Araceae) with special reference to inflorescence characters,
Ph.D. thesis, Univ. Reading (1986) & Acta Bot. Brasilica 1
(2) (Sup!.): 27-40 (1989, "1988") & Bot. J. Linn. Soc. 100:
139-172 (1989) & Kew Bull. 45: 37-71 (1990) & Kew Bull.
46: 601-681 (1991), rev. subg. Mecanostigma; Grayum,
Systematic Botany Monographs val. 43 (1995); Croat, Ann.
Mo. Bot. Gar d. 84: 311-704 ( 1997).
Thaumatophyllum Schott (1859).
Sometimes gigantic; internodes usually long,
intravaginal squamules present, foliage leaves nu-
55
merous (subgen. Pteromischum) or solitary per
article; petiole rarely geniculate; leaf blade linear,
ovate, elliptic, oblong, cordate, sagittate, hastate,
trifid to almost trisect, or pinnatifid, bipinnatifid
or rarely pedatisect; inflorescences 1-11, secreting
resin at anthesis from spathe and/or spadix; pe-
duncle usually short; spathe usually constricted
between convolute tube and upper ± boat-
shaped, gaping blade; spadix sessile to stipitate,
female and male zones usually separated by
sterile zone of staminodial flowers, apical sterile
staminodial zone also rarely present; male flower
2-6-androus; ovary (2-)4-8( -47)-locular, ovules
1-50 per locule, hemianatropous to hemi-
atropous, placenta axile to basal; berry white, or-
ange or red; seed tiny to fairly large, ovoid-oblong
to ellipsoid, testa often costate; endosperm starch-
free. 2n = 28, 30, 32, 34, 36, 48 (26). Over 500 spp.,
tropical and subtropical America (Mexico to
northern Argentina); tropical humid forests,
savannas, swamps, usually root-climbers, hemi-
epiphytes or epiphytes, less often lithophytic,
terrestrial or helophytic; important in horticulture
as foliage ornamental plants, e.g. P. bipin-
natifidum Schott ex Endl., P. scandens C. Koch &
H. Sello; three subgenera and numerous sections
recognised (Mayo 1990, Croat 1997).
6. Tribe Homalomeneae (Schott) M. Hotta
(1970).
Laticifers present, simple; resin canals present in
roots, stems and leaves; sclerotic hypodermis
present in roots; terrestrial or rheophytic; stem
usually epigeal; petiole geniculum absent (very
rarely present in Homalomena); primary lateral
veins pinnate forming 1 marginal vein, higher-
order venation parallel-pinnate; spathe often
boat-shaped, constricted or unconstricted, closing
after anthesis and persistent; flowers unisexual,
perigone absent; anther thecae with cell-wall
thickenings in endothecium; endosperm copious.
46. Furtadoa M. Hotta
Furtadoa M. Hotta, Acta Phytotaxon. Geobot. 32: 142 (1981) &
Gard. Bull. Singapore 38: 43-54 (1985).
Small, stem repent; leaf blade elliptic; inflores-
cences 1-3; peduncle relatively long; spathe boat-
shaped, not constricted; spadix: female and male
zones with a zone of few sterile flowers in between;
male flowers: stamen 1, overtopped by flask-
shaped pistillode, thecae dehiscing by longitudi-
nal slit; female flowers: gynoecium with 1 truncate
56
staminode, ovary 1-locular, ovules many,
hemianatropous, placenta basal; berry light green;
seed ellipsoid, testa smooth, thin. 2n = 40. Two
spp., Sumatra and Malay Peninsula, rheophytes
on rocks in streams. The structure of the male
flower is unique.
47. Homalomena Schott
Homalomena Schott in Schott & Endlicher, Melet. Bot.: 20
(1832); Furtado, Gard. Bull. Singapore 10: 183-238 (1939);
Hotta, Acta Phytotaxon. Geobot. 33: 127-139 (1982) & Gard.
Bull. Singapore 38: 43-54 (1985); Bogner, Aroideana 8: 73-
75 (1986).
Diandriella Engler (1910).
Rarely robust; petiole rarely aculeate; leaf blade
lanceolate to sagittate, rarely peltate; inflores-
cences 1-6; peduncle short; spathe boat-shaped;
spadix stipitate or sessile, female and male zones
contiguous or with sterile male flowers or naked
zone in between; male flower ( 1-)2-4-androus,
thecae usually opening by longitudinal (more
rarely transverse) slit, pollen usually extruded
in strands; female flower usually with 1(-3)
staminodes, ovary incompletely 2-4( -5)-locular,
ovules many, hemianatropous, placentas parietal,
or parietal above and axile below, or basal; seed ±
ellipsoid, testa thick, costate. 2n = 38, 40, 42, 80.
About 110 spp., S and SE Asia to Melanesia, tropi-
cal America (Costa Rica to Bolivia), terrestrial and
rheophytic; five sections recognised.
7. Tribe Anubiadeae Engler (1876).
Roots with sclerotic hypodermis; petiole geni-
culate apically; leaf blade elliptic, lanceolate to
hastate-trifid; primary lateral veins pinnate, form-
ing 1 marginal vein, finer veins parallel-pinnate
with ± distinct transverse veins; spathe boat-
shaped, unconstricted, persistent; stamens con-
nate into ± prismatic synandrium; thecae lateral
or marginal, rarely covering whole synandrium,
dehiscing by longitudinal slit, fused connectives
thickened, ± truncate; ovary (1-)2-3-locular,
ovules many per locule, anatropous, placenta
axile; berry depressed-globose to obovoid; seed
small, ± ovoid to subcylindric, endosperm
copious.
48. Anubias Schott
Anubias Schott, Oesterr. Bot. Wochenbl. 7: 398 (1857); Crusio,
Meded. Landbouwhogesch. Wageningen 79 (14): 1-48
(1979), rev. & Aqua-Planta, Sonderheft 1: 1-14 (1987).
Amauriella Rendle (1913).
Araceae
Rhizome thick, creeping; petiole relatively long,
rarely shortly and sparsely spiny; inflorescences
1-3; peduncle fairly long; spathe closing after an-
thesis; spadix: female and male zones usually con-
tiguous, rarely with few sterile flowers in between;
male flowers: 3-8-androus; berry green to pale
green. 2n = 48, 72. Eight spp. Senegal to Angola,
helophytes, sometimes completely submerged, or
in forest swamps, rarely rheophytic.
8. Tribe Schismatoglottideae Nakai (1943).
Evergreen herbs; usually epigeal; leaves several
per article; petiole sheath usually with long,
marcescent, apical ligule; leaf blade apex with
tubular mucro (except most Schismatoglottis
spp.), primary lateral veins pinnate, forming 1
usually prominent marginal vein, finer venation
parallel-pinnate; inflorescence usually 1; spathe
tube persistent, blade usually white (except
Piptospatha), caduceus at anthesis, rarely
marcescent (Schismatoglottis texturata (Schott)
Engl., Hottarum lucens Bogner); stamens usually
free, sometimes filaments ± connate, often ±
elongated; ovary 1-locular, ovules atropous
to hemitropous (except Schismatoglottis); testa
usually costate, embryo axile, endosperm
copious.
49. Schismatoglottis Zollinger & Moritzi
Schismatoglottis Zollinger & Moritzi in Moritzi, Syst.
Verzeichniss Zollinger: 83 (1846); Bunting, Ann. Mo. Bot.
Gard. 47: 69-71 (1960); Hotta, Mem. Coli. Sci. Univ. Kyoto,
ser. B, 32: 19-30 (1965) & ibid.: 223-238 (1966); Bunting &
Steyermark, Brittonia 21: 187-190 (1969); Bogner & Hotta,
Acta Phytotaxon. Geobot. 34: 48-50 (1983).
Philonotion Schott (1857).
Stem rhizomatous or epigeal, shortly erect; petiole
sheath less than half petiole length, sometimes
ligulate; leaf blade narrowly elliptic to cordate-
sagittate, rarely with apical tubular mucro; inflo-
rescences 1-3; spadix zonation variable: female
zone sometimes bearing sterile organs at the very
base, female and male zones sometimes with
naked zone or zone of sterile flowers in between,
upper part of spadix clavate to cylindric, terminal
appendix of sterile flowers sometimes present;
male flowers: 2-3-androus, filaments usually dis-
tinct, sometimes connate basally; pollen extruded
in strands; staminodes obpyramidal to clavate,
usually truncate; female flowers: sometimes with
whorl of 1-4 staminodes, ovules (1-)few to nu-
merous, anatropous to hemianatropous, placentas
1-4, parietal, usually extending from base to
 Araceae 57

apex of locule; berry oblong to globose, green or 52. Bucephalandra Schott

dull yellow or deep red, few- to many-seeded;
seed shortly ellipsoid, straight. 2n = 26, 39, 52.
About 120 spp., SE Asia to New Guinea, northern
S America, on forest floor, terrestrial or
rheophytic.
50. Piptospatha N.E. Brown
Piptospatha N.E. Brown, Gard. Chron. II, 11: 138 (1879);
Hotta, Mem. Coil. Sci. Univ. Kyoto, ser. B, 32: 19-30 (1965).
Gamogyne N.E. Brown (1882).
Stem decumbent or erect; leaf blade narrowly
lanceolate to elliptic; inflorescence ± nodding;
spathe not constricted, stoutly ellipsoid; spadix
subsessile, sometimes with a few sterile flowers at
extreme base, female and male zones contiguous
or with short zone of sterile male flowers between,
apex sometimes with a few sterile flowers; male
flower 1-2-androus; gynoecia free or superficially
connate (coherent), ovules many, placentas 2-4,
parietal or parietal and basal; berry obovoid; seed
elongate-ellipsoid to cylindric with long micropy-
lar appendage. 2n = 26. Ten spp., Borneo, Malay
Peninsula; terrestrial or rheophytic; closely related
to Hottarum.
Bucephalandra Schott, Gen. Aroid.: t. 56 (1858); Bogner,
Aroideana 3: 134-143 (1980) & Plant. Syst. Evol. 145: 159-
164 (1984).
Microcasia Beccari ( 1879).
Stem rhizomatous, decumbent; leaf blade elliptic,
oblanceolate or obovate; spathe not constricted,
ellipsoid, tube funnel-shaped, blade gaping and
caducous; spadix shorter than spathe, with a few
sterile flowers at extreme base, female zone nar-
rower than upper parts, separated from male zone
by about 2 rows of flattened, scalelike staminodes,
male zone with 2-5 rows of flowers, apical zone
globose, ellipsoid to subcylindr~c, c?mpos~d of
truncate, obpyramidal to subcylmdnc, papillose
sterile male flowers, the uppermost ± connate;
male flower 1-androus; ovary depressed-globose,
ovules many, placenta basal; berry with numerous
seeds; seed narrow-ellipsoid, with long, curved
micropylar appendage. 2n = ca. 26. Three spp.,
Borneo, rheophytes in tropical humid forest.
53. Phymatarum M. Hotta
Phymata rum M. Hotta, Mem. Coli. Sci. Univ. Kyoto, ser. B, 32:
29 (1965); Bogner, Plant. Syst. Evol. 144: 62-66 (1984).

51. Hottarum Bogner & Nicolson Stem rhizomatous, decumbent; leaf blade nar-
rowly elliptic; spathe constricted, blade boat-
Hottarum Bogner & Nicolson, Aroideana 1: 72 (197~, :'1978");
Bogner, Plant. Syst. Evol. 142: 49-55 (1983) & tbtd., 145: shaped, gaping, caducous after anthesis; spadix
159-164 (1984); Bogner & Hotta, Adansonia IV, 5: 27-31 with extreme base usually bearing a few sterile
(1983). flowers, female zone adnate to spathe, conoid to
Microcasia sect. Truncatae M. Hotta, Mem. Coil. Sci. Univ. subcylindric, separated from male zone by cylin-
Kyoto, ser. B, Bioi. 32: 21 (1965).
dric to ellipsoid zone of sterile male flowers, male
zone very short and slightly narrower, terminal
Stem ± erect; leaf blade ± elliptic; spathe con- appendix much longer, elongate-conoid, com-
stricted or not; spadix sometimes adnate to spathe posed of sterile male flowers; male flower 1-
for two-thirds oflength (H. lucens Bogner), female androus; sterile flowers subprismatic, tuberculate,
zone sometimes with sterile flowers at base, flattened or excavated; ovary depressed-globose,
either contiguous with male, or with a few sterile ovules many, placenta basal; berry many-seeded,
male flowers in between, terminal appendix broadly obovate-oblong, greenish white; seed el-
of sterile flowers present; male flower (1- )2- lipsoid, with long micropylar appendage. 2n = 26.
3( -4)-androus, sterile male flowers composed of Three spp., Borneo, rheophytes in forest, ruderal
truncate, prismatic staminodes; female flower habitats, and disused rice paddi.
sometimes with 1 clavate staminode in basal
flowers, ovary depressed-globose, ovules 10-15,
placenta basal; berry ± globose, whitish green, 54. Aridarum Ridley
many-seeded; seed ellipsoid to elongate, with
long, micropylar appendage. 2n = 26. Six spp., Aridarum Ridley, J. Bot. 51: 201 (1913); Hotta, Mem. Coli. Sci.
Univ. Kyoto, ser. B, 32: 19 (1965); Bogner, Aroideana 2: 110-
Borneo; tropical humid forest, rheophytes on 121 (1979) & Aroideana 4: 57-63 (1981) & Blumea 28: 403-
rocks in and along streams; H. lucens is rather 405 (1983).
distinct.
Stem decumbent; leaves sometimes distichous;
leaf blade linear to elliptic; inflorescence often
58 Araceae

somewhat nodding; spathe not constricted, 56. Lagenandra Dalzell

stoutly ellipsoid, blade caducous; spadix normally
Lagenandra Dalzell, Hooker's J. Bot. Kew Gard. Misc. 4: 289
with a few sterile flowers at base, female zone cy- (1852); De Wit, Meded. Landbouwhogesch. Wageningen
lindric, separated from male by zone of sterile 78: 5-45 (1978), rev. & Aquariumplanten: 286-307 (1983);
flowers, short terminal appendix of sterile flowers Bogner & Jacobsen, Aqua-Planta 12: 43-50 {1987);
present; male flower 1-2-androus, thecae apically Nicolson, Fl. Ceylon 6: 75-85 (1988).
narrowed into long or short horn, dehiscing by
apical pore; sterile male flowers of truncate Leaf blade involute in bud, linear to ovate; pe-
staminodes often with small central cavity; ovary duncle usually short; spathe thick-walled, margins
laterally compressed, ovules many, placenta basal; free above kettle, mouth of spathe tube forming a
berry globose or ellipsoid to cylindric; seed ellip- narrow aperture, blade straight or twisted, open-
soid, elongate. 2n = 24. Seven spp., Borneo, ing by narrow slit or widely gaping; spadix: female
rheophytes in humid forest. zone subglobose, separated by slender naked axis
from male zone, appendix small, conoid; gynoecia
free, in spirals or rarely whorled, ovules 1-
55. Heteroaridarum M. Hotta 12( -15), placenta basal, style absent or distinct,
Heteroaridarum M. Hotta, Acta Phytotaxon. Geobot. 27: 63 stigma broad; infructescence usually globose, ber-
(1976). ries free, dehiscent basally with pericarp becoming
revolute; testa costate. 2n = 36, 72. Fourteen spp.,
Stem short; leaf blade elliptic, coriaceous; inflores- Indian subcontinent, Sri Lanka; freshwater aquat-
cence 1; spathe slightly constricted; spadix: ics, usually helophytes, rarely rheophytes.
female zone separated from male by short zone
of sterile flowers, male zone much longer; male 57. Cryptocoryne Fischer ex Wydler
flowers: 3-androus, 2 outermost stamens larger
with shortly horned thecae, central stamen Cryptocoryne Fischer ex Wydler, Linnaea 5: 428 (1830);
smaller with hornless thecae, all thecae opening by De Wit, Aquariumplanten: 141-261 (1983); Jacobsen,
Cryptocorynen, 112pp. (1982) & Nord. J. Bot. 5: 31-50
apical pore; sterile male flowers composed of (1985); Jacobsen & Bogner, Aqua-Planta 12: 96-103 (1987);
basally connate, truncate staminodes; ovary Jacobsen et al., Aqua-Planta 14: 127-132 (1989); Nicolson,
depressed-globose, ovules many, placentas 2, Fl. Ceylon 6: 85-99 (1988).
basal and apical, apical one bearing smaller and
apparently sterile ovules; berry and seed un- Leaf blade convolute in bud, linear to cordate; pe-
known. 2n = ? Only one sp., H. borneense M. duncle short, elongating in fruit; spathe margins
Hotta, Borneo, rheophyte. connate above kettle (except C. spiralis, C.
cognata), blade cordate to lanceolate, variously
coloured, ± spreading or twisted, often with a dis-
9. Tribe Cryptocoryneae Blume (1836).
tinct collar at mouth of spathe tube; spadix: fe-
Aquatic, evergreen, rhizomatous; leaves several; male zone a single whorl of 4-8 gynoecia, usually
primary lateral veins ascending, fusing distally separated from male zone by slender naked axis,
into an inconspicuous marginal vein, finer vena- appendix short, conoid to subclavate; ovaries
tion reticulate; inflorescence 1; spathe tube mar- connate, ovules 5-many, placenta subbasal to ob-
gins basally or entirely connate, usually slightly liquely parietal (morphologically basal); berries
swollen basally forming a "kettle", blade lan- connate forming ovoid syncarp which dehisces
ceolate to cordate, often twisted or long-spiralled apically as one unit to become star-shaped; seed
or long-caudate; spadix entirely enclosed in ellipsoid-oblong, testa rough to subcostate, em-
kettle of spathe tube, sterile terminal appendix bryo well developed. 2n = 20, 22, 28, 30, 33, 34, 36,
apically adnate to spathe at apex of kettle, female 42, 54, 66, 68, 72, 85, 88, 90, 102, 132. About 50 spp.,
zone with a whorl of pistillodial olfactory bodies S and SE Asia to New Guinea, submerged or emer-
situated above the fertile gynoecia (except some gent, freshwater tidal zone, forest streams and
Lagenandra spp.); male flowers 1(-2)-androus, pools; rarely seasonally dormant; Jacobsen (1982)
stamens free, anther sessile or with short filament, recognised 26 informal subgroups.
apex excavated with prominent margins, thecae
ending in prominent horn, dehiscing by apical
10. Tribe Zomicarpeae Schott (1856).
pore; pollen exine smooth; ovary 1-locular,
ovules atropous; embryo elongate, endosperm Laticifers anastomosing; small, evergreen or
copious. seasonally dormant; leaf blade cordate-sagittate
 Araceae 59

(except Zomicarpa); primary veins radiating
from base, finer venation reticulate; spadix longer
than spathe (except Zomicarpa and 1 sp. of
Zomicarpella), sterile terminal appendix present,
female spadix zone adnate to spathe; stamens free;
gynoecia few, ovules anatropous, placenta basal,
style short, stigma discoid.
58. Zomicarpa Schott
Zomicarpa Schott, Syn. Aroid.: 33 (1856); Bogner, Aroideana
3: 10-11 (1980}.
Stem tuberous; leaves few; leaf blade trisect to
pedatisect, leaflets with distinct submarginal
collective veins; inflorescences 3-5, subtended
by cataphylls, appearing before or with leaves;
peduncle subequal to leaf or longer; spathe ±
constricted, tube ± gaping at anthesis, blade
expanded at anthesis, sometimes fornicate, ±
lanceolate; spadix subequal to spathe, female
zone laxly flowered, contiguous with male,
appendix smooth or staminodial; male flower
1-2-androus; ovules 6-9, stigma broad, sub-
sessile; berry depressed-globose, whitish, few-
seeded; seed ovoid to oblong-ellipsoid with
large, strophiolate funicle, testa smooth, thin,
transparent, embryo elongate, partly green,
endosperm copious, green and starchy. 2n =
20. Three spp., NE Brazil, geophytes, seasonally
dormant.
Rhizomatous; leaves few; leaf blade usually varie-
gated; inflorescence 1; peduncle subequalling or
longer than leaves; spathe unconstricted, fully
expanded, margins reflexed at anthesis; spadix:
female zone separated from male by sterile zone
which is basally either naked or with a few scat-
tered sterile flowers and apically bears a short
dense zone of staminodes, male zone short,
few-flowered, sterile terminal appendix present,
staminodial at base, otherwise smooth, sub-
cylindric; male flowers 2-3-androus; gynoecium
slender, ovule 1; testa thin, embryo relatively
large, undifferentiated, endosperm absent. 2n =
14. Only one sp., U. sagittatum Engler, S America
(western Amazonia), on forest floor.
61. Filarum Nicolson
Filarum Nicolson, Brittonia 18: 348 (1967}.
Tuberous; leaves few; inflorescence 1-3, ap-
pearing with leaves; peduncle slender; spathe
oblong-lanceolate, unconstricted, fully expanded,
persistent; spadix very slender, male zone elon-
gate, sparsely flowered, a few sterile rudimentary
flowers above and below male zone, appendix
slender, long; male flowers 1-androus; ovary
oblong, ovule 1; berry subglobose; testa thin,
smooth, embryo large, endosperm absent. 2n = 28
Only one sp., F. manserichense Nicolson, Amazo-
nian Peru, geophyte.

59. Zomicarpella N.E. Brown 11. Tribe Caladieae Schott (1832).

Laticifers anastomosing; terrestrial, geophytic,
Zomicarpella N.E. Brown, Gard. Chron. II, 16: 266 (1881}; ·
Bogner, Aroideana 1: 72 (1979, "1978"}. rarely aquatic (]asarum), climber or hemi-
epiphyte (Syngonium); stem tuberous, rhizoma-
Stem rhizomatous; leaves 1-few; leaf blade usually tous or aerial; leaf basal ribs well developed,
variegated; inflorescences 1-2, appearing with venation pinnate, forming submarginal collective
leaf; peduncle equalling petiole; spathe oblong- vein, 1 or more marginal veins also usually
lanceolate, completely expanded or lower part present, finer venation reticulate; spadix: female
partially convolute, not or slightly constricted; zone often separated from male by subconical
spadix longer than or subequal to spathe, female to attenuate zone of synandrodes, male zone
zone contiguous with male or with short sterile subcylindric to subclavate, sterile appendix absent
zone in between, appendix slender, long; male (except Hapaline appendiculata); flowers uni-
flowers 1-androus; pollen extruded in strands; sexual, perigone absent; stamens connate forming
ovary ellipsoid-oblong, ovules 1-6; berry white, 1- synandrium; pollen extruded in strands (except
few-seeded; testa thin, smooth, embryo small, en- Scaphispatha); ovules anatropous to hemi-
dosperm copious. 2n = 26. Two spp., northern S anatropous; endosperm copious (except ]asarum,
America (western Amazonia), geophytes, season- Syngonium, Hapaline).
ally dormant.
62. Scaphispatha Brongn. ex Schott
60. Ulearum Engler
Scaphispatha Brongn. ex Schott, Prodr. Syst. Aroid.: 214
Ulearum Engler, Bot. Jahrb. 37: 95 (1905}. (1860}; Bogner, Aroideana 3: 4-12 (1980).
60 Araceae
Stem a subglobose tuber; leaves 1-2; leaf blade
peltate, ovate-subcordate to -subsagittate; inflo-
rescence 1, appearing before leaves; peduncle
long, slender; spathe ovate, fully expanded at an-
thesis, lower part later convolute and persistent,
upper part marcescent; spadix shorter than
spathe, male and female zones contiguous;
synandrium 4-androus; ovary 1-locular, ovules
3-5, placenta basal; berry subglobose to obovoid,
1-seeded; seed subglobose, testa smooth and
thin. 2n = 28. Only one sp. S. gracilis Brongn.
ex Schott, Bolivia, Brazil, geophyte, seasonally
dormant.
63. Caladium Vent.
Caladium Vent., Descript. Plant. Nouv. Jard. Cels: 30 (1801);
Madison, Phytologia 35: 103-107 (1976) & Selbyana 5: 342-
377 (1981), rev.; Bogner, Aroideana 7: 4-5 (1984); Grayum,
Ann. Mo. Bot. Gard. 73: 462-474 (1986); Mayo & Bogner,
Willdenowia 18: 231-242 (1988).
Aphyllarum S. Moore (1895).
Stem a subglobose tuber; leaf blade usually
peltate, often variegated, cordate-sagittate or sag-
ittate, rarely trisect; inflorescences 1-2, appearing
with or before leaves; peduncle subequal to peti-
ole; spathe tube usually ventricose-globose; spadix
subequal to spathe; synandrium 3-5-androus,
margins sinuous; ovary 1(-3)-locular, ovules l-ea.
20, placentas 1-3, subbasal to intrusive-parietal;
berry white, 1- to many-seeded; seed ovoid to
ellipsoid, testa longitudinally costate. 2n = 22,
26, 28, 30, 32. About 12 spp., tropical America,
geophytes, seasonally dormant or evergreen;
C. bicolor (Ait.) Vent. and its many cultivars are
horticulturally important worldwide and often
naturalised in tropical regions.
64. ]asarum G.S. Bunting
]asarum G.S. Bunting, Acta Bot. Venez. 10:264 (1977, "1975");
Bogner, Aroideana 8: 55-63 (1985).
Stem hypogeal, rhizomatous, erect; leaves several;
leaf blade linear, ::!:: bullate; peduncle long;
spathe borne above water level, blade caudate;
female zone of spadix basally adnate to spa-
the; synandrium 3-4-androus; ovary 1-locular,
( 1-)2-3( -4)-ovulate, ovule anatropous, placenta
basal; berry 1(-2)-seeded; seed large, obovoid,
testa smooth, embryo large, plumule well
developed, endosperm very sparse. 2n = 22. Only
one sp., f. steyermarkii G.S. Bunting, Venezuela,
Guyana, submerged aquatic in blackwater
streams.
65. Xanthosoma Schott Fig. 15B-K
Xanthosoma Schott in Schott & Endlicher, Melet. Bot.: 19
(1832); Madison, Phytologia 35: 103-107 (1976) & Selbyana
5: 342-377 (1981); Grayum, Ann. Mo. Bot. Gard. 73: 462-
474; Mayo & Bogner, Willdenowia 18: 231-242 (1988).
Sometimes gigantic and arborescent; tuberous;
leaf blade cordate-sagittate, hastate, tri- or
pedatisect, rarely linear-lanceolate or peltate;
inflorescences 1-several, always appearing with
leaves; peduncle usually rather short; spathe
tube usually ventricose; synandrium 4-6-androus;
pollen shed in tetrads; ovary incompletely ( 1-)2-
4-locular, ovules ca. 20-many, placentas (1- )2-4,
intrusive-parietal; berry many-seeded; seed ovoid,
testa costate. 2n = 26, 39, 52. About 57 spp.,
Mexico to northern Argentina, on forest floor
or seasonally flooded sites, sometimes weedy,
evergreen or seasonally dormant. The nutritious
tubers of the X. sagittifolium complex are an
important subsistence food throughout the humid
tropics.
66. Chlorospatha Engler
Chlorospatha Engler, Gartenflora 27: 97 (1878); Madison,
Selbyana 5: 342-377 (1981), rev.; Bogner, Aroideana 8: 48-
54 (1985); Grayum, Ann. Mo. Bot. Gard. 73:462-474 (1986);
Mayo & Bogner, Willdenowia 18: 231-242 (1988); Grayum,
Novon 1: 12-14 (1991).
Caladiopsis Engler (1905).
Stem caulescent, decumbent to erect; leaves 1-few;
petiole sheath relatively long; leaf blade cordate,
sagittate, hastate, trisect or pedatisect; inflores-
cences 3-8; peduncle slender, supported by peti-
ole sheath; spathe tube narrow, elongate; female
zone of spadix free or adnate to spathe; female
and sterile flowers often distant; synandrium 3-5-
androus; pollen shed in tetrads; ovary incom-
pletely (1- )2-4-locular, ovules few to several,
placenta intrusive-parietal, rarely basal; berry ::!::
3-5-furrowed, white to yellowish; seed minute,
costate. 2n = 26. About 16 spp., Costa Rica to
Peru, on forest floor, evergreen.
67. Syngonium Schott
Syngonium Schott, Wiener Z. Kunst 1829 (3): 780 (1829);
Croat, Ann. Mo. Bot. Gard. 68: 565-651 (1982), rev.
Porphyrospatha Engler (1879).
Climbers or hemiepiphytes with elongated inter-
nodes; leaf blade sagittate, trisect, pedatisect,
rarely pinnatifid; inflorescences several, pendent
in fruit; peduncle short; spadix much shorter than
 Araceae
.;
.
. berry and seed large, testa thin, endosperm
., .\~<r
K
Fig. 15A-K. Araceae-Aroideae. A Colocasia esculenta, habit.
B-F Xanthosoma sagittifolium. B Inflorescence, spathe partly
removed. C Group of synandria. D Synandrium, longitudinal
section. E Pistil, side view. F Same, longitudinal section. G-L
X. platylobum. G Habit. H Spadix. I Synandrium, side view.
J Pistillate flowers, seen from above. K Pistil, side view.
(Takhtajan 1982)
spathe; synandrium 3-4-androus; gynoecia partly
to completely connate, eventually forming syn-
carp, ovary (1-)2(-3)-locular, ovules 1(-2) per
locule, anatropous, placenta axile-basal; berries
61
connate, syncarp stoutly ovoid, usually brown;
seed rather large, testa smooth, thin, embryo
large, endosperm absent. 2n = 28 (24, 26). Thirty
five spp., Mexico to SE Brazil, sometimes natu-
ralised elsewhere in the tropics; some species, e.g.
S. podophyllum Schott, are important foliage
ornamental plants in horticulture.
68. Hapaline Schott
Hapaline Schott, Gen. Aroid.: t. 44 ( 1858), nom. cons.; Bogner,
Plant. Syst. Evol. 144: 59-61 (1984); Boyce, Kew Bull. 51: 63-
82 (1996), rev.
Stem tuberous; leaf 1-few; leaf blade cordate to
sagittate; inflorescence appearing with leaves;
peduncle slender, longer than petiole; spathe not
distinctly constricted, tube short, blade linear-
lanceolate; spadix subequal or longer than spathe,
female zone of2-3 flowers, adnate to spathe, sepa-
rated from male by short zone of sterile flowers,
terminal appendix filiform (when present);
synandrium 3-androus, peltate, thecae almost
pendent from margin; ovary 1-locular, ovule 1,
anatropous, placenta parietal-subbasal; berry
white, 1-seeded; testa smooth, embryo large,
endosperm absent. 2n = 26, 28. Six spp., Borneo,
geophytes, seasonally dormant.
12. Tribe Nephthytideae Engler (1887)
Petiole geniculum present; leaf essentially tripar-
tite, main venation pinnate, fine venation reticu-
late; spathe boat-shaped or fully expanded, not
clearly differentiated into tube and blade; spadix
cylindric, female and male zones usually conti-
guous, sterile flowers ± absent (except Pseudo-
hydrosme); stamen filaments lacking or very short
(except Nephthytis); ovules 1 per locule, anatro-
pous, placenta basal (except Pseudohydrosme);
absent. Chromosomes large to very large .
69. Nephthytis Schott
Nephthytis Schott, Oesterr. Bot. Wochenbl. 7: 406 (1857);
Bogner, Aroideana 3: 75-85 (1980); Knecht, Contrib. Arac.
Cote d'Ivoire: 163-171 (1983); Namur & Bogner, Adansonia
IV, 16:71-74 (1994); Ntepe-Nyame, Fl. Cameroun 31:46- 56
(1988); Hay eta!., Novon 4: 365-368 (1994).
Rhizome creeping, usually subepigeal, rarely hy-
pogeal; leaves usually several; leaf blade cordate-
sagittate, deeply sagittate or tripartite, primary
lateral veins ascending, joining into a submarginal
collective vein; inflorescence 1; peduncle relatively
62 Araceae
long; spathe fully expanded, ovate-oblong-elliptic,
often reflexed at anthesis; spadix stipitate; male
flowers 2-4-androus; ovary 1-locular; berry sub-
globose to ellipsoid, orange; seed obovoid or
subglobose, testa absent at maturity. 2n = 40, 60.
Ten spp., W and WC Africa (Liberia to Gabon),
one sp. in Borneo.
70. Anchomanes Schott
Anchomanes Schott, Oesterr. Bot. Wochenbl. 3: 314 (1853);
Knecht, Contrib. Arac. Cote d'Ivoire: 145-151 (1983); Mayo
& Bogner, Fl. Trop. East Africa, Araceae: 24-28 (1985);
Ntepe-Nyame, Fl. Cameroun 31: 22-26 (1988).
Robust to gigantic; rhizome hypogeal, erect or
horizontal; leaf 1; petiole very long, aculeate
(except A. boehmii), geniculate at main blade
divisions; leaf blade dracontioid, ultimate lobes
trapezoid, truncate or shallowly bifid; inflores-
cence 1; peduncle long; spathe boat-shaped; spa-
dix short or subequal to spathe; ovary 1-locular,
style sometimes deflexed; berry oblong-ellipsoid,
red, purplish or whitish; seed obovoid to oblong-
ovoid. 2n = 40. Four to five spp., tropical Africa
(Gambia to Zambia); tropical moist forests, savan-
nas, swamps, seasonally dormant.
71. Pseudohydrosme Engler
Pseudohydrosme Engler, Bot. Jahrb. 15: 455 (1892, "1893");
Bogner, Aroideana 4: 31-37 (1981).
Zyganthera N.E. Brown (1901).
Robust; tuber subglobose, hypogeal; leaf 1; petiole
and blade as for Anchomanes; inflorescence 1; pe-
duncle very short; spathe ovate-elliptic-oblong, ±
boat-shaped, convolute basally into obconic tube,
upper part very broad, fornicate with auriculate
margins; spadix short, sessile, male zone with
apical appendix in P. buettneri Engler; stamens
2-5; staminodes of appendix (when present)
subprismatic, free; ovary 2(-3)-locular, placenta
axile-basal, style short; berry and seed unknown.
2n = ca. 40. Two spp., Central-W Africa (Gabon),
seasonally dormant.
13. Tribe Aglaonemateae Engler (1876).
Leaf blade (linear-)ovate to elliptic-oblong, base
sometimes subcordate; primary veins pinnate,
finer veins parallel-pinnate; spathe boat-shaped,
unconstricted; spadix: female and male zones con-
tiguous; thecae dehiscing by apical pores; ovary 1-
locular, ovule 1, anatropous, style short, narrower
than ovary, stigma broad, discoid, concave; berry
large; seed large, ellipsoid, testa smooth, thin,
endosperm absent.
72. Aglaonema Schott
Aglaonema Schott, Wiener Z. Kunst 1829 (3): 892 (1829);
Nicolson, Smithson. Contrib. Bot. 1: 1-69 (1969), rev.
Stem creeping to erect, internodes green, smooth,
distinct; leaf blade often variegated; inflorescences
1-9; peduncle shorter or longer than petiole,
deflexing in fruit; spadix cylindric to clavate,
subsessile to stipitate; female zone ± few-
flowered, male zone longer; male flower 1-
androus, filament ± distinct; ovary subglobose,
placenta basal; berry red at maturity. 2n = 40,
60, 80, 100, 120 (70, 110). Twenty one spp., S and
SE Asia to New Guinea. Important ornamental
foliage plants.
73. Aglaodorum Schott
Aglaodorum Schott, Gen. Aroid.: t. 58 (1858); Nicolson,
Smithson. Contrib. Bot. 1: 3-4 (1969).
Rhizome much-branched, creeping; leaves several
per article; leaf blade thick, midrib very thick;
inflorescence 1; peduncle long; spadix stipitate,
female zone with 1 whorl of flowers; male
flowers 3-4-androus; gynoecium with whorl of
1-3 short, prismatic staminodes, ovary 1-2-
locular, placenta parietal; berry green with very
thick pericarp; seed plumule well developed.
2n = 40. Only one sp., A. griffithii (Schott) Schott,
Borneo, Malay Peninsula, Sumatra, in open
swamps, especially freshwater tidal zones and
brackish water.
14. Tribe Culcasieae Engler (1887).
Usually climbers or hemiepiphytes with slender
stems, sometimes erect or prostrate terrestrial
plants; roots, stems and leaves containing resin
canals; sclerotic hypodermis present in roots;
petiole geniculate apically; primary veins pinnate,
finer venation reticulate; spadix: female zone con-
tiguous with male, or with zone of sterile male
flowers in between; male flower: 2-4 androus, an-
thers sessile, lateral, lacking endothecial thicken-
ings, thecae dehiscing by short apical slit or pore,
connective usually strongly thickened; ovary
locules 1-ovulate; stigma sessile, large, discoid;
berry ± globose, usually red; testa thin, endo-
sperm absent.
 Araceae 63

74. Culcasia Palisot de Beauvais ovules 1-2, anatropous, placenta subbasal; berry
and seed large, plumule well developed, en-
Culcasia Palisot de Beauvais, Fl. Oware 1: 3 (1805), nom. cons.;
Bogner, Misc. Pap. Landbouwhogesch. Wageningen 19:57- dosperm absent.
63 (1980); Knecht, Contrib. Arac. Cote d'Ivoire: 188-227
(1983); Mayo, Fl. Trop. East Africa, Araceae: 16-22 (1985);
Ntepe-Nyame, Fl. Cameroun 31: 76-117 (1988). 76. Montrichardia H. Cruger Fig. 16
Montrichardia H. Cruger, Bot. Zeitung (Berlin) 12: 25 (1854),
Laticifers absent; sometimes erect, terrestrial; leaf nom. cons.; Jonker-Verhoef & Jonker, Fl. Suriname 1: 75-79
blade simple; inflorescences 1-12(-20 or more); (1953).
spathe boat-shaped, convolute basally, deciduous
or persistent; spadix sessile or stipitate, cylindric- Robust to gigantic; rhizomatous; leaves several
clavate, equalling or exceeding spathe; ovary 1-3- in apical crown; petiole sheath apically ligulate;
locular, locules 1-ovulate, ovule anatropous or inflorescence 1(-2); peduncle relatively short;
campylotropous, placenta axile, basal or basal- spathe thick; spadix subequal to spathe; stigma
parietal, berry red to orange, sometimes greenish appearing sessile on stylar region; berry sub-
yellow; seed ovoid to ellipsoid. 2n = 42, 84. About cylindric, pericarp spongiose; seed obovoid to
27 spp., tropical Africa (Senegal to Tanzania), ellipsoid, testa smooth to rough. 2n = 48. Two
root-climbers, hemiepiphytes, or erect and spp., Belize to eastern Brazil, freshwater
terrestrial. helophytes forming extensive stands along river
margins.
75. Cercestis Schott
16. Tribe Zantedeschieae Engler (1887).
Cercestis Schott, Oesterr. Bot. Wochenbl. 7: 414 (1857);
Knecht, Contrib. Arac. Cote d'Ivoire: 172-187 (1983 ); Stem a depressed-globose tuber or short rhizome
Bogner, Aroideana 8: 73-75 (1986); Ntepe-Nyame, Fl. (Z. aethiopica); leaf blade lanceolate to hastate-
Cameroun 31: 64-75 (1988); Bogner & Knecht, Adansonia sagittate; primary lateral veins pinnate, forming 1
IV, 16: 331-335 (1994).
Rhektophyllum N.E. Brown (1882); Mayo, Fl. Trop. East marginal vein, finer venation parallel-pinnate;
Africa, Araceae: 22-24 (1985); Ntepe-Nyame, Fl. Cameroun peduncle long; spathe :±:: obconic, unconstricted;
31: 58-64 (1988). spadix shorter than spathe, male and female zones
contiguous; stamens free, thecae dehiscing by
Laticifers present; sometimes robust, producing apical pore, connective thickened apically; pollen
flagelliform shoots, internodes short in flowering
zone of stem; leaf blade oblong-lanceolate to cor-
date, sagittate, hastate or tripartite, sometimes
laciniate-pinnatifid; inflorescences 1-16; peduncle
short; spathe boat-shaped to subcylindric, convo-
lute basally, persistent; spadix sessile, shorter than
spathe; ovary 1-locular, ovule hemianatropous
to anatropous, rarely campylotropous, placenta
basal-parietal; seed obovoid. 2n = 42. Thirteen
spp., Senegal to western Uganda, root -climbers or
hemiepiphytes, rarely creeping on forest floor.

15. Tribe Montrichardieae Engler (1876).

Arborescent helophytes; sclerotic hypodermis
present in roots; petiole geniculum absent; leaf
blade sagittate, hastate to trifid (rarely trisect),
primary lateral veins pinnate; spathe deciduous
after anthesis, slightly constricted, differentiated
into lower tube and upper, :±:: boat-shaped blade;
spadix: female and male zone contiguous, sterile
flowers absent; male flower: 3-6-androus, stamens Fig. 16A-E. Araceae-Aroideae. Montrichardia arborescens. A
:±:: sessile, connective overtopping thecae, thecae Flowering shoot. B Spadix. C Stamens. D Pistil, side view. E
dehiscing by short, apical slit; ovary 1-locular, Same, longitudinal section. F Seed. (Takhtajan 1982)
64 Araceae

extruded in strands; ovary 1-5-locular, locules certain species of Nephthytis and to tribe
(1-)4( -B)-ovulate, ovules anatropous, placenta ax- Zomicarpeae.
ile to subapical; testa striate, endosperm copious.
18. Tribe Thomsonieae Blume (1837).
77. Zantedeschia K. Sprengel Seasonally dormant; stem tuberous, rarely rhi-
Zantedeschia K. Sprengel, Syst. Veg. 3: 756, 765 (1826), nom. zomatous; petiole long, terete, sheath very short;
cons.; Letty, Bothalia 11: 5-26 (1973), rev.; Perry, S. Afr. J. leaf blade dracontioid, first foliage leaf of seedling
Bot. 55: 447-451 (1989). divided, all divisions upright in bud; ultimate
Pseudohomalomena A.D. Hawkes (1951). lobes usually ovate-elliptic, acuminate, primary
lateral veins forming submarginal collective veins,
Leaves several; leaf blade often variegated; inflo- finer venation reticulate; inflorescence 1, sub-
rescences 1-2; spathe white, cream, yellow, pink tended by cataphylls; spadix with sterile terminal
or rosy purple; spadix sessile or stipitate; male appendix, female zone usually contiguous with
flowers 2-3-androus; gynoecium rarely (only male; pollen extruded in strands; ovary locules 1-
in Z. aethiopica (L.) Sprengel) with whorl of about ovulate, ovules anatropous, testa smooth, embryo
3 ± spathulate staminodes, otherwise staminodes large, endosperm absent.
absent, ovary shortly ovoid; berry obovoid or
subglobose, usually green or orange, rarely yellow
and mucilaginous within (Z. aethiopica); seed 79. Amorphophallus Blume ex Decaisne
subglobose to ovoid, embryo axile. 2n = 32. Eight Amorphophallus Blume ex Decaisne, Nouv. Ann. Mus. Hist.
spp., Angola to S Africa, helophytic or terrestrial, Nat. 3: 366 (1834), nom. cons.; Hetterscheid & Ittenbach,
seasonally dormant, sometimes evergreen (Z. Aroideana 19: 7-131 (1996).
aethiopica); Z. aethiopica is a familiar garden Thomsonia Wallich (1830), nom. rej.; Bogner, Pl. Syst. Evol.
125: 15-20 (1976).
plant with many hybrids and is of considerable Plesmonium Schott (1856); Bogner, Adansonia II, 20: 305-308
importance in the cut flower trade. (1980).

Sometimes gigantic; tuber depressed-globose to

17. Tribe Callopsideae Engler (1906).
Small, evergreen, rhizome slender; leaf blade
cordate-ovate, primary lateral veins forming
single marginal vein, finer venation reticulate;
spathe completely expanded at anthesis, persis-
tent, partially closing in fruit; spadix fertile to
apex, female zone entirely adnate to spathe, con-
tiguous with male zone or with short naked zone
in between; stamens free, anthers sessile, thecae
subopposite, dehiscing by apical pore; pistils few,
slender, ovary 1-locular, ovule 1, anatropous, pla-
centa basal; embryo elongate, endosperm copious.
78. Callopsis Engler
Callopsis Engler, Notizbl. Konigl. Bot. Gart. Berlin 1:27 (1895);
Mayo, Fl. Trop. East Africa, Araceae: 54-56 (1985).
Leaves 2 per article; petiole sheath very short;
inflorescences 2, appearing with leaves; peduncle
slender, equalling or exceeding petiole; spathe
ovate-elliptic to subcircular, white; spadix:
male zone cylindric; gynoecium flask-shaped,
falcate; berry ellipsoid, green; testa ± smooth,
thin. 2n = 36. Only one sp., C. volkensii Engler, E
Africa (Kenya, Tanzania); terrestrial. Systematic
position problematic; C. shows similarities to
cylindric, rarely rhizomatous; leaf usually 1; inflo-
rescence usually appearing before leaf; peduncle
very short to long; spathe either ± undifferenti-
ated into tube and blade, or with campanulate,
cylindric or ventricose, convolute tube and erect
to spreading blade, rarely connate basally (A.
pusillus Hetterscheid & Serebryanyi); spadix male
zone rarely separated from female by zone of
sterile flowers, appendix rarely reduced or absent,
smooth to staminodial; male flowers 1-6-androus,
filaments sometimes connate, connective rarely
prominulent (A. longiconnectivus Bogner), thecae
dehiscing apically by pore or transverse slit;
gynoecia sometimes± remote; ovary 1-4-locular,
locules 1-ovulate, placenta axile to basal; berry
orange to red, rarely blue or white. 2n = 26, 28,
39. About 150 spp., tropical Africa and Madagas-
car, S and SE Asia to Melanesia, the Pacific, and
N Australia, in humid to dry forests and savannas.
A. paeoniifolius (Dennstedt) Nicolson is an im-
portant food plant in the Old World tropics,
especially in India where it is widely cultivated;
A. konjac C. Koch is cultivated in southern Japan
as a source of food gels. About 14 sections or
informal sectional groups are recognised, but
many may be unnatural (W. Hetterscheid, pers.
comm.).
 Araceae
80. Pseudodracontium N.E. Brown
Pseudodracontium N.E. Brown, J. Bot. 20: 193 (1882);
Gagnepain, Fl. gen. Indo-Chine 6: 1152-1157 (1942);
Serebryanyi, Blumea 40: 217-235 (1995), rev.
Tuberous; leaves often more than 1; inflorescence
often appearing with leaf; peduncle long; spathe
boat-shaped, convolute basally; spadix shorter
than spathe, male zone laxly flowered, appendix
composed of clavate, irregularly connate
staminodes, separated from male zone by naked
axis; male flowers 3-6-androus, filaments free or
± connate; ovary 1-locular, placenta basal; berry
ellipsoid. 2n 26; Seven spp., SE Asia
(Indochina), in moist forest.
19. Tribe Arophyteae Bogner (1972).
Seasonally dormant; stem tuberous (except most
Arophyton spp.); primary lateral veins pinnate,
forming 1 submarginal and 1-2 marginal veins,
finer venation reticulate; spadix fertile to apex
(except some Arophyton spp.); male flower a
synandrium of partially or completely connate
stamens (sometimes reduced to 1 stamen); gyno-
ecium surrounded by urceolate synandrodium,
ovary 1-locular, ovule 1, atropous, placenta basal;
testa thin, smooth, embryo large, endosperm
absent.
81. Arophyton Jumelle
Arophyton Jumelle, Ann. Mus. Colon. Marseille 36 annee, IV,
6: 23 (1928); Bogner, Bot. Jahrb. 92: 24-45 (1972), rev.
Synandrogyne S. Buchet (1939).
Humbertina S. Buchet (1941).
Stem usually rhizomatous, rarely tuberous; leaves
1-few; leaf blade cordate, hastate, tripartite or
pedatisect; inflorescences 2-3; spathe not or
slightly constricted; female zone of spadix free
or partly adnate to spathe, contiguous with male
or separated by zone of sterile or bisexual flowers,
terminal appendix sometimes present; synan-
drium 2-7-androus, thecae marginal, subglobose,
dehiscing on upper surface; berry red or green;
seed ellipsoid to globular. 2n = 38, 54, 76 (40).
Seven spp., Madagascar, on forest floor in ever-
green and deciduous forest, epiphytic or in lime-
stone crevices.
82. Carlephyton Jumelle
Carlephyton Jumelle, Ann. Mus. Colon. Marseille 27 annee, III,
7: 187 (1919); Bogner, Bot. Jahrb. Syst. 92: 10-21 (1972).
65
Leaf 1(-2); leaf blade cordate; inflorescences
1-2( -3), appearing with or before leaf; spathe not
constricted; spadix: lower part of female zone
adnate to spathe, contiguous with male or with
few bisexual flowers in between, sometimes (C.
glaucophyllum Bogner) flowers of "female" zone
almost entirely bisexual; synandrium 2-6( -1 )-
androus, elongate, stamens either completely con-
nate with thecae marginal, inverted and dehiscing
on lower surface, or incompletely connate with
filament apices ± free and thecae dehiscing
apically, sometimes remote; berry orange-red.
2n = 54, 108. Three spp., NW Madagascar, on
limestone in crevices or holes with leaf litter.
83. Colletogyne Buchet
Colletogyne Buchet, Bull. Soc. Bot. Fr. 86: 23 (1939); Bogner,
Bot. Jahrb. 92: 21-24 (1972).
Leaf 1(-2); leaf blade cordate; inflorescences 1-3,
appearing before or with leaf; peduncle longer
than spathe; spathe obovate, unconstricted, fully
expanded at anthesis, later closing, persistent;
spadix: female zone entirely adnate to spathe, con-
tiguous with male zone; synandrium reduced to 1
stamen; berry fusiform. 2n = 54. Only one sp., C.
perrieri Buchet, NW Madagascar, on limestone, in
holes or crevices with leaf litter.
20. Tribe Peltandreae Engler (1876).
Rhizomatous; primary lateral veins pinnate, form-
ing submarginal collective vein, 1-2 marginal
veins also present, finer veins ± parallel-pinnate;
inflorescences usually 2, infructescence pendent;
spathe ± constricted centrally, tube ellipsoid to
oblong, persistent, blade gaping at anthesis, later
marcescent; male flower a truncate, prismatic
synandrium of connate stamens, thecae lateral,
dehiscing by apical pore or short slit; ovary 1-
locular, ovules atropous; berry and seeds large,
testa smooth, embryo large, plumule well devel-
oped, endosperm vestigial to absent.
84. Peltandra Raf.
Peltandra Raf., J. Phys. Chim. Hist. Nat. Arts 89: 103 (1819),
nom. cons.; Huttleston, Temp. N. Am. Arac.: 65-86 (1953);
Blackwell & Blackwell, J. Elisha Mitchell Sci. Soc. 90: 137-
140 (1975).
Rhizome short, erect; petiole and sheath relatively
long; leaf blade sagittate-hastate; peduncle
subequalling petiole; spadix: female and male
66 Araceae
zones contiguous or with zone of sterile flowers
in between, terminal appendix of sterile flowers
sometimes present; synandrium 4-5-androus; gy-
noecium surrounded by cuplike synandrodium,
ovules 1-few, atropous to hemiatropous, placenta
parietal to basal; berry green or orange, 1-3-
seeded; seed ovoid. 2n = 56, 112. Three spp., tem-
perate to subtropical eastern N America (Maine to
Florida), helophytes, seasonally dormant.
85. Typhonodorum Schott
Typhonodorum Schott, Oesterr. Bot. Wochenbl. 7: 70 (1857);
Bogner, Fl. Madagascar 31: 22-26 (1975); Mayo, Fl. Trop.
East Africa, Araceae: 38-40 (1985).
Often gigantic, rhizome thick, aerial shoots form-
ing massive false stems; leaves forming terminal
crown; petiole and sheath very long; leaf blade
subtriangular to sagittate; inflorescences 2-3, pe-
duncle shorter than petiole; spadix with 4 zones
(base to apex): female zone, narrower zone of ster-
ile male flowers, subcylindric-conoid male zone,
terminal appendix of sterile male flowers;
synandria 4-6-androus; ovule 1( -2), atropous,
placenta basal, stigma broad, shallowly 3-6-lobed;
berry very large, yellow; seed compressed-
obovoid. 2n = 112. Only one sp., T. lindleyanum,
Madagascar, Comores, Pemba, Zanzibar,
Mauritius (?naturalized), a helophyte in fresh-
water swamps, also in brackish water, forming
huge stands.
21. Tribe Arisareae Dumortier (1827).
Laticifers simple; stem a tuber or rhizome; leaves
1-3; primary lateral veins of leaf blade forming
submarginal collective vein, finer venation reticu-
late; inflorescence 1, appearing with leaves; spathe
not constricted, tube margins connate, blade for-
nicate, gaping; spadix with usually smooth, sterile,
terminal appendix; flowers unisexual, perigone
absent; male flower 1-androus, thecae apically
confluent, dehiscing by 1 continuous slit; female
flowers 2-5, ovary 1-locular, ovules several,
atropous, placenta basal; endosperm copious,
starch abundant.
86. Arisarum P. Miller
Arisarum P. Miller, Gard. Diet. abr. ed., [121] (1754); Boyce,
Kew Mag. 7: 14-21 (1990).
Seasonally dormant; leaf blade cordate to sub-
hastate; spathe: tube ± cylindric or subventricose,
margins connate, blade fornicate, gaping; spadix:
female zone adnate to spathe, very short, contigu-
ous with male, male zone laxly flowered, appendix
slender to massive and knob like, hidden or shortly
exserted; berry flattened at apex, several-seeded;
seed longitudinally rugose, strophiolate. 2n = 28,
42, 56. Three spp., Mediterranean, Macaronesia.
22. Tribe Ambrosineae Schott (1832).
Stem tuberous; leaves few; leaf blade ovate-
elliptic, primary lateral veins arising basally, arcu-
ate, running into apex, finer veins reticulate;
inflorescence 1, on recurved peduncle; spathe
boat-shaped, unconstricted externally, basal tube
convolute, apical part narrowly gaping, rostrate,
internally divided into 2 chambers, an upper,
morphologically ventral one and a lower, morpho-
logically dorsal one; spadix shortly appendiculate,
adnate to spathe by longitudinal, septumlike lat-
eral dilations, ventral chamber containing female
zone consisting of only 1 gynoecium, dorsal cham-
ber containing 8-10 biseriate male flowers; male
flower a 2-androus, sessile synandrium, thecae
confluent, dehiscing by single continuous slit;
female flower 1, ovary 1-locular, ovules many,
atropous, placenta basal, style long, curved,
stigma discoid; berry many-seeded; seed ellipsoid,
testa costate, with large white conical strophiole,
endosperm copious, starch abundant.
87. Ambrosina Bassi
Ambrosina Bassi, Ambrosina Nov. Plant Gen.: 3 (1763) &
Bononiensi Sci. Inst. Acad. Comment. 5: 82-86 (1766);
Killian, Bull. Soc. Hist. Nat. Afr. N. 20: 257-278 (1929) & 24:
259-294 (1933).
Very small, seasonally dormant; inflorescence
borne at ground level; peduncle relatively long;
spathe ± horizontal; berry depressed-globose.
2n = 22. Only one sp., A. bassii L., western
Mediterranean.
23. Tribe Areae
Seasonally dormant geophytes; stem usually
tuberous; leaves several (often 1 in Sauromatum);
primary lateral veins of leaf blade or lobes or
leaflets forming submarginal collective veins,
finer venation reticulate; inflorescence usually 1,
usually appearing with or after leaves (except
Sauromatum and Biarum); spathe usually
strongly constricted, with convolute (connate in
Sauromatum, some Typhonium spp. and Biarum)
 Araceae 67

tube and gaping, erect to reflexed blade; spadix:

terminal sterile appendix always present, usually
smooth; stamens usually free; ovary 1-locular,
ovules atropous; berry subglobose; seed usually
with rather large, fleshy strophiole (aril), testa
rugose-reticulate (except Sauromatum and some
Biarum spp.), embryo ± small, endosperm
copious, starch abundant.

88. Arum L. Fig. 17F-I

Arum L., Sp. Pl.: 964 (1753); Boyce, The genus A rum: 1-196
(1993) (rev.).

Tubers elongated in some species; leaf blade sagit-

tate to hastate; peduncle short to long; spadix: fe-
male and male zones usually separated by zone of
filamentous pistillodes, rarely contiguous or inter-
mediate zone naked, male zone rather short,
usually separated from appendix by zone of fila-
mentous staminodes, appendix often stipitate;
male flowers 3-4-androus, anthers subsessile;
ovules 6 or more, placenta parietal-basal, stigma
exuding nectar droplet at anthesis; berry many-
seeded, red, pericarp juicy. 2n = 28, 42, 56, 70, 84.
Twenty six spp., temperate Europe, Macaronesia,
Mediterranean, Middle East to Central Asia, in
forests, orchards, open ground, rock crevices.
Boyce (1993) recognised two subgenera.

89. Eminium (Blume) Schott Fig. 17A-E

Eminium (Blume) Schott, Aroideae: 16 (1855, "1853"); Riedl,
Fl. Iranica, Araceae: 5- 7 (1963) & Ann. Naturh. Mus. Wien
73: 103-121 (1969) & Fl. Iraq 8: 187-203 (1985); Lobin &
Boyce, Willdenowia 20: 43-51 (1991).

Leaf blade thick, hastate to pedatifid or pedatisect,

rarely oblong-elliptic, posterior lobes of pedatifid
and pedatisect leaves ± erect and spirally twisted;
spadix: female zone separated from male by zone
of subulate sterile flowers; male flowers 2-
androus, anthers sessile; ovules 2, placenta basal Fig. 17A-1. Araceae-Aroideae. A-E Eminium alberti. A Habit.
to subbasal; infructescence subglobose, borne at B Spadix. C Stamen. D Pistil. E Same, longitudinal section. F-
or below soil level, berry white to pale lilac, peri- 1 Arum korolkowii. F Habit. G Spadix. H Stamen. I Pistil,
carp firm, not juicy. 2n = 28. Eight spp., Turkey, longitudinal section. (Takhtajan 1982)
Middle East, Central Asia.

pedatifid to pedatisect; peduncle longer than peti-

90. Dracunculus P. Miller
Dracunculus P. Miller, Gard. Diet., Abr. ed., [455] (1754);
Boyce, Thaiszia 4: 175-182 (1994).
Large; tuber stoloniferous in D. canariensis; peti-
ole sheath very long, forming false stem; leaf blade
ole; spadix: female zone contiguous with male,
sterile flowers absent, appendix very long, long-
stipitate, fistulose; male flower 3-4-androus, fila-
ments connate basally; pollen extruded in strands;
ovules few, placentas 2, apical and basal; berry
orange-yellow to red, few-seeded. 2n = 28. Two
68
spp., Mediterranean, Macaronesia, macchie, olive
groves, woodland.
91. Helicodiceros Schott ex K. Koch
Helicodiceros Schott ex K. Koch, Index Sem. Hort. Berol. 1855,
Appendix: 2 (1856), nom. cons.; Arrigoni et a!., Le piante
endemiche della Sardegna: 416-420 (1982); Boyce, Thaiszia
4: 175-182 (1994), rev.
Leaf blade pedatifid to pedatisect, posterior lobes
± erect and spirally twisted; peduncle short;
spathe held obliquely, deflexed at constriction;
spadix: female zone separated from male by zone
of long, curved, subulate sterile flowers, appendix
stipitate, long, lying on spathe blade, covered with
curved sterile flowers varying from large and
subulate at the appendix base to setiform at the
apex; male flower 2-3-androus, anthers sessile;
ovules ca. 6, placentas 2, apical one with 3-4
ovules and basal one with 1-2 ovules; berry
orange-coloured, (1- )2-3-seeded. 2n = 56. Only
one sp., H. muscivorus (L.) Engler, western
Mediterranean, on stony ground, in rock crevices.
92. Theriophonum Blume
Theriophonum Blume, Rumphia 1: 127 (1837, "1835");
Sivadasan & Nicolson, Kew Bull. 37: 277-290 (1982), rev.
Pauella Ramamurthy & Sebastine (1967).
Small; leaf blade ovate, sagittate to hastate, some-
times linear-lanceolate; peduncle usually short,
deflexing in fruit; spathe tube marcescent; spadix:
female zone few-flowered, separated from male by
rather long sterile zone partly or completely cov-
ered by filiform to subulate sterile flowers situated
either basally, centrally or apically, male zone
cylindric, sometimes with zone of sterile flowers
above, appendix stipitate or not, cylindric to long-
subulate; male flower 1-2-androus; ovules 3-9 (or
more), placentas 2, apical and basal; berry few-
seeded. 2n = 16 (14, 18). Five spp., India, Sri
Lanka, on forest floor, grassy places, damp sites.
93. Typhonium Schott
Typhonium Schott, Wiener Z. Kunst 1829 (3): 732 (1829);
Nicolson & Sivadasan, Blumea 27: 483-497 (1981); Murata,
Am. J. Bot. 77: 1475-1481 (1990); Hay, Blumea 37: 345-376
(1993); Sriboonma et al., J. Fac. Sci. Univ. Tokyo, sect. III,
15: 255-313 (1994), rev.
Lazarum A. Hay (1992).
Sometimes minute; tuber, rarely stoloniferous or
creeping rhizome; leaf blade cordate, sagittate,
hastate, trifid, trisect, pedatifid or pedatisect; in-
Araceae
florescence 1 or more; spathe tube sometimes
with connate margins, rarely septate internally
forming two chambers; spadix: female zone
separated from male by sterile zone covered en-
tirely or only basally with sterile flowers, appendix
usually stipitate, rarely with spoonlike insertion
(T. cochleare A. Hay), conic to filiform-subulate;
male flower 1-3-androus; ovules 1-2, placenta
basal; infructescence subglobose to subcylindric,
borne above soil level, berry dark red to purple.
2n = 16, 18, 20, 26, 36, 52, 54, 65, > 100 (14). About
40 spp., S and SE Asia to N and E Australia, in
humid and seasonal forests, some species weedy
and naturalised outside Asia in tropical, sub-
tropical and temperate regions; usually seasonally
dormant, sometimes evergreen; two sections
recognised.
94. Sauromatum Schott
Sauromatum Schott in Schott & Endlicher, Melet. Bot.: 17
(1832); Mayo, Fl. Trop. East Africa, Araceae: 56-60 (1985).
]aimenostia Guinea & Gomez Moreno (1946).
Leaves 1-4; leafblade pedatifid or pedatisect; inflo-
rescence appearing before leaf; spathe only slightly
constricted, tube with connate margins; spadix:
female zone separated from male by rather long
sterile zone bearing remote, patent, filiform to
clavate staminodes at base and naked above, ap-
pendix rather slender; male flower few-androus;
ovules 1-4, placenta basal; infructescence sub-
globose, partly hypogeal, berry purplish red,
darker at apex; testa smooth. 2n = 26, 52, 104. Two
spp., tropical Africa, Saudi Arabia, Yemen, S Asia,
Myanmar, SW China, in upland forest and grass-
land, occasionally epiphytic.
95. Biarum Schott
Biarum Schott in Schott & Endlicher, Melet. Bot.: 17 (1832),
nom. cons.; Bogner & Boyce, Willdenowia 18: 409-417
(1989).
Petiole sometimes hardly distinct from blade; leaf
blade linear to obovate or elliptic-oblong; inflores-
cence usually appearing before leaves, sometimes
borne partly below ground level; spathe tube mar-
gins partially to almost completely connate, rarely
free (B. aleppicum J. Thiebaut); spadix: female
zone subhemispherical, separated from male by
sterile zone bearing filiform to conic, suberect to
decurved sterile flowers at base only or through-
out (rarely naked), male zone sometimes sepa-
rated from appendix by short zone of subulate
sterile flowers, terminal appendix slender, elon-
 Araceae 69

gate to conoid, sometimes stipitate; male flowers

1-2-androus; ovule 1, placenta basal; infructe-
scence subglobose borne at or slightly below soil
level, berry white to pale lilac, pericarp firm, not
juicy; testa rugose-reticulate, rarely smooth. 2n =
16, 20, 22, 24, 26, 32, 36, 74, 96, 98. Twenty two
spp., Mediterranean, Middle East, on stony
ground, under trees and in rock crevices.

24. Tribe Arisaemateae Nakai (1943).

Laticifers simple; tuberous, rarely rhizomatous;
leaf blade usually compound: trisect, pedatisect or
radiatisect, rarely cordate or trifid; spathe tube
usually cylindric, blade usually fornicate; spadix
with terminal appendix, appendix often cylin-
dric, clavate or elongate-tapering; flowers uni-
sexual, perigone absent; ovary 1-locular, ovules
atropous, placenta basal; testa rough, endosperm
copious.

96. Pinellia Tenore

Pinellia Tenore, Atti Acad. Sci. Napoli 4: 69 (1839), nom. cons.;
Li Heng, Fl. Reip. Pop. Sin. 13:200-206 (1979); Ulharz, Beitr.
Bioi. Pflanz. 60: 277-291 (1986) & ibid. 61: 241-282 (1986);
Boyce, Kew Mag. 5: 18-21 (1988).

Petiole sometimes bearing tubercles; leaf blade

usually trisect or pedatisect, rarely entire and
ovate-oblong to cordate; peduncle equal or longer
than petiole; spathe weakly constricted, tube nar-
row, persistent, almost closed at mouth by trans-
verse septum, blade ± fornicate; spadix: female
zone adnate to spathe, separated from male by
spathe septum and short, naked zone, appendix
smooth, subulate, slender, long-exserted; male
flower 1-2-androus, stamens free or sometimes
irregularly connate; ovule 1, ± atropous; testa
rugulose. 2n ± 26, 52. Six spp., temperate E Asia,
P. ternata often naturalised elsewhere, in forests
or on rocks, also weedy in fields.
Fig. 18A-M. Araceae-Aroideae. A-E Arisaema amurense. A
Habit. B Staminate spadix. C Synandrium. D Pistillate spadix.
E Pistil. F, G A. thunbergii. F Inflorescence with long exserted
97. Arisaema Mart. Fig. 18 appendix. G Infructescence. H-N A. griffithii. H Habit. I Inflo-
Arisaema Mart., Flora 14: 459 (1831); Huttleston, Temp. N. rescence with flexuose appendix. J Staminate spadix. K
Am. Arac.: 87-154 (1953) (N Amer. spp.); Li Heng, Fl. Reip. Synandrium. L Pistillate spadix. M Pistil, longitudinal seciton.
Pop. Sinicae 13: 116- 194 (1979); Ohashi & Murata, J. Fac. (Takhtajan 1982)
Sci. Univ. Tokyo, Bot. 12: 281-336 (1980); Murata, J. Fac.
Sci. Univ. Tokyo, Bot. 13: 431-482 (1984); Mayo & Gilbert,
Kew Bull. 41: 261-278 (1986) (African spp.); Murata, Pl. Sometimes large; tuber depressed-globose, rarely
Sp. Bioi. 2: 57-66 (1987) & Bot. Mag. Tokyo 103: 371-382 rhizomatous, sometimes stoloniferous; petioles
(1990).
Flagellarisaema Nakai (1950). usually long, often forming false stem; leaf blade
Heteroarisaema Nakai (1950). trisect, pedatisect or radiatisect, rarely cordate or
Pleuriarum Nakai (1950). trifid; peduncle short to long; spathe tube usually
Ringentiarum Nakai (1950). cylindric, mouth often flaring with revolute to
70
grotesquely auriculate margins, blade usually for-
nicate, often with elongate or filamentous apex;
spadix unisexual (paradioecious) or monoecious,
male zone usually laxly flowered, contiguous with
female, sometimes separated from appendix by
short zone of filamentous or subulate staminodes,
appendix hidden or exserted, rarely absent (A.
exappendiculatum H. Hara), often cylindric, clav-
ate or elongate-tapering, sometimes drawn out
into an extremely long flexuose thread, rarely
composed of several long threads; male flowers
remote, 2-5-androus, stamens connate, syn-
andrium ± sessile to long-stipitate; ovules 1-9 per
ovary; berry red, few-seeded; testa rough. 2n = 20,
22, 24, 26, 28, 39, 42, 48, 52, 56, 72, 112, 140, 168
(64). About 170 spp., eastern and southeastern
USA, northern Mexico, E and NE Africa, Arabian
Peninsula, temperate E Asia, tropical S, SE and
E Asia, Indonesia, in the tropics only at high
altitudes. Murata (1990) recognized 11 sections.
25. Tribe Colocasieae Engler (1876).
Laticifers anastomosing (except Ariopsis); pri-
mary lateral veins of leaf blade or leaflets
(Protarum) pinnate, forming submarginal collec-
tor vein, at least 1-2 other marginal veins also
present; spathe usually constricted centrally (ex-
cept Ariopsis, sometimes constricted twice in
Remusatia) with lower part forming a convolute,
persistent tube and upper part a gaping, ± boat-
shaped, marcescent to caducous blade; flowers
unisexual, perigone absent; male flower a synan-
drium of several connate stamens; endosperm
copious, with abundant starch.
98. Ariopsis Nimmo
Ariopsis Nimmo in Graham, Cat. Pl. Bombay: 252 (1839).
Laticifers simple; plant small; stem a partially hy-
pogeal tuber; leaves !-several; leaf blade peltate,
ovate-cordate to -emarginate; inflorescences 1-3,
appearing either before or with the leaves; pe-
duncle slender, longer than spathe; spathe ovate,
boat-shaped, fornicate, not constricted, persis-
tent; spadix: female zone adnate to spathe, few-
flowered, separated from male by short naked
axis, terminal appendix absent; synandria con-
nate, peltate, stipitate, 6-thecate; ovary 1-locular,
ovules many, atropous, placentas 4-6, parietal;
berry many-seeded; testa costate, endosperm
copious. 2n = 28, 84. Two spp., tropical S Asia
(Myanmar to Western Ghats); in evergreen forest,
seasonally dormant.
Araceae
99. Protarum Engler
Protarum Engler, Bot. Jahrb. 30, Beibl. 67: 42 (1901); Bogner,
Palmengarten 37: 40 (1973); Robertson, Fl. Pl. Seychelles:
259 (1989).
Tuber globose; leaf 1; leaf blade subpedatisect to
almost radiatisect, leaflets shortly stalked, lan-
ceolate, acuminate to acute; inflorescence 1; pe-
duncle short; spadix shorter than spathe, female
zone conoid, male zone obconic, female and male
zones separated by slender sterile zone, terminal
appendix thick, obtuse; synandrium 3-androus, ±
6-lobed; female flower with whorl of 4-6 free
staminodes surrounding gynoecium, ovary 1-
locular, ovules 4, hemiatropous, placenta basal;
berry 1-seeded; seed ellipsoid, testa costate, em-
bryo discoid, endosperm copious. 2n = 28. Only
one sp., P. sechellarum, Seychelles Is., seasonally
dormant.
100. Steudnera K. Koch
Steudnera K. Koch, Wochenschr. Giirtnerei Pflanzenk. 5: 114
(1862).
Sometimes robust; stem epigeal, stout, erect or
sometimes creeping; leaves 1-several; leaf blade
peltate, ovate-subcordate; inflorescences 1-2;
spa-the ovate or ovate-lanceolate, not constricted,
barely convolute at base, ± fully expanded at an-
thesis, upper part later reflexed; spadix: female
zone mostly adnate to spathe, contiguous with and
longer than male zone, terminal appendix absent;
synandrium 3-6-androus; gynoecium usually sur-
rounded by 2-5 short staminodes, ovary !-locular,
ovules many, hemiatropous, placentas 2-5, pari-
etal; berry many-seeded; testa costate. 2n = 28, 56.
Eight spp., tropical S to SE Asia (Assam to
Vietnam), on forest floor, sometimes seasonally
dormant.
101. Remusatia Schott
Remusatia Schott in Schott & Endlicher, Melet. Bot.: 18 (1832);
Mayo, Fl. Trop. East Africa, Araceae: 40-42 (1985); Li Heng,
Aroideana 10: 23-26 (1988); Li Heng & Hay, Acta Bot.
Yunnanica, Suppl. 5: 27-33 (1992).
Gonatanthus Klotzsch (1841).
Stem tuberous, subglobose, producing shoots
bearing small cataphylls and axillary, clustered
bulbils covered by recurved scales; leaves 1-2, leaf
blade peltate, lanceolate to ovate-cordate; inflo-
rescence 1, appearing before leaf; peduncle short;
spathe with 1 or 2 constrictions, blade broad or
 Araceae
narrow and very long, reflexed at anthesis or
erect-sigmoid; spadix: female zone separated from
male by much narrower zone of sterile synand-
rodes, terminal appendix absent; synandrium 2-
3-androus; pollen extruded in strands; female
flowers lacking staminodes, ovary 1-locular or
partially 2-3-locular apically, ovules several to
many, hemiatropous, placentas 2-4, parietal or
basal or both (R. yunnanensis H. Li & A. Hay);
berry many-seeded; testa thick, succulent, ver-
ruculose to irregularly furrowed. 2n = 28, 42, 56.
Four spp., tropical Africa, Madagascar, S and SE
Asia, Indonesia, N Australia, in seasonal or ever-
green forest, terrestrial, rupicolous or epiphytic,
seasonally dormant.
102. Colocasia Schott Fig. 15A
Colocasia Schott in Schott & Endlicher, Melet. Bot.: 18 (1832),
nom. cons.; Plucknett in Wang, Taro: 14-19 (1983); Shaw in
Chandra, Edible Aroids: 160-168 (1984).
Sometimes gigantic and arborescent; stem either
hypogeal and tuberous or mostly epigeal and mas-
sive; leaves several; leaf blade peltate, ovate-
cordate; inflorescences usually many; peduncle
much shorter than petiole; spathe constricted; spa-
dix: female zone separated from male by much
narrower zone of sterile synandrodes, terminal
appendix erect, ± smooth, sometimes reduced
to tiny stub; synandrium 3-6-androus; pollen
extruded in strands; female flowers lacking
staminodes, ovary 1-locular, ovules many, hemi-
atropous, placentas 2-4, parietal; berry greenish,
many-seeded; testa thickish, costate. 2n = 28, 42.
Eight spp., NE India, SE Asia, Indonesia, seasonally
dormant or evergreen. C. esculenta (L.) Schott is
cultivated and naturalised throughout the tropics
with hundreds of cultivars; it is the world's most
economically important aroid - for details see re-
cent reviews by Wang (1983) and Chandra (1984)
cited above. Two sections are recognised.
103. Alocasia (Schott) G. Don
Alocasia (Schott) G. Don in Sweet, Hart. Brit., ed. 3:
631 (1839), nom. cons.; Burnett, Aroideana 7: 67-162
(1984); Hay, Aroids of Papua New Guinea (1990); Hay &
Wise, Blumea 35: 499-545 (1991) (reg. rev. New Guinea and
Austral.)
Xenophya Schott (1863); Nicolson, Blumea 16: 115-118 (1968).
Schizocasia Schott ex Engler (1880).
Panzhuyuia Z.Y. Zhu (1985).
Sometimes gigantic and arborescent; stem hypo-
geal to epigeal and erect; leaves several; leaf blade
71
ovate-cordate to sagittate, sometimes peltate, mar-
gin rarely sinuate to deeply pinnatifid; inflores-
cences several; spathe constricted, blade usually
boat -shaped and erect, rarely fornicate, sometimes
reflexed, rareJy persistent; spadix: female zone
separated from male by much narrower zone of
sterile synandrodes, terminal appendix always
present; synandrium 3-8-androus; pollen ex-
truded in strands; female flowers lacking stami-
nodes, ovary 1-locular or partially 3-4-locular at
apex, ovules hemiatropous or anatropous, few,
placenta basal, style usually distinct, narrower
than ovary; berry 1-few-seeded, usually reddish;
testa smooth. 2n = 28, 42, 56, 70, 84. About 70 spp.,
S and SE Asia to northern Australia, Melanesia,
Pacific [A. macrorrhizos (L.) Schott widely
naturalised throughout the tropics], seasonally
dormant to evergreen. Two sections recognised by
Krause (in Engler and Krause 1920); Xenophya is
reduced here following Hay and Wise (1991) but
merits recognition as a separate section.
26. Tribe Pistieae Blume (1836).
Laticifers absent; acaulescent; leaves rosulate,
densely pubescent; petiole very short; leaf blade
obovate-cuneate to oblong, midrib absent; inflo-
rescence 1, very small; spathe ± constricted, lower
margins connate forming tube, free margins be-
tween tube and blade folded to form a partition
separating upper male chamber from lower fe-
male one, blade gaping; spadix: female zone with a
single basal gynoecium and thin pouch-shaped
flap below spathe partition, male zone with 2-8
flowers in a whorl, appendix naked, very short;
synandrium sessile, of 2 connate stamens; ovary
obliquely adnate to spadix axis, 1-locular, ovules
numerous, atropous, placenta parietal (morpho-
logically basal), style distinct, curved; berry
thin-walled, several-seeded, pericarp eventually
decaying; seed cylindric, testa thick, reticulate-
alveolate, with micropylar operculum formed by
both integuments, embryo minute, endosperm
copious.
104. Pistia L.
Pistia L., Sp. Pl.: 963 (1753); Buzgo, Bot. Jahrb. Syst. 115:557-
570 (1994).
Plant small, stoloniferous; roots much-branched,
feathery; leaf blade somewhat spongy; peduncle
very short. 2n = 28. Only one sp., P. stratiotes L.,
pantropical, free-floating evergreen aquatic.
72 Araceae
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Bromeliaceae
L.B. SMITH 1 and W. TILL
Bromeliaceae Juss., Gen. Pl. 49 (1789), "Bromelieae".
Perennial rosette herbs with a short axis, more
rarely with an elongated stem, or rosette trees,
from few centimeters to more than lOrn, terres-
trial, saxicolous, or epiphytic. Roots absorbing in
terrestrials, modified to holdfasts in epiphytes and
epilithes, rarely lacking. Leaves spirally arranged,
dilated-sheathing at the base, undivided, entire or
spinose-serrate, usually bearing peltate trichomes
at least when young. Stipules lacking. Inflores-
cence terminal, often becoming pseudolateral by
sympodial growth, scapose or sessile, compound
or simple, usually bearing brightly colored con-
spicuous flower bracts or "petaloid" scape bracts.
Flowers perfect or sometimes functionally uni-
sexual (all Hechtia, Catopsis p.p.), usually
actinomorphic. Perianth heterochlamydeous, the
3 sepals and 3 petals free or connate, the petals
often with scalelike basal appendages and/or lon-
gitudinal callosities. Stamens 3 + 3; filaments free,
or joined to the petals and/or to each other; an-
thers introrse, bithecal, 4-loculate, latrorsely de-
hiscent. Ovary superior to inferior, 3-carpellate,
3-loculate; placentae axile, extending the length of
the ovary or variously reduced; ovules anatro-
pous, naked or with a chalaza! appendage; style 1,
stigma usually 3-lobed; septal nectaries always
present. Fruit usually a septicidal capsule or
baccate; seeds winged, plumose, or naked; embryo
small, cylindrical, situated laterally at the base of
the copious starchy endosperm.
A neotropical family of 56 genera and more
than 2600 species distributed from N America
(Virginia) to southern America (Patagonia); a
single species in W Africa.
CHARACTERS OccuRRING IN FEw GENERA AND
SPECIES. Rheophytic plants in Pitcairnia and
Guzmania; climbing ones in Pitcairnia; distichous
phyllotaxis in Tillandsia, mainly in subgenus
Diaphoranthema; petiolate leaves in Pitcairnia,
Ronnbergia, Bromelia, and Cryptanthus, cuticular
1 Deceased.
 Bromeliaceae 75

water absorption without functional peltate tri-

chomes in Acanthostachys; lateral inflorescence in
Dyckia; reduction to 1 flower in Tillandsia; pres-
ence of glandular hairs in Puya, Deuterocohnia,
and Pitcairnia; zygomorphic flowers in Pitcairnia
and Tillandsia; !epidote petals in Pitcairnia,
Dyckia, and Hechtia; paired petal appendages in
the pitcairnioid genera Brewcaria and Steyer-
bromelia; more than 1 vascular branch in each
filament in Aechmea, Billbergia, Bromelia, Dyckia,
Encholirium, and Vriesea; connectives with an
apical appendage in Androlepis or an apical glan-
dular outgrowth in Neoregelia eleutheropetala
(Ramirez 1991); semiintrorse anther dehiscence in
Dyckia and Vriesea; undivided stigma in Wer-
auhia and Aechmea; additional apical loculicidal
dehiscence of the capsules in Deuterocohnia,
Dyckia, Encholirium, Fosterella, Hechtia, Pit-
cairnia, and Puya; seeds with a strongly elongated
chalaza! appendage in A canthostachys.

VEGETATIVE MoRPHOLOGY. The roots are often

much modified due to functional adaptations.
Intracauline roots are present in species with a
distinct stem (Fig. 19).
The usually erect stem ranges from extremely
short in species with rosulate leaves to the stout 8-
9.5-m-high columns of Brocchinia micrantha and
Puya raimondii, it is usually branched and ligni-
fied to various degree. Tillandsia usneoides is
distinctive by its pendent stems without roots.
Rhizomes may travel several meters underground
(Bromelia, Deinacanthon, Neoglaziovia, Pit-
cairnia, and Pseudananas) or be completely aerial
as in species of Aechmea, Cryptanthus, Neoregelia,
Tillandsia, and Vriesea.
Leaves vary from lingulate to narrowly triangu-
lar or even filiformous, from fiat and soft to rigid
and distinctly succulent. The leaf sheaths are usu-
ally dilated but often merge in the leaf blades,
rarely both are separated by a distinct petiole. The
green coloration is often masked or ornamented
by anthocyanin or by a dense indumentum of tri-
chomes. In Tillandsioideae the leaves are always
entire, while in Pitcairnioideae and Bromelioideae
the leaves are predominantly spinose-serrate.
In Pitcairnia leaf dimorphism or even polymor-
phism is observed in which the outer leaves are
narrow and spiny and the inner ones broad and
even pseudopetiolate. In some long-caulescent
species one type may grade into the other, some-
times repeatedly every year. In rosulate species the
leaves often form a tank (phytotelm, Janetzky and
Vareschi 1993; Krugel1993) which holds water, a
feature that is lacking in caulescent species with
Fig. 19. Bromeliaceae. Schematic longitudinal section of an
axis showing intracauline adventitious roots growing down-
wards in the cortex and branching in the substrate. (Weber
1954)
trichoma! moisture and nutrient absorption. Dis-
tichous phyllotaxis is rare (e.g., Till a ndsia subge-
nus Diaphoranthema).
In terrestrial Pitcairnioideae, tank rosettes are
lacking with the exception of Brocchinia species
which have well-developed "pitchers". In Tilland-
sioideae and Bromelioideae tanks of various types
are common.
VEGETATIVE ANATOMY. A comprehensive and
critical treatment was given by Tomlinson (1969),
and modern data are supplied by Benzing (1980,
1990), who should be consulted also for functional
and ecological aspects.
From the periphery to the center, the roots con-
sist of a piliferous layer with well-developed root
hairs in soil roots, of a multiseriate exodermis of
suberized cells, of the outermost cortical layers
immediately within the exodermis which are
always differentiated as a wide sclerotic cylinder
forming a conspicuous mechanical layer, of the
middle cortex of thin-walled parenchyma, of
the lacunose inner cortex, of a uniseriate thick-
walled endodermis, of a 1-(2)-layered thin-walled
76 Bromeliaceae
pericycle, and of a stele with the vascular tissues
and pith cells. The pith is strongly lignified in spe-
cialized taxa (Strehl and Winkler 1983). The xylem
always contains at least a few vessels with scalari-
form perforations only or tracheids (Wagner
1977); only for Pitcairnia have specialized vessels
been recorded (Cheadle 1955). The outer tissues
are separated from the inner ones by the clearly
defined endodermis. Intracauline roots are
located in the stem cortex and differ from external
roots in the absence of root hairs, exodermis, and
lacunae. Roots of epiphytic species are often
modified from the above scheme.
The stem anatomy is quite uniform within the
family. A thin cuticle covers an epidermis which
contains spherical silicia bodies. The narrow
cortex has narrow, elongated, thick-walled, ligni-
fied cells in the outer hypodermal layers which
eventually are suberized in old stems; the inner
hypodermal layers are transitional to thinner-
walled, more isodiametric cells of the inner cortex.
Periderm is developed from secondary meristem-
atic layers. The central cylinder is delimited from
the cortex by a narrow, 2-4-layers-wide zone of
elongated cells which become sclerotic in older
stems. The peripheral sclerenchyma is separated
from the vascular tissue by a narrow, thin-walled
layer, termed pericycle by some authors. The vas-
cular bundles lie diffuse in the center but are more
congested in the periphery of the central cylinder,
and the bundles have a more or less distinct
bundlesheath. The xylem contains mainly trache-
ids, vessels are rare and less often encountered
than in roots (Cheadle 1955; Wagner 1977). The
sieve elements of the phloem contain protein crys-
talloids of the Pile type in their plastids (Behnke
1981). The anatomical character of the stem is
somewhat confused by the strong suppression of
internodial growth and by the junction of adventi-
tious roots. The arrangement of the stem bundles
conforms to the meristele type, but the collateral
vascular bundles are sometimes reversed and may
be sickle-, crescent-, or annular-shaped. The stem
of the Bromeliaceae is capable of transition to pri-
mary radial growth through a cambial cap at the
stem apex. This growth tissue apparently can con-
tinue to develop after flower formation.
The fundamental character of the leaf of the
Bromeliaceae is the small-celled and small-
lumened epidermis with variously thickened
walls. Another unique feature of the epidermis is
that the greater cell dimension is often at right
angles to the long axis of the leaf. Some species are
distinguished by lateral walls strengthened by
sinuous folds so that adjacent cells interlock with
one another. The entire surface of the epidermis
has a smooth coating of the cuticle (Harms 1930).
Epicuticular wax crystalloids of the Strelitzia type
(Barthlott and Frolich 1983) occur in several spe-
cies. Adequate protection and strength for the
leaves of most Tillandsioideae and Pitcairnioideae
seem to be provided by the epidermis. In Puya,
Ananas, and others the epidermis is further
strengthened by a sclerenchymatous hypodermis
whose cells are generally longer than the epider-
mal cells. These hypodermal cells often have
brownish contents and sometimes contain sili-
ceous crystal fragments. Opaque yellowish drop-
lets of tannins also occur in the hypodermal cells
and seem to be characteristic of the family.
Chlorophyll is generally not contained in typical
palisade cells, which are completely absent from a
number of species. Chlorophyll-bearing cells are
nearly always separated from the upper leaf
surface by water-storage cells and their position
otherwise includes a great variety of types. Corru-
gated leaves with longitudinal furrows on their
lower side provide mechanical strength which is
caused by vascular bundles and bundle-sheath
fibers that make up the convex portion of the cor-
rugations. The stomata usually lie on the sides,
and the stalks of the trichomes lie in the bottom of
the concave portion. In cross-section, leaves are
often crescent-shaped, which enhances surfacial
water conduction. In addition, such leaves are 51
times more resistant to bending than a flat blade
of the same surface area.
The stomata of the Bromeliaceae are located
almost entirely on the underside of the leaf, usu-
ally at the same level as the epidermis, although
raised accessory cells sometimes produce pseudo-
stomata. Stomata are characterized by especially
narrow lumina and the presence of at least 1 lat-
eral and 1 polar pair of accessory cells. Guard cells
seem to be relatively uniform throughout the fam-
ily but the substomatal pores are distinctive
enough for Robinson (1969) to have used them as
part of a classification system for Navia. Stomatal
movement is usually lacking, with the exception of
Ananas; guard cells are not separated.
The peltate trichomes have a basic function in
the absorption and retention of moisture by the
leaves (Benzing 1980; Benzing et al. 1985). They
consist of a foot cell, a stalk of few living cells and
a shield of dead cells. In Brocchinia reducta, how-
ever, the cells of the shield are alive at maturity.
Simple hairs without a shield (Fig. 20A,B) are rare
(Lindmania, Navia, Pitcairnia, Strehl and Winkler
1982). The most primitive forms of trichomes are
probably simple (Lindmania, Navia, Pitcairnia)
 Bromeliaceae
and stellate hairs (Pitcairnia, Puya) in Pitcairnio-
ideae, the latter type with long, slender, slightly
fused cells radiating from 1 central cell. Trichomes
of most Pitcairnioideae, however, have an undi-
vided central cell or a central cell group which
differs in size and/or content from the peripheral
cells. Pitcairnia, Lindmania, and Fosterella have
primitive trichomes from which radiate long
filamentous cells only slightly fused to each other
(Fig. 20D,E). In Dyckia the central cells are sur-
rounded by concentric circles of rounded cells
that are only slightly elongated radially. In most
Bromelioideae the cells are polygonal so that the
outer and inner ones differ little (Fig. 20K). In
several species the trichomes are differentiated
into a disk and a wing, especially in Neoregelia
and some Aechmea. However, in Bromelia and
Orthophytum the trichomes are similarily primi-
tive as in Pitcairnioideae. The trichomes of the
Tillandsioideae, especially the genus Tillandsia,
form the highest stage of development (Fig. 20F-I)
and are constructed according to a regulary pat-
tern. The stalk consists of 1-3 thin-walled cells
(but up to 8 in Glomeropitcairnia) and an apical
dome cell. The shield consists of 4 right-angled
77
Fig. 20A-K. Bromeliaceae. Epidermal hairs (A, B) and scales
(C-K). A-C Navia fontoides. ASimple and B branched hairs. C
Surface view. D, E Fosterella penduliflora. D Transversal
section. E Surface view. F, G Tillandsia flexuosa. F Transversal
section. G Surface view. H, I T. recurvata. H Transversal
section. I Surface view. J, K Nidularium innocentii. J Transver-
sal section. K Surface view. (Tomlinson 1969)
cells in the center surrounded by a ring of 8, then
a ring of 16, and in many species an outer ring of
64 cells. The trichome may be fused to the epider-
mis as far as the 8-celled ring. Although the cell
division formula of 4 + 8 + 16 + 64 is the general
rule, many exceptions occur, noteworthy being
the 4 + 8 + 64 in Catopsis, Glomeropitcairnia, and
some Vriesea. The shield varies from perfectly
orbicular to extremely excentrical.
INFLORESCENCE STRUCTURE. Inflorescences are
paniculate and are made up of racemes or spikes.
Reductions of the axes lead to corymbs and heads
and even to single flowers. The primary axis usu-
ally flowers only once with a flowering period
from a few days to over 1 year in large many-
flowered species of Aechmea. In Deuterocohnia
78 Bromeliaceae

(except Abromeitiella) the inflorescence becomes succulent in the lower parts as in Racinaea. They
woody, has a cambiumlike meristematic cylinder, may be plicate in several species of Tillandsia.
and flowers continuously for several years, with Anthers are dorsifixed or basifixed, usually
buds opening at the beginning of each rainy linear, rarely with an elongated connective
season. Usually, the renewing shoots are formed (Androlepis).
during the period of flowering, thus providing The ovary is superior to inferior in the
additional vegetative propagation. A minority of Pitcairnioideae, superior in the Tillandsioideae
Bromeliaceae is hapaxanthous and survives only (except of Glomeropitcairnia), and inferior in
by seed. the Bromelioideae. However, Bohme (1988) con-
The sequence of flowering is usually acro- cluded from the position of the septal nectaries
petalous. Exceptions occur in Canistrum, where that all gynoecia of the Bromeliaceae are more or
the middle region flowers first, and in species of less inferior. The carpels are only weakly fused at
Aechmea subgenus Ortgiesia, where acropetalous the edges in Pitcairnioideae but distinctly in the
and basipetalous flowering, as well as such as in other two subfamilies. Septal nectaries are devel-
Canistrum, can be found among closely related oped at various degrees and very rarely reduced
species. in Araeococcus (Bohme 1988).
The style is slenderly cylindric and straight or
FLOWER STRUCTURE. In general, the sepals are curved according to the direction of the corolla
contorted with the right margin covered, and the (Pitcairnia), or short and stout or even lacking
median sepal is in abaxial position. However, in (Dyckia, Hechtia, Racinaea, Tillandsia).
Brocchinia, Dyckia, and others the sepals are Stigma morphology (Fig. 21) has proved to be a
imbricate with both margins of the median sepal valuable taxonomic character with far-reaching
covered, and in Cottendorfia florida cochlear, consistency in Tillandsia subgenus Tillandsia
where the median sepal covers both margins (Gardner 1982, 1986b) and in the thecophylloid
of the adaxial ones. Sepals vary from symme- species of Vriesea (now Werauhia; Utley 1978,
trical to strongly asymmetrical (Racinaea, most 1983). Brown and Gilmartin (1984, 1989b), as well
Bromelioideae ). as Schill et al. (1988), defined the types of stigma
The petals are nearly always regular but in many structure: free erect untwisted lobes (simpleerect),
species of Pitcairnia and Werauhia, one is curved, joined erect untwisted lobes lacking papillae and
forming a hood over the stamens. Form varies forming a cup (cupulate), conduplicately folded
from linear to ovate; they may be entire, finely lobes twisted together spirally (conduplicate-
crenate or even denticulate and the apex may be spiral), bladelike lobes folded in an irregular
round, retuse, apiculate, or mucronate. Color may convolute fashion (convolute-blade), and a prolif-
be white, red, purple, rose, yellow, green, blue, or eration of irregular growth in the marginal stig-
violet of various shades and intensities. The most matic line of each lobe (coralliform).
interesting feature of the petals is ligular append-
ages on the inside base which have been used for Pollen Morphology. The pollen grains (Fig. 22) are
taxonomic classification. The structure may be a generally single, only in Androlepis and Hohen-
simple membranaceous scale or flap (Pitcairnio- bergiopsis are they known to be united in tetrads.
ideae) or it may consist of 2 separate scales The exine is subtectate and usually reticulate, less
(Brewcaria, Steyerbromelia, Bromelioideae, Til- often scrobiculate or pertectate, with the sexine
landsioideae) which are usually serrate or !acini- thicker than the nexine. Grains are either sul-
ate (Brown and Terry 1992). Harms (1930) cate or 2-4-porate to pantotreme. Erdtman and
suggested that the evolution of the 2 more or less Praglowski (1974) found the porate type restricted
separate scales is explained by pressure of the fila- to subfam. Bromelioideae (12 genera studied), al-
ment on a single simple ligular primordium. though, apart from this, sulcate pollen is present
Arrais (1989) suggested a staminodial nature of in 11 genera of this subfamily; Aechmea being het-
the petal appendages. The function of the append- erogeneous in this respect. Subfamily Pitcairnio-
age is not known with certainty. The early inter- ideae and Tillandioideae have invariably sulcate
pretation as a mechanism to avoid loss of nectar grains, although the data basis is still small (12
has been queried. genera of Pitcairnioideae and three of Tillandio-
The stamens are usually free and contain 1 vas- ideae studied). With respect to both exine sculp-
cular bundle, rarely 2, or up to 6 in Encholirium ture and number and position of apertures, a
(Arrais 1989). The filaments are linear, fiat, or certain similarity between Bromeliaceae and
thickened throughout or apically, or distinctly Amaryllidaceae, especially subfam. Hypoxido-
 Bromeliaceae
A,B
D,E
ideae, was noted by Erdtman and Praglowski
(1974).
Since the time ofErdtman and Praglowski, more
palynological data have accumulated (Ehler and
Schill 1973), but broader systematic or anthe-
cological relationships have not yet been per-
ceived (Benzing 1994). However, Halbritter (1988,
1992) successfully demonstrated the value of the
aperture morphology for classification at generic
and subfamiliallevel. A simple sulcus with smooth
margins is typical of Pitcairnioideae and Catopsis
while porate pollen is restricted to Bromelioideae.
A "distal germination area" seems to occupy
an intermediate position and is typical of Pitcair-
nia, Tillandsioideae, Billbergia, Cryptanthus, and
Orthophytum. Inaperturate pollen is rare
(Guzmania, Aechmea, Canistrum).
EMBRYOLOGY. The anther tapetum is secretory,
pollen mother cell division is successive, the
pollen grains are 2-celled, ovules are bitegmic
and crassinucellate, the embryo sac is of the
Polygonum type, occasionally proliferation of the
antipodals occurs, endosperm is Helobial, and
79
c
F
Fig. 21A-F. Bromeliaceae. Stigma morphology. A Pitcairnea
andreana, conduplicate-spiral. B Tillandsia castellanii,
simple-erect. C Vriesea psittacina, convolute-blade. D
Werauhia gigantea, cupulate. E Tillandsia cyanea, coralli-
form. F Canistrum aurantium, conduplicate-spiral. Bar
SOO[!m. (Photo G. Gortan)
embryogeny follows the Asterad type (J ohri et al.
1992). Polyembryony has been found in several
species of Tillandsia subgenus Diaphoranthema
(Subils 1973).
KARYOLOGY. Chromosome numbers have been
tabulated by McWilliams (1974). The chromo-
somes are very small; Pitcairnioideae have uni-
form complements of equal-sized chromosomes
mostly based on x = 25. The same base number
predominates in the Bromelioideae, in which
varying degrees of bimodality are observed. In
Aechmea tillandsoides n = 21 was explained as
having arisen through a combination of n = 25
and n = 17 (unique for Cryptanthus) at an early
date (Marchant 1967); other numbers reported in
this subfamily include n = 24, 36, and 48.
 80 Bromeliaceae
A,B
D,E
Fig. 22A-F. Bromeliaceae, pollen grains. Pitcairneoideae: A
Puya spathacea, simple sulcus. B Pitcairnia grafii, insulae
type. Tillandioideae: C Tillandsia jucunda var. jucunda,
operculum type. D Guzmania acorifolia, diffuse sulcus.
Bromelioideae: E Lymania corallina, biporate. F Hohenbergia
stellata, tri- to tetraporate. Bar lOf!m. (Photo H. Halbritter)
Uniformly bimodal complements characterize the
Tillandsioideae, in which a greater variety of base
numbers is reported, which include, apart from n
= 25, also n = 16, 17, 27, and 32. Polyploidy is
quite rare within the family (Bromelia, Fosterella,
Guzmania, Nidularium, Pseudananas, Tillandsia,
McWilliams 1974; Brown and Gilmartin 1986,
1989a; Till 1992) and the base number x = 25
appears to be paleopolyploid itself. Recent studies
(Brown and Gilmartin 1986, 1989a) suggest a
hybridization of 2 paleodiploids with n = 7 and
n = 8, respectively, and a subsequent secondary
hybridization step with the n = 8 lineage yielding
x = 25. Decreasing base numbers are explained
with aneuploid reductions. Elevated chromosome
numbers, however, do not provide reliable evi-
dence for polyploidy in the history of a taxon
(Greilhuber 1995) as the isozyme numbers in
Bromeliaceae are typical of diploids (Soltis et al.
1987).
PoLLINATION. Hummingbirds (Trochilidae) are
the major pollinators ofbromeliads (Harms 1930;
Utley 1983; Gardner 1986a; Kraemer et al. 1993;
Araujo et al. 1994; Martinelli 1994; Seres and
c
F
Ramirez 1995) with Trochilinae predominant in
montane regions. Honeycreepers (Coerebidae)
prevail in highland areas. Species of Puya subge-
nus Puya have the outer part of each branch sterile
and this forms a distinct perch that is used by
icterids (oriole-blackbird family).
Hymenoptera (bees, bumble bees) and Lepi-
doptera (moths, butterflies) are listed as visitors
by several authors (Harms 1930; Gardner 1986a;
Till1992; Martinelli 1994). Hawkmoth pollination
may be supposed for Tillandsia xiphioides with
long and fragrant flowers.
Bat pollination is reported for Encholirium
(Sazima et al. 1989), Alcantarea, Vriesea, and
Werauhia (Utley 1983; Martinelli 1994) with
glossophagine bats as the pollinator.
The size and structure of the septal nectary
(Bohme 1988) and the amount and quality of the
nectar offered (Freeman et al. 1985; Bernardello et
al. 1991, 1994) are crucial for the pollination type
and the visitors. In entomophilous flowers the
septal nectaries are smaller and less nectar is
offered than in ornithophilous ones. On the other
hand, entomophilous flowers often enhance their
attraction function by enlargement of the petals
(salver flowers) and/or flower fragrance.
REPRODUCTIVE SYSTEMS. The majority of species
in the family are certainly outbreeders. Self-
fertility has been found in Tillandsia subgenus
Tillandsia, and facultative autogamy in Tillandsia
subgenus Diaphoranthema (Till 1992). Vriesea
 Bromeliaceae
and Werauhia species are capable of inbreeding
and numerous Guzmania species are self-
pollinated. In the Bromelioideae, Acanthostachys
pitcairnioides, Aechmea bromeliifolia, and
Ronnbergia nidularioides are examples of self-
pollinated species, whereas Ae. fulgens and Ae.
nudicaulis appear not to be. Most of the species
of Billbergia subgenus Helicodea seem to be
inbreeders, whereas subgenus Billbergia, the
more primitive group, is primarily outcrossing.
Fosterella penduliflora, Pitcairnia andreana, P.
xanthocalyx, and Puya mirabilis are a few
examples of inbreeders in Pitcairnioideae.
Natural hybrids appear to be rather rare under
field conditions (Harms 1930) but have been
found between several species of Tillandsia
subgenus Tillandsia (Gardner 1982). The great
abundance of artificial hybrids produced by
horticulturists attests to the fact that genetic in-
compatibility systems are of minor importance in
isolating species under natural conditions. Differ-
ent phenologies and tight adaptation to pollina-
tors seem to be the most effective barriers against
hybridization in the field (Araujo et al. 1994;
Martinelli 1994).
FRUIT AND SEED. Detailed modern studies in the
seed morphology have been made by Nemirovich-
Danchenko (1983) and GroB (1988a). In the
Pitcairnioideae the fruit is generally a septicidal
capsule with an additionalloculicidal apical dehis-
cence in some genera. The pericarp is not differen-
tiated into exocarp and endocarp. Seeds are
usually small and appendaged (Fig. 24H), rarely
naked (Navia, Pepinia, Varadarajan 1986), the ap-
pendage being a completely surrounding wing or
the seeds are bicaudate. Tillandsioideae have cap-
sular fruits with the pericarp splitting into a
stramineous exocarp and a dark lustrous
endocarp and have plumose seeds (Figs. 25G,
26G). In Glomeropitcairnia the outer integument
forms plumose appendages at both ends, which
suggests an intermediate position between
Pitcairnioideae and Tillandsioideae. In Catopsis
single hairs grow from the chalaza to form a folded
coma. In the remaining genera of Tillandsioideae,
the outer integument splits up into rows of elon-
gated cells forming the pseudopappus (Fig. 26G).
The seed coat formed by the inner integument
usually consists of 2 cell layers and seems to have
a characteristic structure for individual genera
(Smith and Downs 1974). In the Bromelioideae the
fruits are berries with a variously fleshy pericarp,
the seeds unappendaged and often with a
sarcotesta; Acanthostachys is a remarkable excep-
81
tion with seeds that have strongly elongated cha-
lazal appendages.
Modern information about germination is
found in GroB (1988b). In Pitcairnioideae and
Bromelioideae, the primary root first appears with
the formation of a crown of root hairs, but never
in Tillandsioideae, where the formation of a crown
of root hairs is rare. In the epigeal Pitcairnioideae
the cotyledon moves out of the seed and becomes
green while it remains in the seed in the two other
subgenera which are both hypogeal.
DISPERSAL. Wind-borne seeds prevail in Pit-
cairnioideae and are obligatory for Tilland-
sioideae, both with capsular fruits. Seeds of
Pitcairnioideae are less adapted to anemochory
and are bicaudate or winged at most. The trans-
port of the naked seeds of members of this sub-
family, e.g., Navia, is unknown; perhaps rain
serves as an agent. In Tillandsioideae an elaborate
and very effective flight apparatus (pseudo-
pappus) allows dispersal over long distances. The
berry fruits of Bromelioideae are dispersed mainly
by birds and less often by bats (Muller in MOller
1921). Ants appear to disperse seeds of several
bromelioid genera. Dispersal in the vegetative
state is common in Tillandsia usneoides, as it is
used by birds for nest construction.
PHYTOCHEMISTRY. Flavonols, flavones (the latter
as highly methoxylated exudates), triterpenes,
and steroids have been isolated from leaves. The
flavonoid pattern is similar to that of the
Commelinaceae, with which the Bromeliaceae
agree also in the occurrence of silica bodies
(Hegnauer 1963, 1986). This reflects the isolated
position of the family within the monocotyledons
(Williams 1978). Cyanidin is the prevaling antho-
cyanin, proanthocyaninidins seem to be absent.
Flower fragrance includes the triterpenes cycloart-
23-en-3~, 25-diol, citronellol, geraniol, and nerol
(Hegnauer 1963). Cell wall UV fluorescence is due
to the combined occurrence of p-coumaric acid,
ferulic acid, and diferulic acid (Harris and Hartley
1980). Exudated gums contain galactose, xylose,
and arabinose. Ananas comosus contains brome-
lain, a mixture of at least 5 proteolytic enzymes;
the odor of its fruit is mainly caused by
undecatrien and undecatraen (Carle 1992).
SUBDIVISION AND RELATIONSHIPS WITHIN THE
FAMILY. Bentham and Hooker (1883) have been
the first to propose a modern subdivision with
three tribes which have been widely accepted as
natural units. Pitcairnioideae were soon further
82 Bromeliaceae
divided. Harms (1930) first recognized subfami-
lies, including the fourth and newly erected Navi-
oideae, to which he attributed a primitive status.
The peculiarities of Navia were confirmed and
extended on the genus Cottendorfia by Robinson
(1969). The three-partition of Bentham and
Hooker ( 1883) was accepted at subfamiliallevel by
most researchers up to the most recent mono-
graphs (Smith and Downs 1974, 1977, 1979).
Generally, the terrestrial Pitcairnioideae are
considered as the least specialized group with
epigeal germination, most primitive trichomes,
predominantly superior ovaries, the capsular
fruits, and the weakly elaborated seeds.
Tillandsioideae with hypogeal germination, most
highly elaborated trichomes, prevailing superior
ovaries, capsules, plumose seeds, and the atmo-
spheric growth habit, as well as Bromelioideae
with hypogeal germination, moderately elabo-
rated trichomes, inferior ovaries, and baccate
fruits, are interpreted as derived lineages. Both
Pittendrigh (1948) and Tomlinson (1969) have
emphasized that the three subfamilies are best re-
garded as separate derivatives of a common ances-
tral group, so that distinct evolutionary trends
must be sought within each subfamily. A mole-
cular analysis has indicated the monophyly of
Tillandsioideae and Bromelioideae and the
sister group relationship of Bromelioideae and
Pitcairnioideae, the latter supposedly being
paraphyletic (Ranker et al. 1990). A cladistic study
of Varadarajan and Gilmartin (1988a) led to the
recognition of three monophyletic tribes within
Pitcairnioideae: Brocchinieae, Pitcairnieae, and
Puyeae (Varadarajan and Gilmartin 1988b). The
relationships among the genera of Bromelioideae
are insufficiently understood and the arrangement
presented under Genera of the Bromeliaceae
(below) follows the supposed progressions from
sulcate to porate pollen, from a lacking to a large
epigynous tube, and from appendaged to naked
petals, but remains somewhat arbitrary.
AFFINITIES. The family Bromeliaceae is widely
recognized as monophyletic (Gilmartin and
Brown 1987). Synapomorphies of the family
include stellate and scalelike multicellular hairs;
solitary silica bodies; a conspicuously sepaloid
and petaloid perianth; conduplicate-spiral stig-
mas; and possibly the chromosome base number
of x = 25. Molecular analyses support a concept
of a close relationship between the Bromeliifiorae,
Commelinifiorae, and Zingiberifiorae, which form
a monophylum to which the palms are basal (Clark
et al. 1993). At the family level, rbcL data indicate
that Bromeliaceae are closely related to Rapatea-
ceae and Mayacaceae (Chase et al. 1993; Clark et al.
1993, Givnish et al. 1995), or to Commelinaceae
(Duvall et al. 1993). A relationship to Velloziaceae
as proposed by Dahlgren ( 1983) and Dahlgren et al.
(1985) cannot be confirmed. Takhtajan (1980) is
alone in his suspicion of a relationship with the
agavaceous stock ofLiliales.
DISTRIBUTION, HABITATS, AND THE EVOLUTION
OF EPIPHYTISM. Bromeliaceae are native to the
tropical and subtropical regions of the neotropics
extending to Virginia in N America and to north-
ern Patagonia in S America. They range from sea
level to over 4500 m in the central Peruvian Andes.
Only Pitcairnia feliciana is endemic to western
tropical Africa; its occurrence there is interpreted
by long-distance dispersal. This pattern of distri-
bution is equalled by Rapateaceae, with Marsch-
alocephalus as the only African member. A few
Bromeliaceae live far off the South American sub-
continent on islands: Racinaea insularis on the
Galapagos Islands, Greigia berteroi and Ochagavia
elegans on the Juan Fernandez Islands.
The Brocchinieae are restricted to the Guayana
Highlands, Pitcairnieae have their center in north-
ern S America extending into Mesoamerica, the
Antilles, and southeastern Brazil, and Puyeae have
one center along the Andean cordillera and
another in eastern Brazil; only Brewcaria and a
few Puya species are represented in the Guayana
Highlands. Hechtia is remarkable for its restric-
tion to Mexico and Guatemala. Tillandsioideae
have their center in northwestern S America ex-
tending into Mesoamerica, the Antilles, south-
western S America, and southeastern Brazil,
to which latter area Alcantarea is restricted.
Bromelioideae are centered in eastern Brazil.
Some genera exhibit long-distance disjunctions
like Greigia, Billbergia, some subgenera of
Aechmea, Hohenbergia, and Neoregelia, or are
wide-ranging like some subgenera of Aechmea
and Bromelia. For detailed distribution maps at
generic and subgeneric levels see Smith and
Downs (1974, 1977, 1979).
Bromeliads are found in a great variety of habi-
tats. Mesophytes prevail in environments with
high humidity like lowland rainforests, montane
forests, and cloud forests, while CAM xerophytes
are able to settle in dry areas like savannahs,
campos rupestres, semideserts, fog deserts and
even on bare rock. For more details see Benzing
(1980, 1990) and Rauh (1990).
The evolution of epiphytism has been explained
controversely. Schimper (1888) concluded that
 Bromeliaceae
evolution of the epiphytic life form had occurred
within the rainforest itself as proceeding from the
forest floor to the lower limbs, and then ultimately
to the relatively dry upper canopy of the forest as
certain bromeliads became more and more inde-
pendent of roots through increasing specialization
of the adsorbing trichomes. Pittendrigh (1948)
believed in a moderate xeric ancestor which gave
rise to both fully xeric and mesic derivatives.
Tomlinson (1969), however, pointed out that
the anatomical evidence may be in conflict with
Pittendrigh's evolutionary interpretation. Medina
(1974) proposed an intermediary position of a
heliophilic C3 ancestor between completely meso-
phytic and xerophytic derivatives. However, the
disucssion has not yet come to an end (Benzing
et al. 1985).
AssociATED BIOTA. An extensive literature has
developed on the animals which live in bromeliad
tanks and leaf bases (Kriigel1993). The spectrum
of phytotelm inhabitants comprises animals and
prokaryota, algae, myxomycota, and higher plants
(Laessle 1961), and ranges from occasionals to
organisms restricted to these microhabitats like
the crustacea Elpidium bromeliarum or the frog
Syncope antenori. Many malaria mosquito species
use the bromeliad ~anks during their life cycles.
FossiLS. Six different specimens of fossils have
been assigned tentatively to the Bromeliaceae.
According to the evaluation by Smith and Downs
(1974), for none of them is the association with the
family proven, or at least likely. The probably
only true bromeliad fossil is Karatophyllum
bromelioides from Costa Rica (Gomez P. 1972).
EcoNOMIC IMPORTANCE. Ananas comosus, the
pineapple, is the species of primary economic im-
portance and is cultivated for its fruits worldwide
in tropical regions. It also provides the proteolytic
enzyme bromelain which is pharmaceutically
used (Carle 1992). Tillandsia usneoides, Spanish
moss, was formerly important, especially in Loui-
siana, as a squrce of pillow and cushion stuffing,
but has been supplanted by plastic. Neoglaziovia
variegata, the car6a or carua, and Ananas lucid us,
the curagua, are a source of heavy fiber in north-
ern S America. Some Bromelia species have edible
fruits (Arenas and Arroyo 1988; Correa and Bernal
1990). Many bromeliads are cultivated as
ornamentals.
CoNSERVATION. Seven species of Tillandsia have
been protected by appendix II of CITES. Many
83
more may be endangered by commercial collect-
ing on a large scale, especially in Guatemala and
Brazil. A main source of endangering, however, is
the destruction of the natural habitats by unscru-
pulous land use.
CONSPECTUS OF CLASSIFICATION OF
BROMELIACEAE
I. Subfamily Pitcairnioideae Harms (1930).
1. Tribe Brocchinieae G.S. Varad. & Gilmartin (1988).
Genus 1.
2. Tribe Pitcairnieae K. Koch (1874).
Genera 2-10.
3. Tribe Puyeae Wittm. (1888).
Genera 11-16.
II. Subfamily Tillandsioideae Harms (1930).
Genera 17-25.
Ill. Subfamily Bromelioideae Harms (1930).
Genera 26-56.
KEY TO THE GENERA
1. Mature seed appendaged or if not (Pitcairnia in part and
Navia and Brewcaria), then the fruit capsular and dehis-
cent; ovary superior or largely so in most genera to infe-
rior; fruit capsular or if not (Pitcairnia in part), then the
seed appendaged; germination epigeal or hypogeal 2
- Mature seed unappendaged; fruit baccate; ovary inferior;
leaves mostly spinose-serrate; indument almost always of
obvious scales; germination hypogeal; plants often terres-
trial (subfam. Bromelioideae) 27
2. Seed appendages entire or slightly divided (Brocchinia
paniculata) or lacking (Pepinia aphelandriflora) and
Navia); fruit usually dehiscent; leaves mostly spinose-
serrate; indument of finely to scarcely divided scales;
germination epigeal; plants usually terrestrial (subfam.
Pitcairnioideae) 3
- Seed appendages finely divided and forming a coma,
always present; fruit dehiscent; leaves always entire;
indument almost always of obviously radially symmetric
scales; germination hypogeal; plants mostly epiphytic
(subfam. Tillandsioideae) 19
3. Plants monoecious or, if rarely dioecious (Cottendorfia) or
polygamodioecious (Dyckia maritima, D. selloa, and D.
hebdingii), then the petals yellow or orange and plants
native to northeastern and southernmost Brazil 4
- Plants dioecious with functionally unisexual flowers;
petals rose or white; plants of Texas, Mexico, and northern
Central America 16. Hechtia
4. Bases of the filaments separate from each other, but some-
times individually adnate to the petals and sepals 5
- Bases of the filaments forming a tube and adnate to the
petals; petals yellow to orange; plants of Brazil, Uruguay,
Paraguay, and Argentina 15. Dyckia
5. Seeds obviously and persistently appendaged 6
- Seeds naked at maturity 17
6. Petal blades tightly spiraled after anthesis, broad, distinct
from the claw; leaf blades narrowly triangular, never con-
tracted at base; ovary superior or slightly inferior; Andean
plants of open slopes and summits from Costa Rica and
Guayana to Chile and Argentina 13. Puya
- Petal blades remaining separate after anthesis or, if
sometimes slightly spiraled (Deuterocohnia), then not
clawed 7
84 Bromeliaceae
7. Ovary wholly superior; petals regular 8 19. Ovary nearly or quite superior; seeds plumose on base or
- Ovary partially to wholly inferior, or, if superior, then the
petals zygomorphic (Pepinia and Pitcairnia spp.) 14
8. Petals naked or with paired appendages (Steyerbromelia)
9 appendaged at both ends; flowers polystichous. Lesser
- Petals each bearing a single basal scale; xerophytic plants
of the southern Andes from Peru to Chile, Argentina, and
W Brazil 12. Deuterocohnia
9. Seeds with a falciform appendage; petal blades narrow,
indistinct; plants of NE Brazil 14. Encholirium
- Seeds bicaudate-appendaged 10
10. Anthers basifixed, linear, coiled at anthesis, inner fila-
ments adnate to the base of the petals; leaf blades thin,
more or less contracted at base; mesophytic plants of
Mexico to Argentina and W Brazil 8. Fosterella
- Anthers subbasifixed to equitant, stout, straight; filaments
usually free; leaf blades firm, not contracted at base; plants
of the Guayana Highlands 11
11. Flowers perfect; sepals convolute, each with its left side
covering the right side of the next one 12
- Flowers unisexual, dioecious; sepals cochlear with the
abaxial overlapping both the adaxial, plants of NE Brazil
6. Cottendorfia
12. Petals naked; stigmas straight, erect; flowers mostly
pedicellate 13
- Petals with paired appendages; stigmas broad, strongly
contorted; flowers sessile; inflorescence compound, lax
4. Steyerbromelia
13. Anthers subbasifixed; petals brightly colored, more or less
massed together after anthesis but not twisted; sepals large
and firm 7. Connellia
- Anthers equitant; petals white or rose, separate after
anthesis; sepals not over lOmm long, thin, flat
5. Lindmania
14. Petals large, naked, or appendaged, usually zygomorphic
and forming a hood over the anthers; sepals convolute
with the left side of each overlapping the right of the next
one; plants of Mexico and the West Indies to Argentina
and Brazil 15
- Petals minute, regular; sepals cochlear with both adaxial
ones overlapping the abaxial; plants of the Guayana
Highlands 16
15. Seeds 2-tailed; foliage usually dimorphic to polymorphic;
inflorescence a raceme, panicle, spike, or frequently
strobiliform; flowers and floral bracts occasionally secund
and reflexed; leaf blades often broadly elliptic, ovate, lan-
ceolate, or oblanceolate 10. Pitcairnia
- Seeds winged; foliage monomorphic to rarely dimorphic,
never polymorphic; inflorescence a raceme or panicle,
never strobiliform; flowers and floral bracts always erect to
spreading; leaf blades narrowly elliptic to lanceolate
9. Pepinia
16. Epigynous tube lacking; inflorescence open and definitely
branched 1. Brocchinia
- Epigynous tube well developed; inflorescence sessile,
capitate 3. Ayensua
17. Sepals convolute; petals zygomorphic (Pepinia
aphelandriflora) 15
- Sepals cochlear with both adaxial ones overlapping
the abaxial; petals regular; ovary superior (in most
species) to nearly inferior; plants of the Guayana
Highlands 18
18. Petals naked; inflorescence scapose, pinnate, and more or
less open or sessile and capitate 2. Navia
- Petals with 2 appendages; inflorescence long-scapose,
simple, densely cylindric 11. Brewcaria
apex or largely on the base and only slightly on the apex
20
- Ovary only half superior; seeds equally plumose-
Antilles, Trinidad, adjacent Venezuela
18. Glomeropitcairnia
20. Appendage of the seed wholly or largely basal, straight at
maturity 21
- Appendage of the seed largely apical ( = chalaza!), folded
at maturity; sepals strongly asymmetric in most species;
flowers in at least slightly more than 2 ranks; leaves often
cretaceous-coated on the inside. Florida, Mexico, and the
West Indies to Brazil and Peru 17. Catopsis
21. Petal claws free or with very short tube exceeded by the
sepals; flowers distichous in most species 22
- Petal claws conglutinated in a tube, equaling the sepals or,
rarely, the petals entirely included in the sepals 26
22. Petals naked; inflorescence of 1 or more distichous-
flowered spikes or racemes or rarely reduced to 1 or more
polystichous-flowered spikes or to a single flower; plants
of southern United States to Argentina and Chile 25
- Petals bearing scales ("lateral folds" of Mez) on the inside
of the claw; inflorescence as in Tillandsia. Mexico and the
West Indies to Argentina and Uruguay 23
23. Seed with the apical appendage divided into a short coma;
petals linear long, fusiform, usually 10-15 times longer
than wide, soon flaccid and drooping 21. Alcantarea
- Seed with the apical appendage minute and undivided;
petals elliptical, usually 5-10 times longer than wide, usu-
ally firm and remaining more or less erect after an thesis
24
24. Flowers with brilliant coloration in most species, bright
yellow, orange, or red, rarely dull to white, light yellow, or
light orange; the adaxial petal pair arranged apically in
respect to the abaxial; petal appendages tongue-shaped;
stigma with the convolute blade type morphology, that is,
3 obviously spreading lobes covered with papillae
22. Vriesea
- Flowers generally dull in color, white, greenish white, light
green yellowish green, yellow, or light orange; the adaxial
petal pair arranged basally in respect to the abaxial;
petal appendages dactyloid with 1-5 fingers of varying
length; stigma with the cupulate type morphology, that is,
3 apical, capitate, cup-shaped lobes, without papillae
23. Werauhia
25. Sepals symmetric or if slightly asymmetric, then ovate
or lanceolate and broadest below the middle, free or
variously connate; seeds usually with a distinct apical
appendage 24. Tillandsia
- Sepals asymmetric, free or nearly so, broadest near apex,
not over 12mm long; seeds without apical appendage
25. Racinaea
26. Petal claws always naked; spikes always polystichous-
flowered. Florida, Mexico, and the West Indies to Brazil
and Bolivia 20. Guzmania
- Petal claws bearing scales on the inside; flowers polystic-
hous rarely secund or distichous. Colombia to Peru
19. Mezobromelia
27. Sepals symmetric or nearly so 28
- Sepals asymmetric 51
28. Filaments forming a tube to which the fleshy petals are
joined along their centers but with their margins free;
sepals mostly free or nearly so; leaves very laxly and
coarsely spinose-serrate 29
- Filaments not connate but sometimes adnate 31
 Bromeliaceae 85

29. Sepals with soft, usually broad apices; inflorescences 49. Petals connate or agglutinated in a tube the height of the
compound. Mexico and the West Indies to Argentina and sepals 52. Nidularium
Uruguay 36. Bromelia - Petals free or nearly so 50
- Sepals spinose-mucronate 30 50. Petals completely free; inflorescence often scapose
30. Inflorescence simple, subsessile. Argentina 45. Canistrum
37. Deinacanthon - Petals short-connate at or a little above the base
- Inflorescence branched with terminal strobils, scapose. S 46. Wittrockia
Mexico, Guatemala 49. Hohenbergiopsis 51. Ovaries coalescing to form a compound fruit; inflores-
31. Terminal axes of the inflorescence visible 32 cence simple, strobilate 52
- Terminal axes of the inflorescence covered by leaves or - Ovaries always remaining distinct 53
bracts 38 52. Inflorescence with a small, inconspicuous coma, never
32. Petals naked; sepals 0.5-7mm long 33 producing basal shoots; plant propagating by elongate rhi-
- Petals appendaged; sepals mostly much larger 34 zomes; petals bearing vertical folds. Paraguay and adjacent
33. Inflorescence compound; sepals broadly ovate or oblong, areas 39. Pseudananas
0.5-2mm long. Costa Rica and Trinidad to Amazonian - Inflorescence usually with a large conspicuous coma (lack-
Brazil 28. Araeococcus ing in A. monstrosus), often with basal shoots; rhizomes
- Inflorescence simple; sepals narrowly elliptic, 7mm long; lacking; petals usually bearing well-developed scales.
flowers subsessile or pedicellate. Mount Itatiaia area in E Probably native from Paraguay to the Amazon Basin, now
Brazil 31. Fernseea pantropical 40. Ananas
34. Petals zygomorphic or tightly recoiled and flowers sessile. 53. Flowers pedicellate 54
W Mexico and Central America to Argentina and Uruguay - Flowers sessile or subsessile 59
35 54. Inflorescence nidular, simple in most species; petals
- Petals regular (actinomorphic) 36 naked. Amazonia, E Brazil 53. Neoregelia
35. Epigynous tube usually well developed 34. Billbergia - Inflorescence scapose 55
- Epigynous tube shallow. W Mexico 33. Ursulaea 55. Sepals more or less connate, long-mucronate; petals
36. Petals erect. E Brazil 37 appendaged. E Brazil 47. Portea
- Petals apically recoiled 33. Ursulaea - Sepals free or unarmed 56
37. Flowers sessile 44. Quesnelia 56. Inflorescence simple; sepals unarmed 57
- Flowers pedicellate 35. Neoglaziovia - Inflorescence compound 58
38. Inflorescence simple, strobilate; flowers solitary in the axil 57. Petals naked. Colombia 48. Pseudaechmea
of each bract 39 - Petals appendaged. Colombia and Guyana to NE Brazil
- Inflorescence compound 46 43. Aechmea subg. 2. Lamprococcus
39. Scape short or none; strobils nidular or axillary 40 58. Sepals 1.5-3mm long; inflorescence glabrous; petals
- Scape well developed, obvious 44 naked. Colombia to Suriname and Amazonian Brazil
40. Floral bracts foliaceous. NE Brazil 26. Orthophytum 28. Araeococcus
- Floral bracts bracteiform 41 - Sepals 3.5-22mm long; inflorescence !epidote; petals
41. Scape distinct, its bracts shorter than the floral bracts; appendaged. Mexico to Peru
petals naked. Mexico and Venezuela to Chile 43. Aechmea subg. 1. Podaechmea
30. Greigia 59. Petals appendaged with well-developed scales 60
- Scape none or very short 42 - Petals naked or with lateral folds or rudimentary or
42. Epigynous tube shallow, crateriform (A. pitcairnioides). reduced scales 66
Brazil: Bahia 41. Acanthostachys 60. Epigynous tube shallow or lacking; flowers in terete
- Epigynous tube cylindric, deep. Chile 43 strobils; inflorescence mostly pinnate and lax, rarely digi-
43. Sepals obtuse, carinate 29. Fascicularia tate or simple (H. littoralis). Antilles to Venezuela and
- Sepals acute to attenuate, pungent to mucronate, not Brazil 42. Hohenbergia
carinate 32. Ochagavia Epigynous tube well developed; inflorescence various 61
44. Scape erect, without bracts (A. strobilacea). S Brazil, 61. Sepals unarmed 62
Paraguay, Argentina 41. Acanthostachys - Sepals armed. N and S America 43. Aechmea subg.
- Scape covered with bracts 45 3. Aechmea, 43. Aechmea subg. 4. Ortgiesia,
45. Scape bracts foliaceous, scape erect. NE Brazil and 43. Aechmea subg. 6. Pothuava
26. Orthophytum 62. Floral bracts attached basally, not decurrent nor forming
- Scape bracts bracteiform; scape prostrate. French Guiana pouches; flowers polystichous 63
and adjacent Brazil 56. Disteganthus - Floral bracts decurrent and forming pouches around the
46. Inflorescence obviously compound with several strobils flowers; flowers often distichous. N and S America
on an elongate floral axis 47 43. Aechmea subg. 5. Platyaechmea
- Inflorescence pseudosimple with hands or fiat fascicles in 63. Inflorescence branched 64
the axils of large bracts 48 - Inflorescence simple 65
47. Floral bracts foliaceous, serrulate; strobils sessile or 64. Leaves 2-ranked; blades marked with spots or bands; floral
subsessile. NE Brazil 26. Orthophytum bracts minute; ovules obtuse (Qu. marmorata). Brazil:
- Floral bracts bracteiform, entire; strobils on Espirito Santo to Sao Paulo 44. Quesnelia
distinct scapes. Mexico and Venezuela to Chile - Leaves more than 2-ranked or the blades concolorous;
30. Greigia floral bracts large to lacking; ovules long-caudate. Colom-
48. Outer bracts of the inflorescence foliaceous; sepals high bia, Venezuela, Amazonian Brazil 43. Aechmea subg.
connate; petals naked. NE Brazil 27. Cryptanthus 2. Lamprococcus
- Outer bracts of the inflorescence bracteiform, large, and 65. Ovules obtuse (no further distinction possible without
covering most of the flowers. E Brazil 49 keying by species). E Brazil 44. Quesnelia
86 Bromeliaceae
- Ovules apiculate to caudate. Central America to Brazil and
Argentina 43. Aechmea subg. 7. Macrochordion
66. Ovary deeply sulcate; inflorescence simple or compound.
NE Brazil 51. Lymania
- Ovary evenly rounded 67
67. Inflorescence lax; axes visible 68
- Inflorescence dense 71
68. Inflorescence simple. Costa Rica to Peru 55. Ronnbergia
- Inflorescence pinnately compound 69
69. Flowers minute; sepals not over 3mm long; ovules few;
epigynous tube none. Costa Rica, Venezuela, Trinidad,
Tobago, Guyana to Amazonian Brazil 28. Araeococcus
- Flowers small to large; sepals more than 3 mm long; epigy-
nous tube distinct 70
70. Branches elongate, many-flowered; flowers perfect;
anthers unappendaged. E and Amazonian Brazil and
adjacent areas 54. Streptocalyx
- Branches short, digitately few-flowered; flowers function-
ally unisexual on different plants; anthers appendaged.
Central America: Guatemala to Costa Rica 50. Androlepis
71. Flowers 2 or more in the axil of each bract 72
- Flower single in the axil of each bract 73
72. Inflorescence involucrate; sepals only slightly asymmetric,
unarmed or mucronulate. E Brazil 52. Nidularium
- Inflorescence strobilate; sepals strongly asymmetric,
mucronate. E and Amazonian Brazil and adjacent areas
54. Streptocalyx
73. Petals naked or with lateral folds; bracts chartaceous or
coriaceous; leaf blades often petiolate. Costa Rica to Peru
55. Ronnbergia
- Petals bearing rudimentary or reduced appendages; bracts
mostly thick and ligneous; leaf blades never petiolate;
pollen sulcate. Mexico to Peru and Amazonian Brazil, E
Brazil 38. Chevaliera
GENERA OF THE BROMELIACEAE
I. Subfam. Pitcairnioideae Harms (1930).
subfam. Navioideae Harms (1930).
Nearly always terrestrial or saxicolous. Roots
present and largely functional. Leaves nearly
always spinose-serrate. Ovary superior or
largely so to rarely inferior (in Ayensua, many
Brocchinia spp., and a few species of Navia
and Pitcairnia). Pollen sulcate, sulcus simple
except of Pepinia and Pitcairnia with a distal
germination area. Stigma of the conduplicate-
spiral type morphology except in Brewcaria,
Cottendorfia, Hechtia, Brocchinia (in part), and
Fosterella (in part). Fruit capsular or rarely
indehiscent.
1. Tribe Brocchinieae G.S. Varad. & Gilmartin
(1988).
Seeds elongate, apical and basal ends dissimilar,
basal appendage variously modified, surface cellu-
lar pattern of seed appendage and seed body alike;
foliar trichomes symmetric with stalk of six or
more cells; leaf photoassimilatory tissue with
abundant stellate chlorenchyma; leaves almost
always entire (except Brocchinia serrata L.B. Sm.);
inflorescence a raceme or profusely branched
panicle.
1. Brocchinia Schult. & Schult. f.
Brocchinia Schult. & Schult. f., in Roem. & Schult., Syst. veg.
7(2): lxx, 1250 (1830); L.B. Sm., Ann. Mo. Bot. Gard. 73:689-
721 (1986).
Terrestrial or rarely epiphytic, stemless to mas-
sively caulescent; leaves rosulate; blades usually
entire, glabrous or subglabrous; scape conspic-
uous; inflorescence compound, mostly ample;
flowers perfect, minute, pedicellate; sepals free,
imbricate with the 2 adaxial covering the abaxial;
petals free, naked; stamens included at anthesis;
filaments more or less adnate to the sepals and
petals; stigma conduplicate-spiral or simple erect;
ovary from 1/3 to wholly inferior; placentae linear;
ovules few, long-appendaged; capsule wholly
septicidal or rarely loculicidal as well (B. ser-
rata L.B. Sm.). Twenty one spp., Guayana
Highlands.
2. Tribe Pitcairnieae K. Koch (1874).
Seeds usually spherical, lenticular, cuneate, po-
lygonal, reniform, or ellipsoidal; if elongate, then
apical and basal ends similar, surface cellular pat-
tern of seed appendage and body often different;
foliar trichomes symmetric or asymmetric with
a stalk of 2 or 3 cells; leaf photoassimilatory
tissue without stellate chlorenchyma but with
abundant palisade and spongy elements or
undifferenciated chlorenchyma; leaves entire to
mostly spinulose or serrulate, typically mem-
branous, occasionally coriaceous; foliar trichome
arrangement irregular and nonoverlapping; epi-
dermis smooth (except for some Pepinia); water-
storage tissue peripheral only; hypodermal
sclerenchyma absent (except for Ayensua and
some Pepinia); inflorescence a globose head,
spike, raceme, or panicle.
2. Navia Mart. ex Schult. & Schult. f. Fig. 23
Navia Mart. ex Schult. & Schult. f., in Roem. & Schult., Syst.
veg. 7(2): lxv, 1195 (1830); L.B. Smith, Ann. Mo. Bot. Gard.
73: 689-721 (1986); emend. Steyerm., J. Bromeliad Soc. 39:
54 (1989).
Stemless or caulescent; leaves rosulate or in a
dense spiral along the stem; blades entire or more
 Bromeliaceae
Fig. 23A-E. Bromeliaceae-Pitcairnioideae. Navia acaulis. A
Habit. B Flower with subtending bract. C Pistil. D Dehiscing
capsule. E Seed. (Takhtajan 1982)
or less serrulate; scape usually lacking, rarely well
developed; inflorescence usually capitate but
sometimes paniculate; floral bracts conspicuous;
flowers sessile or subsessile, perfect; sepals imbri-
cate with both adaxial ones overlapping the
abaxial, free to high-connate; petals free to high-
connate, naked; stamens of the second series
highly adnate to the petals; anthers linear; ovary
usually superior but rarely to wholly inferior; style
slender; placentae linear; ovules naked or nearly
so; capsule usually dehiscent; seeds reticulate,
naked or soon losing the vestigial wing. Ninety
eight heliophytic spp., Guayana Highlands from
Colombia to Suriname.
3. Ayensua L.B. Sm.
Ayensua L.B. Sm., Mem. N. Y. Bot. Gard. 18 (2): 29 (1969);
emend. L.B. Sm. & Read, Phytologia 38: 137 (1977).
Caulescent, terrestrial and saxicolous in dense
colonies; leaves densely imbricate along the stem;
sheaths distinct; blades attenuate, not contracted
at base, deciduous; scape lacking; inflorescence
capitate; flowers perfect, sessile or subsessile;
87
sepals free, imbricate with the abaxial wholly in-
cluded by the 2 adaxial; petals free, naked; sta-
mens included, the 2 series unequal; ovary wholly
inferior, crowned with a large epigynous tube;
ovules caudate; fruit indehiscent on the outside,
dividing into 3 equal valves inside. Only one sp., A.
uaipanensis (Maguire) L.B. Sm., Auyan-tepui in
Guayana Highlands of southern Venezuela.
4. Steyerbromelia L.B. Sm.
Steyerbromelia L.B. Sm., Acta Bot. Venez. 14: 8 (1984), nom.
illeg.; Ann. Mo. Bot. Gard. 73: 689-721 (1986).
Large heliophytic herbs; leaves rosulate; sheaths
short; blades narrowly triangular, not constricted
at base, more or less spinose-serrate; scape devel-
oped, stout; scape bracts subfoliaceous; inflo-
rescence amply pinnate-paniculate, lax; flowers
perfect, sessile; sepals imbricate, free; petals
appendaged with vertically attached scales, free;
stamens equal; ovary superior; stigmas broad,
strongly contorted; ovules caudate. Three spp.,
open summits, Guayana Highlands.
5. Lindmania Mez
Lindmania Mez, in A. & C. DC., Monogr. phan. 9: 535 (1896);
L.B. Sm., Ann. Mo. Bot. Gard. 73: 689-721 (1986).
Cottendorfia sensu LB. Sm. (1960), (1974); in part but not as
to type.
Small to large heliophytic herbs; leaves rosulate;
blades mostly entire but sometimes more or less
serrulate; scape usually but not always developed;
inflorescence compound or rarely simple; flowers
perfect, small, white, or greenish; sepals con-
volute, free; petals free, naked; filaments mostly
free; anthers versatile; stigma simple-erect; ovary
wholly superior, glabrous; style slender; placentae
short, basal; capsule short, ovoid, septicidal; seeds
slenderly fusiform, bicaudate. Thirty eight spp.,
open habitats, Guayana Highlands.
6. Cottendorfia Schult. & Schult. f.
Cottendorfia Schult. & Schult. f., in Roem. & Schult., Syst. veg.
7(2): lxiv, 1193 (1830).
Heliophytic herb; leaves rosulate; blades entire,
pungent; inflorescence tripinnately paniculate,
lax; branches densely polystichous-flowered;
flowers unisexual, dioecious, sessile, small; sepals
cochlear, the abaxial overlapping both the adaxial,
free; petals free, naked; stamens included; fila-
ments free; anthers ovate, mediofixed; ovary
88 Bromeliaceae

superior; style cylindric, short; stigmas 3, linear, 10. Pitcairnia L'Her. Fig. 24A-H
erect; placentae subscutate, basal; ovules few,
Pitcairnia L'Her., Sert. angl.: 7 (1788); nom. cons.; G.S. Varad.
apically short-caudate. Only one sp., C. florida & Gilmartin, Syst. Bot. 13: 294-299 (1988).
Schult. & Schult. f., restricted to xeric habitats in
NE Brazil. Mostly terrestrial herbs but sometimes saxicolous
or epiphytic, mostly stemless to long-caulescent;
leaves polystichous or rarely distichous; blades
7. Connellia N.E. Br.
Connellia N.E. Br., Trans. Linn. Soc. Lond. Bot. 6: 66 (1901);
L.B. Sm., Ann. Mo. Bot. Gard. 73: 689-721 (1986).

Stemless or short-caulescent; leaves rosulate, co-

riaceous; blades spinose-serrate to entire; scape
developed; inflorescence simple or compound;
flowers perfect; sepals convolute, free; petals free,
naked; stamens included; anthers basifixed; ovary
wholly superior; style elongate; capsule septicidal;
seeds long-caudate. Five spp., open slopes and
summits, Guayana Highlands.

8. Fosterella L.B. Sm.

Forsterella L.B. Sm., Phytologia 7: 171, pl. 1 (1960); L.B. Sm. &
Read, Bradea 6: 134-140 (1992).
Lindmania sensu Mez (1896), (1935); sensu L.B. Sm. (1960);
in part but not as to type.

Mesophytic terrestrial herbs, stemless or nearly

so; leaves rosulate; blades entire or weakly
serrulate; scape always well developed; inflores-
cence usually laxly paniculate; flowers small to
minute, perfect; sepals convolute, free; petals free,
unappendaged; stamens barely included to ex-
serted; filaments free or the inner adnate to the
bases of the petals; anthers basifixed, linear; pollen
dry, wind-borne; stigma conduplicate-spiral or
simple-erect; ovary superior; placentae linear the
length of the cell; capsule septicidal, globose to
subpyramidal; seeds narrow, bicaudate. (2n = 50,
100). Sixteen spp., mesophytic habitats, Mexico to
Argentina and W Brazil.

9. Pepinia Brongn. ex Andre

Pepinia Brongn. ex Andre, Ill. Hort. 17: 32, pl. (1870); G.S.
Varad. & Gilmartin, Syst. Bot. 13: 294- 299 (1988).

Foliage monomorphic to dimorphic; leaves nar-

rowly elliptic to lanceolate; inflorescence a raceme
or panicle; flowers and floral bracts erect to
spreading; seeds winged to rarely naked. (2n =
50). Sixty eight species, terrestrial and saxicolous
in open savannas and scrub forests. Guayana
Highlands and adjacent lowlands, N Andes,
Central America.
Fig. 24A-M. Bromeliaceae-Pitcairnioideae. A-H Puya
mirabilis. A Habit. B Leaf. C Stamen. D Pistil. E Flower. F Style
with stigma. G Capsule. H Seed. 1-M Puya berteroiana. I
Habit. J Flower. K Stamen. L Pistil. M Ovary, cross-section.
(Takhtajan 1982)
 Bromeliaceae
from linear to broadly lanceolate and petiolate,
usually strongly dimorphic, entire or serrate,
sometimes deciduous; scape usually developed;
inflorescence simple or compound; flowers
perfect; sepals convolute, free; petals free, usually
zygomorphic, naked or with a single scale or
its remnant at base; stamens elongate, from
slightly shorter to slightly longer than the petals;
anthers linear; ovary from almost wholly sup-
erior to wholly inferior; ovules numerous,
appendaged or rarely obtuse; fruit a capsule or
rarely indehiscent on the outside; seeds bica-
udate or broad and winged. (2n = 50). Two hun-
dred and eighty five spp., highly varied habitats.
Mexico and the West Indies to Argentina and
Brazil.
3. Tribe Puyeae Wittm. (1888)
Seeds usually spherical, lenticular, cuneate,
polygonal, reniform, or ellipsoidal, if elongate,
then apical and basal ends similar, surface cell-
ular pattern of seed appendage and body often
different; foliar trichomes symmetric or asym-
metric with a stalk of 2 or 3 cells; leaf photo-
assimilatory tissue without stellate chlorenchyma
but with abundant palisade and spongy ele-
ments or undifferentiated chlorenchyma; leaves
strongly spinose-serrate, succulent or semisuc-
culent; foliar trichomes in epidermal grooves
arranged in overlapping tiers; epidermis with
ridges and furrows; water-storage tissue periph-
eral and internal; hypodermal sclerenchyma
present.
11. Brewcaria L.B. Sm., Steyerm. & H. Rob.
Brewcaria L.B. Sm., Steyerm. & H. Rob., Acta Bot. Venez. 14:
10 (1984); L.B. Sm., Ann. Mo. Bot. Gard. 73: 689-721
(1986).
Large heliophytic herbs; leaves rosulate; blades
narrowly triangular, spinose-serrate; scape erect,
stout; inflorescence spicate, dense; flowers perfect;
sepals imbricate; petals bearing 2 horizontally at-
tached scales, free; stamens equal, included;
stigma simple-erect; ovary slightly inferior; ovules
exalate or very narrowly alate. Two spp., Guayana
Highlands.
12. Deuterocohnia Mez
Deuterocohnia Mez, in Mart., Fl. bras. 3(3): 506 (1894); M.A.
Spencer & L.B. Sm., Bradea 6: 141-146 (1992).
Abromeitiella Mez (1927).
89
Plants growing in ring formations or forming
cushions or polsters; leaves densely rosulate,
spinose-serrate; scape developed and the inflores-
cence perennial, or scape lacking and the inflores-
cence sessile; flowers perfect; sepals convolute,
free; petals free, each bearing a single basal ap-
pendage; stamens free; ovary superior, ovules ob-
tuse to very short caudate; seeds fusiform,
bicaudate. (2n = 50). Fourteen spp., extreme xero-
phytic habitats, southern Andes, Peru, Bolivia,
Argentina, and Chile.
13. Puya Molina Fig. 241-N
Puya Molina, Sag. stor. nat. Chili: 160, 351 (1782).
Stemless or more generally long-caulescent; leaves
coriaceous; blades never constricted at base,
coarsely spinose-serrate; inflorescence simple or
paniculate; flowers perfect; sepals convolute, free;
petals free, unappendaged or with a pair of vertical
folds, twisted together in a spiral after anthesis;
stamens generally somewhat shorter than the pet-
als; filaments free; ovary superior or slightly infe-
rior; style long and slender; capsule loculicidal and
generally tardily septicidal as well; seeds large,
bearing a dorso-apical wing. (2n = 50). One hun-
dred and ninety spp., open slopes and summits
from Costa Rica and Guyana to Chile and Argen-
tina. Divisible into two subgenera, Puyopsis and
Puya, according to absence or presence of sterile
branch ends serving as perches for pollinating
birds.
14. Encholirium Mart. ex Schult. & Schult f.
Encholirium Mart. ex Schult. & Schult. f., in Roem. & Schult.,
Syst. veg. 7(2): lxviii, 1233 (1830); L.B. Sm. & Read, Bradea 5:
296-311 (1989).
Coarse xerophytes; leaves spinose-serrate; scape
well developed; inflorescence usually simple or
rarely few-branched, of many pedicellate flowers;
flowers perfect; sepals convolute or imbricate, free;
petals convolute, free, naked; stamens about equal-
ling the petals to much exceeding them; filaments
free; ovary wholly superior; capsule short, stout;
seeds each with a single dorso-apical wing much
extended backwards. Twenty nine spp., xerophytic
habitats, northeastern Brazil to Mato Grosso.
15. Dyckia Schult. & Schult. f.
Dyckia Schult. & Schult. f., in Roem. & Schult., Syst. veg. 7(2):
lxv, 1194 (1830).
Prionophyllum K. Koch (1874).
90 Bromeliaceae

Xerophytes, often repent; leaves usually spinose-

serrate, rigid; scape usually lateral; inflorescence
amply paniculate to simple; flowers perfect and
uniform to rarely dimorphic with some function-
ally unisexual; sepals imbricate, usually free;
petals imbricate; stamens included to exserted;
filaments basally connate among themselves and
with the petals, free or connate above; ovary
wholly superior; ovules many, each with a single
wing; capsule short and broad, septicidal and
sometimes partially loculicidal as well, seeds nu-
merous, broad, alate. (2n =50, 100). One hundred
and twenty four spp., xeric habitats, Central and E
Brazil, Argentina, Bolivia, Paraguay, and Uruguay.

16. Hechtia Klotzsch

Hechtia Klotzsch, Allg. Gartenzeitung 3: 402 (1835).

Stem short; leaves densely rosulate; blades

coarsely spinose-serrate in most species, rarely
serrulate or even entire; scape usually lateral; in-
florescence paniculate, bipinnate or more divided,
branches from capitate to lax and very long; flow-
ers functionally unisexual on different plants; se-
pals free; petals naked, usually free but sometimes
joined to a ring of filaments by their centers; sta-
mens exserted to included; anthers mostly ovate;
stigma simple-erect; ovary from wholly superior
to 3/4 inferior; capsule septicidal and loculicidal;
seeds many, oblong, narrowly winged or some-
times almost naked, often sculptured. (2n = 50).
Fifty one spp., Texas, Mexico, and northern
Central America.
II. Subfam. Tillandsioideae Harms (1930)
Largely epiphytic, caulescent, or acaulescent.
Roots often functioning only as holdfasts and
sometimes completely lacking. Leaves rosulate or
fasciculate or distributed along stem, entire;
indumentum of radially symmetric scales. Pollen
sulcate, usually with a distal germination area,
rarely inaperturate (Guzmania) or sulcus simple
( Catopsis). Stigma morphology variable, all types
present. Ovary superior or nearly so except
in Glomeropitcairnia. Fruit capsular, septicidal;
seeds with a plumose appendage at base or apex or
both.
17. Catopsis Griseb. Fig. 25
Catopsis Griseb., Nachr. Konig!. Ges. Wiss. Georg Augusts-
Univ. [1] : 10, 12 (Jan 1864).
Fig. 25A-G. Bromeliaceae-Tillandsioideae. Catopsis ber-
teroniana. A Fruiting plant. B Flower with subtending bract. C
Pistil in longitudinal section, showing one row of ovules. D
Cross-section of ovary, showing two rows of ovules in each
locule. E Three ovules showing elongated integument below
and distal tuft of hairs developing into terminal appendage of
seed. F Dehiscing capsule. G Seed with long terminal append-
age of hairs. (Smith and Wood 1975)
Leaves densely utriculose-rosulate, often creta-
ceous-coated; scape conspicuous; inflorescence
usually bipinnate, rarely simple or tripinnate,
the branches polystichous-flowered; flowers per-
fect or dimorphic and functionally dioecious at
least as to the staminate plants (Mexico and
Central America); sepals free, usually strongly
asymmetric with the right side produced;
petals free, naked; stamens included, the 2 series
usually very unequal; anthers ovate or elliptic;
ovary superior; style short or none; stigma
conduplicate-spiral or simple erect; ovules few
to several, apically long-caudate, a small basal
coma of agglutinated hairs appearing as a spur on
the funicle; seeds with a folded apical coma. (2n =
 Brameliaceae
50). Seventeen spp., xeric (canopy) epiphytes,
Florida, Mexico and the West Indies to Brazil and
Peru.
18. Glomeropitcairnia Mez
Glomeropitcairnia Mez, Bull. Herb. Boissier II, 5: 232 (1905);
Gilmartin & al., Syst. Bot. 14:339-348 (1989); Till, Halbritter
& Gortan, J. Brameliad Soc. 47: 65-72 (1997).
Stemless; leaves rosulate, the epidermal scales
with a limb of regularly concentric cells; scape
erect; inflorescence of sessile or short-stipitate
heads or short spikes, bipinnate or tripinnate;
flowers perfect; sepals free; petals free, bearing 2
scales at base; stamens included; stigma papillose,
of the convolute-blade type; ovary about half infe-
rior; capsule imperfectly dehiscent; seeds with
a pappiform coma at both ends. Two spp.,
heliophytic habitats, Lesser Antilles, Trinidad and
adjacent Venezuela.
19. Mezobromelia L.B. Sm.
Mezobromelia L.B. Sm., Proc. Am. Acad. Arts 70: 151, pl. 1,
figs. 10, 11 (1935); H. Luther, Bromelia 2: 3-5 (1995).
Leaves densely rosulate; inflorescence bipinnate;
scape well developed; flowers polystichous or dis-
tichous and deflexed-secund, subsessile or pedi-
cellate, perfect; sepals subfree or connate; petals
tightly conglutinated, bearing 2 scales on the in-
side; filaments adherent to the petals; stigma con-
duplicate-spiral; ovary superior. Nine epiphytic
spp., mesophytic habitats, Greater Antilles to
Peru.
20.. Guzmania Ruiz & Pav.
Guzmania Ruiz & Pav., Fl. peruv. 3: 37, pl. 261 (1802).
Sodiroa Andre (1877).
Stemless or rarely caulescent; leaves densely
polystichous; scape usually conspicuous; inflores-
cence mostly bipinnate or simple, its branches
polystichous-flowered; flowers perfect; sepals free
to high-connate; petals naked, agglutinated in a
tube; stamens equal, more or less agglutinated to
the petals; ovary superior; style elongate; stigma
simple-erect or of the convolute-blade type;
ovules numerous; seeds with a basal, usually
brown coma. (2n = 50). One hundred and sixty
seven species, mainly mesophytic habitats,
Florida, Mexico, and the West Indies to Brazil and
Bolivia.
91
21. Alcantarea (E. Morren ex Mez) Harms
Alcantarea (E. Morren ex Mez) Harms, Notizbl. Bot. Gart.
Berlin-Dahlem 10: 802 (1929); J.R. Grant, Trop. subtrop.
Pflanzenwelt 91: 7-15 (1995).
Leaves forming large rosettes, inconspicuously
!epidote; inflorescence usually paniculate; flowers
distichous, secund, perfect; sepals free, convolute;
petals free, soon becoming flaccid, with 2 entire
appendages; stamens and style exserted, stigma
with the convolute-blade type morphology; ovary
cryptically half-inferior; seeds with both basal and
apical comas. (2n = 50). Twelve spp., mainly
heliophilic epilithes in lowland and montane for-
ests, SE Brazil.
22. Vriesea Lindl.
Vriesea Lind!., Bot. Reg. 29: pl. 10 (1843), nom. cons.; J.R.
Grant, Trap. subtrap. Pflanzenwelt 91: 5-57 (1995).
Acaulescent, usually epiphytic; leaves rosulate;
scape usually conspicuous; inflorescence usually
of distichous-flowered spikes; flowers perfect; se-
pals convolute, free or nearly so; petals
free or connate in a tube much shorter than
the sepals, bearing 2 scales at base, firm and
erect; stamens included or exserted; stigma con-
duplicate-spiral, simple-erect, or of the convolute-
blade type; ovary nearly or quite superior; ovules
many, usually caudate; seeds fusiform with a long
straight basal coma. (2n = 50). One hundred
and ninety-three species in mainly mesophytic
habitats. Mexico and theW est Indies to Argentina
and Uruguay. Divisible into two sections, Vriesea
and Xiphion, according to characters of the
flowers.
23. Werauhia J.R. Grant
Werauhia J.R. Grant, Trap. subtrop. Pflanzenwelt 91: 16-57
(1995).
Usually stemless; leaves rosulate, inconspicuously
!epidote; inflorescence paniculate with often re-
duced branches; flowers usually with nocturnal
anthesis, often secund, sessile to pedicellate, usu-
ally zygomorphic; sepals free, ecarinate; petals
free, with 2 badsal appendages; stamens and style
not exserted from the corolla; stigma cupulate, its
cup-shaped lobes erect and without papillae. (2n
= 50). Sixty six species, mainly epiphytic in mon-
tane and cloud forests, Central and N Andes,
northern S America, Central America, and Greater
Antilles.
92 Bromeliaceae

24. Tillandsia L. Fig. 26 44, 50, 100). Five hundred and forty spp., extreme
xeric to extreme mesophytic habitats, terrestrial
Tillandsia L., Sp. pl.: 286 (1753); W. Till, Sippendifferen-
zierung innerhalb Tillandsia subgenus Diaphoranthema. or frequently epiphytic. Southern United States to
Doct. Diss. Univ. Wien (1984); C.S. Gardner, Selbyana 9: Argentina and Chile. Divisible into six subgenera,
130-146 (1986); Gilmartin & G.K. Br., Am. J. Bot. 73: 387- Allardtia, Anoplophytum, Phytarrhiza, Diaphor-
397 (1986); R.S. Beaman, Systematics of Tillandsia subge- anthema, Tillandsia and Pseudalcantarea accord-
nus Pseudalcantarea. M.Sc. Thesis Univ. Florida (1989). ing to characters of petals, stamens, and styles.
Dendropogon Raf. (1825).
Allardtia A. Dietr. (1852).
25. Racinaea M.A. Spencer & L.B. Sm.
Caulescent or acaulescent; leaves rosulate or dis-
tributed along a stem, polystichous or distichous; Racinaea M.A. Spencer & L.B. Sm., Phytologia 74: 152 (1993).
blades lingulate to narrowly triangular or linear;
scape usually distinct; inflorescence usually of dis- Mainly stemmless with rosulate foliage; inflores-
tichous-flowered spikes, rarely reduced to a single cence usually compound; flowers hermaphrodi-
polystichous-flowered spike or to a single flower; tic, small, actinomorphic, distichously arranged,
flowers perfect; sepals convolute, free or equally or sessile to short pedicellate; sepals convolute,
adaxially joined; petals free, naked; stamens of asymmetric and broadest towards the apex, usu-
various lengths relative to the petals and to the ally free; petals naked, filaments succulent at the
pistil; stigma conduplicate-spiral, simple-erect, base; ovary superior, style subsessile to short,
coralliform, or of the convolute-blade type; ovary stigma papillose, of the simple-erect type; seeds
superior; ovules many, caudate or obtuse; seeds with a badsal appendage only. (2n = 50). Fifty six
with a plumose, white, basal appendage. (2n = 38, mesophytic spp., mainly in montane and cloud
forests, C and N Andes, Mesoamerica, Greater
Antilles, Guayanas, and SE Brazil.

III. Subfam. Bromelioideae Harms ( 1930)

Herbs, largely epiphytic, usually acaulescent.

Roots often functioning as holdfasts. Leaves
rosulate or fasciculate, usually spinose-serrate or -
serrulate; indumentum of peltate scales of irregu-
larly arranged cells or very rarely of multicellular,
cylindric trichomes. Pollen sulcate with a distal
germination area, bi- to poly-porate, or inaper-
turate (Canistrum, some Aechmea). Ovary com-
pletely inferior or nearly so (Acanthostachys);
stigma of the conduplicate-spiral type morphol-
ogy except in Cryptanthus and Orthophytum;
F ovules obtuse to long-caudate. Fruit baccate but
often dry, indehiscent; seeds naked.

26. Orthophytum Beer

Orthophytum Beer, Flora 37: 347 (1854).

Stemless or caulescent; leaves polystichous; blades

Fig. 26A-G. Bromeliaceae-Tillandsioideae. A Tillandsia bar-
tramii, flower, showing tubular corolla and imbricate
sepals. B-G T. utriculata. B Habit. C Style with stigmas. D Part
of inflorescence with three nearly mature fruits. E Ovule, the
apex caudate, the micropyle below (at right). F Nearly mature
seed, oriented as in E. G Mature seed with basal appendage of
hairs, oriented as in E and F. (Smith and Wood 1975)
serrate; scape erect or lacking; scape-bracts folia-
ceous; inflorescence bipinnate from several very
dense heads or rarely simple; primary bracts folia-
ceous or subfoliaceous; floral bracts large, pun-
gent; flowers perfect; sepals free; petals free,
bearing 2 scales; stamens shorter than the petals
and the second series adnate to them; pollen 4-
pored or ellipsoid without distinct pores, reticu-
 Bromeliaceae
late; epigynous tube usually lacking; placentae
central; ovules obtuse. Twenty four spp., saxi-
colous and terrestrial on rocky ground, NE Brazil.
27. Cryptanthus Otto & A. Dietr.
Cryptanthus Otto & A. Dietr., Allg. Gartenzeitung 4: 297
(1836), nom. cons.; non Cryptanthus Osbeck (1757).
Stemless or caulescent; leaves rosulate or evenly
polystichous along the stem; blades narrowly tri-
angular or slightly contracted to subpetiolate at
base, spinose-serrate; inflorescence sessile in the
center of the rosette or at the apex of the stem,
compound; flowers polygamous in some species
at least; sepals much connate; petals connate by
their claws, naked or with two callosities; stamens
exserted from the throat of the corolla, usually
highly adnate to the petals; pollen sulcate; ovules
usually few, obtuse; placentae apical or median;
epigynous tube none or very shallow; fruit small,
rather dry; seeds small, flat. (2n = 34, 54). Forty
one spp., xerophytic, saxicolous, or terrestrial, NE
Brazil.
28. Araeococcus Brongn.
Araeococcus Brongn., Ann. Sci. Nat. Bot. II. 15: 370 (1841).
Stemless; inflorescence laxly paniculate; flowers
perfect; sepals free to connate for half of their
length; petals free, naked; stamens shorter than the
petals; filaments free; anthers longer than the fila-
ments; pollen grains ellipsoid, sulcate; epigynous
tube none; style longer than the petals; stigma
lobes scarcely twisted; ovules few (up to 10), borne
at the top of the locule, long-caudate; seeds nar-
rowly ovoid-cylindric, acute at both ends. (2n =
50). Six spp., mesophytic epiphytes, Costa Rica,
Colombia, Venezuela, Trinidad, Guyana, Brazil.
29. Fascicularia Mez
Fascicularia Mez in Mart., Fl. bras. 3(3): 627 (1891).
Terrestrial or rarely on three trunks, stemless;
leaves very densely rosulate; blades spinose-
serrate; inflorescence sunk in the center of the
rosette, simple; flowers perfect; sepals free; petals
free, bearing 2 small scales near base; epigynous
tube conspicuous; stamens included; filaments
free; anthers linear, elongate, dorsifixed near base;
pollen globose, sulcate, smooth; style elongate;
stigmas not contorted; placentae central to
subapical; ovules rather few, globose, short-
93
appendaged; fruit dry; seeds globose, naked. Five
spp., Chile.
30. Greigia Regel
Greigia Regel, Gartenflora 14: 137 (1865).
More or less caulescent; leaves imbricate along the
stem, partially or completely serrate or serrulate;
scape very short; inflorescence lateral or central,
simple or compound, subglobose; sepals free or
connate; petals connate, usually naked; filaments
adnate to the petals; anthers narrow, acute; pollen
large, spherical, sulcate, smooth; fruit fleshy.
Twenty eight spp., saxicolous in open ground or
terrestrial in forest or paramo, Chile to Mexico
and Venezuela.
31. Fernseea Baker
Fernseea Baker, Handb. Bromel.: 20 (1889); emend. E. Pereira
& Moutinho, Bradea 3: 343, fig. (1983).
Stemless; leaves forming a dense pseudobulb;
blades spinose-serrate; inflorescence simple, race-
mose; flowers perfect; sepals free; petals free,
naked; stamens free, included at anthesis; pollen
sulcate; epigynous tube large; ovules numerous,
unappendaged; fruit succulent; seeds unappend-
aged. Two spp., alpine habitats, Brazil; Mount
Itatiaia region in NW Rio de Janeiro and adjacent
Sao Paulo.
32. Ochagavia Phil.
Ochagavia Phil., Anales Univ. Chile 13: 168 (May 1856).
Suffruticose, amply proliferating; leaves tank-
forming; blades spinose-serrate; inflorescence
sunk in the center of the leaves, simple; flowers
perfect; sepals free; petals free, naked; stamens
usually exserted; anthers subelliptic, dorsifixed;
pollen eporate with a single longitudinal fold; epi-
gynous tube very conspicous; placentae extending
the whole length of the locule; ovules rather nu-
merous, unappendaged; fruit crowned by the per-
sistent sepals; seeds large, globose. Three spp.,
coastal cliffs, Juan Fernandez Islands, Chile.
33. Ursulaea Read & H.U. Baensch
Ursulaea Read & H.U. Baensch, J. Bromeliad Soc. 44: 206
(1994).
Podaechmea (Mez) L.B. Sm. & W.J. Kress (1990) in part, not as
to type.
94 Bromeliaceae

Stemless; leaves rosulate, serrate; scape elongated;

flowers perfect, subactinomorphic, pedicellate;
sepals free, nearly symmetric, mucronate; petals
apically recoiled, with two fimbriate scales at the
base; stamens and style exserted; epigynous tube
shallow; pollen sulcate; ovary inferior, placentae
subapical, ovules caudate. Two spp., epiphytic and
terrestrial, Mexico: Jalisco.

34. Billbergia Thunb. Fig. 27A-F

Billbergia Thunb., Pl. bras. 3: 30 (1821).
Helicodea Lem. (1864).

Stemless; leaves forming broadly funnelform to

long-tubular rosettes; blades serrate or serrulate
or rarely entire; scapebracts large, mostly red; in-
florescence compound or simple; flowers perfect;
sepals free; petals free, actinomorphic or slightly
zygomorphic, bearing 2 scales near base; epigy-
nous tube usually developed; stamens exserted,
both series or the second adnate to the petals only
as high as the scales; pollen sulcate, sometimes
subporate as well (B. elegans Mart. ex Schult. &
Schult. f.); style exceeding the stamens; ovules
many. (2n = 50). Sixty two spp., terrestrial, saxi-
colous, and epiphytic, Mexico and the West Indies
(Bahamas) to Uruguay and Argentina. Divisible
into two subgenera, Billbergia and Helicodea, ac-
cording to the opening of the petals at anthesis,
and somewhat to the form of the inflorescence.

35. Neoglaziovia Mez

Neoglaziovia Mez, in Mart., Fl. bras. 3(3): 426 (1894).

Stemless; leaves few; inflorescence simple, race-

mose; flowers perfect; sepals free, unarmed; petals
free; stamens included; filaments free; pollen sul-
cate; ovules few, short-caudate; fruit succulent.
Three spp., terrestrial and saxicolous in dry open
areas or low thorn forest (caatinga). NE Brazil.

Fig. 27A-K. Bromeliaceae-Bromelioideae. A-F Billbergia

36. Bromelia L.
Bromelia L., Sp. pl.: 285 (1753).
Coarse terrestrials or rarely epiphytes, spreading
by rhizomes; leaves coarsely spinose along the
margins; inflorescence sessile or scapose, com-
pound; sepals free to high-connate, unarmed; pet-
als united into a tube by the filaments but their
margins free, without appendages; stamens in-
cluded, their filaments forming a tube; epigynous
tube conspicuous to nearly lacking; fruit succu-
nutans. A Plant with pollinator. B Flower. C Same, longitudi-
nal section. D Stamen. E Style with stigma. F Base of petal with
scales. G-L Nidularium scherem etiewii. G Habit. H Flower
bud. I Opened ovary and floral tube. J Stigma. K Stamen.
(Takhtajan 1982)
lent, relatively large; seeds few to many, flattened,
naked. (2n = 50, 100). Forty nine spp., mostly in
dry open ground but sometimes submesophytic,
Central and S America.
 Bromeliaceae
37. Deinacanthon Mez
Deinacanthon Mez, in A. & C. DC., Monogr. phan. 9: 12 ( 1896).
Stemless, rhizomatose; leaves squarrose-rosulate;
blades spinose-serrate; inflorescence simple,
sessile; flowers perfect; sepals free, spinose-mucr-
onate; filaments connate with the petals; pollen
sulcate; ovules numerous. Only one sp., D.
urbanianum (Mez) Mez, dry open woodland to
rocky crests, Paraguay, NW Argentina.
38. Chevaliera Gaudich. ex Beer
Chevaliera Gaudich. ex Beer, Pam. Bromel. 22, 257 (1856); L.B.
Sm. & W.J. Kress, Phytologia 66: 77-79 (1989).
Large xerophytes, leaves rosulate, serrate; inflo-
rescence strobilate, usually simple; floral bracts
coriaceous or ligneous; flowers in many ranks;
petals with reduced or lacking appendages; pollen
sulcate. Twenty two spp., terrestrial in lowland
and montane forests, rarely epiphytic, Northwest-
ern S America, Central America, SE Brazil.
39. Pseudananas Hassi. ex Harms
Pseudananas Hassi. ex Harms, in Engl. & Prantl, Nat.
Pflanzenfam. ed. 2, 15a: 153 (1930).
Plant propagating by elongate basal stolons;
leaves rosulate; blades with spinose margins;
scape evident; inflorescence simple, strobilate, the
apical coma inconspicuous or lacking; flowers
connate by their ovaries and forming a syncarp;
sepals free; petals bearing lateral folds which over-
lap the filaments; fruit large, succulent. (2n =
100). Only one sp., P. sagenarius (Arruda)
Camargo, terrestrial on open ground to dense
forest, from near sea level to 1400 m alt, Ecuador,
Bolivia, Brazil, Paraguay, Argentina.
40. Ananas Mill.
Ananas Mill., Gard. diet. abr. ed. 4 (1754).
Plant not producing stolons; leaves rosulate; scape
evident; inflorescence densely strobiliform, usu-
ally crowned with a tuft of sterile foliaceous bracts,
often producing slips at the base; flowers sessile;
sepals free; petals free, each typically bearing 2
slenderly funnelform scales; stamens included;
pollen ellipsoid with 2 pores; ovaries coalescing
with each other and with the bracts to form a
fleshy syncarp; epigynous tube short; placentas
apical; ovules caudate. (2n = 50, 3n = 75). Eight
95
spp., terrestrial in humid or dry open or forested
areas, native to S America south to Uruguay.
41. Acanthostachys Klotzsch
Acanthostachys Klotzsch, in Link, Klotzsch & Otto, Icon. pl.
rar. 1: 21, pl. 9 (1840); Rauh & Barthlott, Trop. subtrop.
Pflanzenwelt 39: 5-35 (1982).
Rhizomatose; leaves fasciculate or rosulate; scape
well developed or lacking; inflorescence simple,
densely polystichous-flowered; flowers sessile,
perfect; sepals free; petals free, bearing 2 scales;
stamens included; pollen biporate; ovary com-
pletely inferior, the thickened style base of A.
strobilacea (Schult. & Schult. f.) Klotzsch partially
superior; epigynous tube none or crateriform (A.
pitcairnioides (Mez) Rauh & Barthlott); ovules
long-caudate; seeds with a strongly elongated
apical appendage. (2n = 50). Two spp., epiphytic
and saxicolous, E Brazil, Paraguay, NE Argentina.
42. Hohenbergia Schult. & Schult. f.
Hohenbergia Schult. & Schult. f. in Roem. & Schult., Syst. Veg.
7(2): lxxi, 1251 (1830).
Stemless; leaves rosulate; blades spinose-serrate;
scape well developed; inflorescence of strobilate
spikes; flowers perfect; sepals mostly very short-
connate; petals with 2 appendages; stamens in-
cluded, series I free, series II partly adnate to the
petals; pollen with 2 or 4 pores; placentas mostly
apical; ovules obtuse to long-caudate. Forty seven
spp., terrestrial, saxicolous or epiphytic, West
Indies to E Brazil. Divisible into two subgenera,
Hohenbergia and Wittmackiopsis, according to
ovule shape, leaf anatomy, and inflorescence
branching.
43. Aechmea Ruiz & Pav.
Aechmea Ruiz & Pav., Fl. peruv. prodr.: 47 (1794); nom. cons.
Macrochordion de Vriese (1853).
Hoplophytum Beer (1854), incl. Platyaechmea (Baker) L.B. Sm.
& W.J. Kress.
Lamprococcus Beer (1856).
Ortgiesia Regel (1867).
Pothuava Gaudich. ex L.B. Sm. & W.J. Kress (1989).
Podaechmea (Mez) L.B. Sm. & W.J. Kress (1990).
Acaulescent, frequently propagating by rhizomes;
leaves usually forming a tank, usually spinose-
serrate; scape well developed; inflorescence com-
pound or simple; flowers distichous or polystic-
hous, sessile or stipitate, perfect; sepals free or
96 Bromeliaceae

connate, usually strongly asymmetric; petals free, 46. Wittrockia Lindm.

bearing 2 more or less adnate basal appendages;
Wittrockia Lindm., Ofvers. Forh. K. Sven. Vetenskapsakad.
stamens shorter than the petals, free or the second III. 24(8): 15, 20 (1891).
series more or less adnate to the petals; anthers
dorsifixed; pollen with 2, 3, 4 or numerous pores, Leaves rosulate, serrate or serrulate; scape usually
rarely sulcate; ovules mostly caudate; seeds evident; inflorescence compound, very densely
small. (2n = 42, 50). One hundred and eighty two capitiform in the center of an involucre of
species, largely terrestrial or epiphytic in sub foliaceous bracts; flowers perfect; sepals free or
forest. Mexico and the West Indies to Argentina. more or less connate; petals more or less connate
Divisible into seven subgenera, Podaechmea, or agglutinated near base, appendaged; stamens
Lamprococcus, Aechmea, Ortgiesia, Platyaechmea, included; style long but included; placentas cen-
Pothuava, and Macrochordion, according to inflo- tral or nearly; ovules obtuse. Eleven spp., terres-
rescence form, floral order, pedicels, and sepal trial, saxicolous, and epiphytic, E Brazil.
connation and armament.
Aechmea includes some very discordant ele-
ments and is very likely of polyphyletic origin. 47. Portea Brongn. ex K. Koch
Further research is likely to divide it with some Portea Brongn. ex K. Koch, Index Sem. Horti Bot. Berol.
parts becoming independent genera and others "1856", App.: 7 (1857).
merging with genera now considered distinct.
Leaves rosulate, serrulate; scape bracts brightly
44. Quesnelia Gaudich. colored; inflorescence compound; flowers slen-
derly pedicellate, perfect; sepals more or less con-
Quesnelia Gaudich., Voy. Bonite, Bot., Atl. pl. 54 (1842). nate; petals free, bearing 2 fimbriate scales at base;
stamens of the second series highly adnate to the
Stemless or rarely long-stemmed; leaves serrulate; petals; pollen with more than 5 pores; placentas
scape developed, inflorescence usually simple; various; ovules numerous, long-caudate. (2n =
flowers perfect; sepals free or short -connate; 50). Nine spp., saxicolous, terrestrial, or epiphytic
petals free, bearing 2 scales near base; stamens in forest, E Brazil.
shorter than the petals and generally included;
filaments of the second series highly adnate to the
petals; pollen ellipsoid, biporate; epigynous tube 48. Pseudaechmea L.B. Sm. & Read
distinct; placentas central to apical; ovules obtuse, Pseudaechmaea L.B. Sm. & Read, Phytologia 52: 53, pis. 3, 4
numerous. (2n = 50?, 100). Fifteen spp., terres- (1982).
trial, saxicolous, or epiphytic, from sea strand to
upland rainforest, E Brazil. Divisible into tw-o sub- Leaves rosulate, serrulate; scape elongate; inflo-
genera, Quesnelia and Billbergiopsis, according to rescence simple, laxly racemose; pedicels very
form of floral bracts and inflorescence. long; flowers polystichous, perfect; sepals subfree;
petals naked; stamens included; filaments free;
pollen biporate; epigynous tube none; placentas
45. Canistrum E. Morren 1
subapical; ovules caudate. Only one sp., P.
Canistrum E. Morren, Belgique Hort. 23: 257 (1873). ambigua L.B. Sm. & Read, native to tropical forest
in NW Colombia.
Mostly epiphytic; leaves rosulate, armed; scape
evident; inflorescence compound, densely corym-
bose, involucrate with the bracts covering all but 49. Hohenbergiopsis L.B. Sm. & Read
the apices of the flowers; flowers perfect; sepals Hohenbergiopsis L.B. Sm. & Read, Phytologia 33: 440 (1976).
free or nearly so; petals free, appendaged; stamens
included; pollen hi- or multiporate; ovules obtuse Coarse leaves rosulate; blades spinose-serrate;
or short-caudate, numerous; seeds many, sub- scape developed; inflorescence compound, the ul-
fusiform. (2n = SO). Ten spp., rainforests, E timate branches strobilate; flowers sessile, perfect;
Brazil. Divisible into two subgenera, Canistrum petals naked, attached to the filament tube but not
and Cucullatanthus, according to petal and ovule to each other; pollen in tetrads; epigynous tube
morphology (Leme 1997). lacking; ovules caudate. Only one sp., H.
guatemalensis (L.B. Sm.) L.B. Sm. & Read, on trees
1 Two species have recently been segregated to be members in dense wet forest. S Mexico, Guatemala.
of the new genus Edmundoa Leme (Leme 1997).
 Bromeliaceae
50. Androlepis Brongn. ex Houllet
Androlepis Brongn. ex Houllet, Rev. Hort. 42: 12 (1870).
~eaves rosulate, spinose-serrate; scape erect;
mflorescence compound with simple or divided
branches; spikes polystichous-flowered; flowers
functionally unisexual on different plants, sessile;
sepals free; petals short -connate, naked; stamens
included; filaments short, adnate; anthers each
bearing 2 foliaceous appendages at apex; pollen
biporate, in tetrads; epigynous tube short; ovules
numerous; borne along the upper part of the
locule, acute. Only one sp., A. skinneri Brongn. ex
Houllet, epiphytic in forest, S Mexico to Costa
Rica, and Peru.
51. Lymania Read
Lymania Read, J. Bromeliad Soc. 34: 201 (1984); Leme, J.
Bromeliad Soc. 39: 262-264.
Slenderly stoloniferous or stout-rhizomatous;
leaves few, forming a crateriform or utriculate
rosette, minutely serrulate; scape erect; inflores-
cence simple or bipinnate, lax or dense; flowers
sessile, polystichous, perfect; ovary deeply sulcate
or alate; epigynous tube very short but distinct;
sepals about half connate; petals free, naked; sta-
mens included; anthers sagittate, apiculate; pollen
porate, smooth; filaments connate or free the
.
mner series adnate to the petals; placentae
' nian Brazil and adjacent areas. Probably to be
su~apical; ovules obtuse or variously caudate;
frmt globose, ridged. Six mesophytic spp., Brazil:
Bahia.
52. Nidularium Lem. 2 Fig. 27G-L
Nidularium Lem., Jard. Fleur. 4 (Misc.): 60, pl. 411 (1854).
Andrea Mez (1896).
Stemless; leaves forming funnelform tanks,
serrate or serrulate; scape short; inflorescence
compound with flowers in flat fascicles and
covered by large bracts; flowers sessile or rarely
pedicellate, perfect; sepals more or less connate;
petals naked or rarely with vertical folds, tubular-
connate; stamens included; pollen biporate;
ovules numerous, obtuse. (2n = 50, 100).
Fifty spp., epiphytic or terrestrial in rainforest, E
Brazil.
A revision of the genus is forthcoming (Leme in
press).
2Pa~t of t~e .genus is being segregated as a separate genus
Camstropsts (mel. Andrea; Leme in press).
97
53. Neoregelia L.B. Sm.
Neoregelia L.B. Sm., Contr. Gray Herb. 104: 78 (1934); I.
Ramirez, Systematic Revision of Neoregelia subgenus
Hylaeaicum, St. Louis: M.Sc. Thesis (1991).
Leaves rosulate, usually spinose-serrate; inflores-
cence sunk in the center of the rosette, simple in
most species, very densely capitiform, umbellate
or corymbose; flowers perfect, pedicellate; sepals
asymmetric; petals free or connate, naked; sta-
mens and pistil included; ovules obtuse. (2n = 50).
Ninety seven spp., epiphytic in forest to saxicolous
and heliophytic, E Brazil to Amazonian S America.
Three subgenera, Hylaeaicum Neoregelia, accord-
mg to petal connation, pedicel morphology, and
polle~ morphology. Hylaeaicum probably merits
genenc status.
54. Streptocalyx Beer
Streptocalyx Beer, Flora 37: 348 (1854).
Terrestrial or epiphytic; leaves densely rosulate,
serrate; scape elongate to almost none; inflores-
cence 2-4-pinnate, laxly paniculate to densely
subglobose; flowers sessile; sepals free to distinctly
connate; petals free, naked; stamens and pistil in-
cluded; pollen 2- or 4-porate; epigynous tube dis-
tinct; seeds ellipsoid or ovoid, naked. (2n = 50).
Twenty spp., mostly in forest, Eastern and Amazo-
merged with Aechmea.
55. Ronnbergia E. Morren & Andre
Ronnbergia E. Morren & Andre, Ill. Hort. 21: 120, pl. 177
(1874).
Lea_ves fasciculate or rosulate, often petiolate,
entire or serrulate; scape well developed; inflores-
cence simple, elongate; flowers sessile, perfect; se-
pals more or less connate; petals free, naked or
with lateral folds; stamens included, the second
series free or partially adnate to the petals; pollen
biporate; epigynous tube large; placentas apical.
Twelve spp., epiphytic in rainforest, Costa Rica to
Peru. Probably to be merged with Aechmea.
56. Disteganthus Lem.
Disteganthus Lem., Fl. Serres Jard. Eur. 3: pl. 227 (1847).
Caulescent or acaulescent, propagating by stolons
and rhizomes; leaves petiolate, armed; scapes
several, borne beneath the rosette, bracteate;
inflorescence strobilate, simple; flowers perfect;
98 Bromeliaceae
sepals high-connate; petals naked, free; stamens
included; pollen porate; epigynous tube large; pla-
centas apical; ovules obtuse. Three spp., swampy
forest, Surinam, French Guiana, and adjacent
Brazil.
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100 Butomaceae
Butomaceae
C.D.K. CooK
Butomaceae Rich., Mem. Mus. Hist. Nat. 1: 364-372 (1815-
1816); in the sense of M. Pichon, Notul. Syst. 12: 170-183
(1946).
Aquatic, freshwater, rhizomatous perennial. Rhi-
zome dorsiventral, creeping, apparently mono-
podia!, axillary buds sometimes developing
into short-stalked bulbils. Leaves distichous or
subdistichous, borne on the apex of the rhizome,
erect, linear, glabrous not differentiated into peti-
ole and blade, and up to 1m or more long, trique-
trous, submerged or usually emergent, the base
somewhat extended and sheathing below; in-
travaginal scales numerous. Scapes developed on
alternate sides of the rhizome, usually separated
by an odd number (5, 7, 9) offoliage leaves, erect,
terete, glabrous. Inflorescence an umbel-like com-
plex of cymes subtended by (2-)3(-4) bracts.
Flowers pedicellate, perfect, protandrous, almost
hypogynous, actinomorphic. Perianth of 6 peta-
loid tepals in 2 whorls, free, white to pink. Stamens
9, the outer whorl with 6 and the inner with 3;
filaments flattened; anthers basifixed, 2-thecate,
latrorsely and longitudinally dehiscent. Gyno-
ecium of 6 conduplicate carpels arranged in 2 al-
ternate whorls of 3, the carpels connate at the base
into a ring, otherwise distinct, each apically nar-
rowed into a crested ventral somewhat decurrent
stigma; basally nectariferous on lateral carpel
walls; placentation laminar. Ovules numerous,
anatropous. Fruit of separate follicles, dehiscing
by widening of the ventral sutures. Seeds numer-
ous with ribbed testa and a straight embryo;
endosperm lacking.
Only Butomus umbellatus L., widespread in
temperate Eurasia and recently naturalised in
eastern N America. Butomus junceus Turcz. is not
considered to be worthy of specific status.
VEGETATIVE MORPHOLOGY. Described in detail
by Tomlinson (1982).
VEGETATIVE ANATOMY. Described in detail by
Stant (1967). Hairs are completely absent from all
parts. The roots may be contractile; the exodermis
is not differentiated into long and short cells. The
rhizome has a narrow outer cortex with numerous
crystals arranged as irregularly polygonal clusters
or as solitary styloids; the endodermis is lacking.
The stomata are paracytic, each has 1 pair of
lateral subsidiary cells.
INFLORESCENCE STRUCTURE. Described by
Charlton and Ahmed (1973) and Wilder (1974).
The scapes develop on alternate sides of the
rhizome, usually separated by an odd number
(5, 7, 9) of foliage leaves; at maturity they are erect
and terete. The inflorescence at maturity appears
to be a simple umbel, but in its development it
is a complex cymose system with trifurcations
followed by bifurcations.
FLOWER STRUCTURE AND ANATOMY. The devel-
opment of the flower is well documented by Singh
and Sattler (1974). All floral parts are clearly in
whorls of 3 except the outer whorl of stamens, in
which the 6 members are paired. The nectaries are
located on the lower radial walls of adjacent
carpels. The pedicel is supplied by many traces
which first become organised into a ring of about
12 vascular bundles and then become aggregated
in the receptacle to form a continuous vascular
cylinder supplying 3 traces to each tepal, 1 to each
stamen and 1 dorsal and 1 ventral trace to each
carpel. The dorsal traces of the carpels remain
unbranched into the style.
EMBRYOLOGY. The anther wall formation is of
the monocotyledonous type. The endothecium
has spiral wall thickenings and the tapetum is
plasmodial. The tapetal cells, when still intact, are
uninucelate and the microsporogenesis is suc-
cessive. The ovules are anatropous, bitegmic,
crassinucellate and have a parietal cell cut off from
the archesporia! cell. The embryo sac is of the
Polygonum type and endosperm formation
Helobial. Embryogeny is of the Caryophyllad
type.
PoLLEN MoRPHOLOGY. Described by Argue
(1972); pollen is sulcate and reticulate.
KARYOLOGY. Investigations on chromosome
numbers are summarised by Krahulcova and
Jarolimova (1993). 2n = 16, 20, 22, 24, 26, 28, 30,
39, 40, and 42 are reported. Triploid plants 2n =
39 are widespread in Europe and often outnumber
diploids (Hroudova and Zakravsky 1993b); they
are usually, but not invariably, sterile. Pollen of
the triploids is viable and hydridisation between
cytotypes is recorded. Under controlled condi-
 Butomaceae 101

tions, up to 34% of triploid cross-combination

seeds germinated.

PoLLINATION AND REPRODUCTIVE BIOLOGY. The

flowers are reported to be protandrous to homo-
gamous but even in protandrous plants the
stigmas are below the anthers (Fig. 28C,D). Self-
pollination or, at least, geitonogamy may take
place. The flowers are relatively large and attrac-
tive, secrete large quantities of nectar and have
exposed pollen. They are visited by many different
kinds of insects: particularly Hymenoptera and
Diptera (Woodhead and Kirchner 1908). A single
plant may develop ± 7000 seeds. In spite of the
possibility of self-pollination and the attraction of
insects, seed set is often poor and large stands may
not set seed at all. Self-sterility may be the result of
chromosomal imbalance, but very often the devel-
oping fruits are damaged by insects (Hejny 1960).
Diploids are self-compatible but triploids are self-
incompatible (Krahulcova and Jarolfmova 1993).
InN America, where B. umbellatus is introduced,
it sets good seed.
Little is known about seed dispersal. The seeds
sink in water within a few minutes. Germination
takes place underwater in spring. Dried seeds do
not germinate. Seeds stored in water germinate
after 3-4 months after collection (Hroudova and
Zakravsky 1993a); when stored at 1-3°C they
showed 68% germination after 5 years.
Vegetative reproduction is efficient. Small
rhizome fragments readily regenerate, axillary
bulbils on the rhizome may also play an important
role, and bulbils have been reported to develop in
the inflorescence in plants from Scandinavia
(Lohammer 1954). Under cultivation the species is Fig. 28A-J. Butomaceae. Butomus umbellatus. A Pseudo-
usually propagated by division of the rhizome. umbel. B Vertical section of leaf. C Pollen-shedding flower. D
Flower with spent stamens and receptive stigmas. E Carpel. F
Same, opened to show laminar placentation. G Stamen before
PHYTOCHEMISTRY. Proanthocyanins occur in the and H after release of pollen. I Gynoecium. J Seed. (Takhtajan
tannin cells. Cyanogenesis is not known in this 1982)
family.

AFFINITIES. The genera Butomopsis, Hydrocleys

and Limnocharis are often included in the
Butomaceae but they are clearly separate from
Butomus and should be placed in a separate
family, Limnocharitaceae, next to Alismataceae.
The monotypic Butomaceae clearly belongs in
Alismatiflorae, where it is distinct in having an
umbel-like inflorescence in combination with
laminar placentation, straight embryo and
monosulcate pollen grains. It shows affinities to
Alismataceae and Limnocharitaceae but lacks
schizogenous ducts and to the Aponogetonaceae
but lacks laticifers. All in all the simplicity and lack
of extreme specialisation of Butomaceae indicates
affinities to Hydrocharitaceae; this is also sup-
ported by work with rbcL (Chase et al. 1995).
DISTRIBUTION AND HABITATS. Butomaceae OC-
cur in most of Europe, but are absent from
many islands (including the Azores, Iceland and
many of the larger Mediterranean islands). The
family extends eastwards to Manchuria with
isolated populations in the Pamirs and eastern
China. It has been introduced into N America,
where it is spreading.
102 Butomaceae

Butomus is found on the banks of still or slowly
flowing water particularly in lakes, backwaters,
ponds, ditches and canals. It is rarely found in
swiftly flowing water, where it develops long, thin,
pale green submerged leaves and does not flower.
Triploids are more common in warm regions
with alkaline and base-rich soils (Hroudova and
Zakravsky 1993b); the diploids are commoner in
cool, acid, base-poor substrata.
PALAEOBOTANY. The fossil genus Butomites
cretaceus Velenovsky, described from leaf and
rhizome fragments from the Cretaceous of
Bohemia, does not resemble modern Butomus and
probably does not belong in the family. Tertiary
remains from the Oligocene to the Pleistocene
have been identified as Butomus (Mai 1985) from
central and E Europe; the remains are rather inad-
equate for confident determination.
EcoNOMIC IMPORTANCE. Butomus is commonly
grown as an ornamental. In the rice fields of the
Danube Delta it is considered an undesirable weed
and in N America there is evidence that it is more
competitive than many native reed-swamp
species.
The rhizomes contain much starch and are
eaten in parts of Russia; the starch is usually
mixed with grain starch for making bread or the
rhizomes may be prepared as a vegetable.
Only one genus:
Butomus umbellatus II. Reproduction, growth and biomass
production. Folia Geobot. Phytotaxon. 28: 413-424.
Hroudova, Z., Zakravsky, P. 1993b. Ecology of two cytotypes
of Butomus umbellatus III. Distribution and habitat differ-
entiation in the Czech and Slovak Republics. Folia Geobot.
Phytotaxon. 28: 425-435.
Krahulcova, A., Jarolimova, V. 1993. Ecology of two cytotypes
of Butomus umbellatus I. Karyology and breeding
behaviour. Folia Geobot. Phytotax on. 28: 385-411.
Lieu, S.M. 1979. Growth forms in the Alismatales. II. Two
rhizomatous species: Sagittaria lancifolia and Butomus
umbellatus. Can. J. Bot. 57: 2353-2373.
Lohammer, G. 1954. Bulbils in the inflorescence of Butomus
umbellatus. Sven. Bot. Tidsskr. 48: 485-488.
Mai, D.H. 1985. Entwicklung der Wasser- und Sumpfpfianzen-
Gesellschaften Europas von der Kreide bis ins Quartiir.
Flora 176:449-511.
Singh, V., Sattler, R. 1974. Floral development of Butomus
umbellatus. Can. J. Bot. 52: 223-230.
Stant, M.Y. 1967. Anatomy of the Butomaceae. J. Linn. Soc.
Bot. 60: 31-60.
Takhtajan, A.L. 1982. See general references.
Tomlinson, P.B. 1982. Helobiae (Alismatidae). In: Metcalfe
C.R. (ed.) Anatomy of monocotyledons, Vol. VII. Oxford:
Clarendon Press.
Weber, H. 1950. Dber das Wachstum des Rhizomes von
Butomus umbellatus L. Planta 38: 196-204.
Wilder, G.J. 1974. Symmetry and development of Butomus
umbellatus (Butomaceae) and Limnocharis flava (Limno-
charitaceae) Am. J. Bot. 61: 379-394.
Woodhead, T.W., Kirchner, 0. 1908. Butomaceae. In:
Kirchner, 0., Loew, E., Schroter, C. (eds.) Lebensgeschichte
der Bliitenpfianzen Mitteleuropas 1 (la). Stuttgart: E.
Ulmer.

Butomus L. Fig. 28
Butomus L., Sp. Pl.: 372 (1753).

Characters of the family.

Selected Bibliography

Argue, C.L. 1972. Pollen ofthe Butomaceae and Alismataceae.

I. Development ofthe pollen wall in Butomus umbellatus L.
Grana 11: 131-144.
Charlton, W.A., Ahmed, A. 1973. Studies in the Alismataceae.
IV. Developmental morphology of Ranalisma humile and
comparison with two members of the Butomaceae,
Hydrocleis [sic] nymphoides and Butomus umbellatus. Can.
J. Bot. 51: 899-910.
Chase, M.W., Stephenson, D.W., Wilkin, P., Rudall, P.J. 1995.
Monocot systematics: a combined analysis. In: Rudall, P.J.,
Cribb, P.J., Cutler, D.P., Humphries, C.J. (eds.) Monocotyle-
dons: systematics and evolution. Royal Botanic Gardens
Kew pp. 685-730.
Hejny, S. 1960. Okologische Charakteristik der Wasser- und
Sumpfpfianzen in der slowakischen Tiefebene (Donau- und
Theissgebiet). Bratislava: Slowak. Akad. Wiss. (SA V).
Hroudova, Z., Zakravsky, P. 1993a. Ecology of two cytotypes of
 Cannaceae 103

Cannaceae cultivated in tropical and more temperate regions

as ornamentals, often escaped and naturalised,
K. KUBITZKI comprising about 25 species (Segeren and Maas
1971) or only about 10 (P. Maas, pers. comm.).

MORPHOLOGY AND ANATOMY (Tomlinson

1969). The fleshy, underground, sympodially
branched rhizome bears distichous cataphylls,
and each segment of the rhizome ends in a leafy
Cannaceae Juss., Gen. Pl.: 62 (1789), nom. cons. shoot. The aerial shoots have well-developed
stems on which bladeless sheaths grading into the
Glabrous, erect, sometimes giant perennial herbs distal foliage leaves are produced. Phyllotaxy on
from sympodial, often tuberous-thickened, star- the aerial stem is distichous or in some species
chy rhizomes; mucilage canals traversing both the spiral, the latter condition maybe secondary due
rhizome and aerial stem. Leaves distichously to to torsion of internodes, although in view of the
spirally arranged, with a short, open, eligulate close relationship to Musa also a primarily spiral
sheath gradually passing into the petiole, and an arrangement would be comprehensible. The inflo-
expanded, simple, entire lamina rolled up from rescences are terminal on the leafy shoots and the
one side to the other in bud; blade with a promi- phyllotaxy of the bracts subtending the partial
nent, compound midrib and numerous lateral inflorescences shifts to tristichy.
veins in a pinnate-parallel arrangement, some- Salient anatomical features of Canna include a
times glaucous or lanuginose from a cover of normal root structure, the absence of hairs, and
extremely long epicuticular wax rodlets. Inflores- epidermis consisting of alternating files of wide
cence a terminal, bracteate, simple or compound and narrow cells, a colourless hypodermis beneath
thyrse with few-flowered cincinni, or by reduction the leaf epidermis, and a layer of hypodermal
a raceme, prophylls 2-keeled; flowers showy, "oblique cells" developed in the abaxial side of the
perfect, epigynous, pentacyclic, asymmetric with petiole and midrib. In the aerial stem no clear
no organ clearly median, basically trimerous but boundary between cortex and central cylinder is
with a modified androecium; sepals 3, free, green noticeable. In contrast to all other families of the
or purplish, not petaloid, persistent in fruit; petals Zingiberales, wide mucilage canals are found in
3, basally connate into a tube, 1 smaller than the the stem and rhizome. Vessels with simple or
other 2; stamen 1, petaloid, bearing a single scalariform perforations with few bars are found
bisporangiate theca along the edge, the other theca in the roots.
suppressed; staminodes 1-4( -5), petaloid, the one Crystals of calcium oxalate are common in
± opposite the functional stamen called labellum, ground tissue but raphide sacs are lacking. Silica
the remaining staminodes of different form and as druselike bodies are found in all parts except
size, called wings; gynoecium of 3 carpels forming the roots, particularly in the leaf hypodermis and
an inferior, trilocular ovary with axile placenta- bundle sheaths. Tannin is common in unmodified
tion; style petaloid; stigmatic surface Wet, papil- cells in all organs; starch is abundant in the
late, terminal and decurrent along an edge of the ground tissue of the rhizomes; in Canna edulis
style; septal nectaries present; ovules anatropous, and C. indica the grains attain a length of 130 f-tm.
bitegmic, crassinucellate, arranged in two alter- The epicuticular wax cover consists of com-
nating rows in each locule. Fruit usually a locu- pound rodlets of the Strelitzia type and is often
licidal, tuberculate-bristly capsule, sometimes particularly well developed on the inflorescences
indehiscent; seeds with a lid next to the radicle, (Frolich and Barthlott 1988).
not arillate; embryo straight, surrounded by a
thin, starchy endosperm and a copious, very hard, FLOWER STRUCTURE AND POLLINATION. The
starchy perisperm ("chalazosperm"). most recent studies on flower structure and devel-
A monogeneric family native to tropical and opment are by Kirchoff (1983) and Kunze (1984).
subtropical regions of the New World from the Still in bud, the (half) anther dehisces and deposits
lowland up to 2700 m above sea level, with some its pollen on the ventral face of the style in a large
species possibly occupying scattered areas in the patch but usually away from the terminal stig-
tropics of the Old World of doubtful indigenous matic strips. The style then shifts into a position
status, now widely cultivated in all tropical regions opposite to the labellum and, together with
for its starchy, edible rhizomes, and many hybrids staminodes, forms the "upper lip". This way, the
104 Cannaceae

totally asymmetric flower attains secondary sym-

metry. In Canna indica (Fig. 29) and some other
wild species, the placement of the pollen away
from the stigmatic areas on the apex of the style
avoids self-pollination; in still other species (or
garden cultivars?), the pollen deposit overlaps the
stigma and self-pollination occurs automatically.
For a fuller description and discussion of this in-
teresting instance of secondary pollen presenta-
tion, see Yeo (1993).
Most species seem to be ornithophilous, but
Vogel (1969) reported C. brittonii to be bat-
pollinated, with an enormous development of the
septal nectaries, and other species with extremely
elongated floral tubes probably phalaenophilous.

EMBRYOLOGY. The anther tapetum is amoeboid.

For pollen mother cells, both simultaneous and
successive cytokinesis have been reported. Pollen
grains are trinuclear when shed. The ovule is ana-
tropous, bitegmic and crassinucellar. The cells of
the nucellar epidermis divide and add to the nu-
cellar tissue. Development of the embryo sac is of
the Polygonum type. The antipodals are ephem-
eral. Endosperm development is Nuclear, but later
the tissue becomes Cellular (Davis 1966; Johri et
al. 1992).

PoLLEN MoRPHOLOGY. The pollen grains of

Canna are spherical, inaperturate, spinulose, and
with 60-90 11m diameter relatively large. The
pollen wall is exceptionally thick (> 10 [lm) and
contains only a minor part of detectable sporo-
pollenin, which is mainly localised in the spinules
at the pollen grain surface and in the in tine plasma
membrane. A coherent exine is absent and the
pollen wall is made up entirely by the intine. The
intine is composed of a thick (3-4 11m) outer chan-
nelled layer, the pectic "ectintine", and a some-
what thinner (1-2 [lm), fibrous band adjacent to
the protoplast, the cellulosic "endintine" (Skvarla
and Rowley 1970; Kress and Stone 1982). The
function of this peculiar pollen wall stratification Fig. 29A-D. Cannaceae. Canna indica. A Leaf and inflores-
B Infructescence. C Fruit, transversal section. D Seed-
may lie in facilitating the development of pollen cence.
ling. (Takhtajan 1982)
tube at any site on the pollen grain wall, apart
from permitting harmomegathy.

KARYOLOGY. Chromosome numbers are based
on x = 9, most species being diploids. Polyploidy,
and particularly triploidy, is restricted to artifi-
cially selected cultivars of wild species. Karyotypes
of some species were described by Sato (1960).
FRUIT AND SEED. After fertilisation, the ovule un-
dergoes a strong development of the chalaza. This
results in a pachychalazal seed in which the
tegumentary part covers only about one sixth of
the seed circumference. While in other zingi-
beralean families a sclerotised, silicified endo-
testa functions as the principal mechanical layer,
such an endotesta would be of little use in Canna,
where it could cover only a small part of the seed
surface. Therefore in Canna we find the develop-
 Cannaceae
ment of an exotestal layer composed of Mal-
pighian cells from tegumentary, raphal and cha-
lazallayers, which provide mechanical protection
and a physical barrier to imbibition. The intact.
seed coat is impermeable. However, the seed is
able to absorb water after the raising of the pre-
formed imbibition lid on the raphe, a structure
unique for Canna. During imbibition, the exotesta
looses its hardness, allowing the embryo to
emerge (Grootjen and Bouman 1988).
The seeds of Canna are able to survive extremely
long dormant periods. Seeds from archaeological
sites of an age of 600 years have proved still viable
(Sivori et al. 1968).
PHYTOCHEMISTRY (Hegnauer 1963, 1986). Apart
from chelidonic acid and aromatic resin (rhi-
zome) and non-fragrant resin (leaves), common
leaf phenolics including procyanidin and pro-
delphinidin are recorded for Canna. Various
species have molluscicidal properties.
AFFINITIES. The Cannaceae are clearly zingi-
beralean, and modern phylogenetic analyses (e.g.
Dahlgren and Rasmussen 1983; Kress 1990) and
morphological studies (Kunze 1984) agree in the
very close relationship between Marantaceae and
Cannaceae. However, differences between the two
families, including the positioning of pollen on
the ventral face in Cannaceae, the dorsal face
in Marantaceae, indicate that both are different
lineages.
PALAEOBOTANY. The silicified "pseudostem"
from India (Trivedi and Verma 1970) ascribed to
Canna deserves further study.
EcoNOMIC IMPORTANCE. Cultivated Cannas
comprise an assortment of over 1000 horticultural
varieties, most of which fall into the main groups,
Canna X generalis Bailey, and C. X orchidoides
Bailey, which each include numerous lineages.
Canna glauca, C. indica, C. iridiflora and C.
warszewiczii have been identified as the likely pro-
genitors of the usually diploid (or triploid) culti-
vars of C. X generalis. Around 1890, some C.
generalis biotypes were crossed with C. flaccida to
yield C. X orchidoides, which due to irregular
meiosis is completely sterile (Khoshoo 1972).
The starchy rhizome of Canna edulis is used as a
source oflivestock for animals and humans. Start-
ing from domestication in the northern Andes
around 2500 B.c. (Gade 1966), it is now cultivated
in warm regions around the world but mainly in
India and Australia, with yields of 25 t/ha. The
105
product from Australia is called Queensland
arrowroot. The starch grains of Canna are the
largest of all almidon-producing plants and may
be distinguished with the naked eye.
Only one genus:
Canna L. Fig. 29
Canna L., Sp. Pl. 1: 1 {1753); Kraenzlin, Pfianzenreich IV.47
(1912), monogr., now largely outdated; Segeren & Maas,
Acta bot. neerl. 20: 663-680 {1971), rev. northern S Amer.
Description as for family. Segeren and Maas dis-
tinguish two weakly defined subgenera based on
number of staminodes; the elaborate infrageneric
classification of Kraenzlin hardly tenable any
longer.
Selected Bibliography
Dahlgren, R., Rasmussen, F.N. 1983. Monocotyledon evolu-
tion: characters and phylogenetic estimation. Evol. Bioi. 16:
255-395.
Davis, G.L. 1966. See general references.
Frolich, D., Barthlott, W. 1988. See general references.
Gade, D. 1966. Achira, the edible Canna, its cultivation and
use in the Peruvian Andes. Econ. Bot. 20: 407-415.
Grootjen, C.J., Bouman, F. 1988. Seed structure in Cannaceae:
taxonomic and ecological implications. Ann. Bot. 61: 363-
371.
Hegnauer, R. 1963, 1986. See general references.
Johri, B.M., Ambegoakar, K.B., Srivastava, P.S. 1992. Com-
parative embryology of angiosperms Vol. 2. Berlin, Heidel-
berg, New York: Springer.
Khoshoo, T.N. 1972. Evolution of garden cannas. In: Vida, G.
(ed.) Evolution in plants. Symp. Bioi. Hungar. Vol. 12 pp.
93-101.
Kirchoff, B.K. 1983. Floral organogenesis in five genera of the
Marantaceae and in Canna. Am. J. Bot. 70: 508-523.
Kranzlin, F. 1912. Cannaceae. In: Engler, A. (ed.) Das
Pflanzenreich IV.47. 74 pp. Leipzig: W. Engelmann.
Kress, W.J. 1990. The phylogeny and classification of the
Zingiberales. Ann. Mo. Bot. Gard. 77: 698-721.
Kress, W.J., Stone, D.E. 1982. Nature of the sporoderm in
monocotyledons, with special reference to the pollen grains
of Canna and Heliconia. Grana 21: 129-148.
Kunze, H. 1984. Vergleichende Studien an Cannaceen- und
Marantaceenbliiten. Flora 175: 301-318.
Pai, R.M. 1965. Morphology of the flower in Cannaceae. J. Bioi.
Sci. 8: 4-8.
Rogers, G.K. 1984. The Zingiberales (Cannaceae, Marantaceae,
and Zingiberaceae) in the southeastern United States. J.
Arnold Arbor. 65: 29-38.
Sato, D. 1960. The karyotype analysis in Zingiberales with
special reference to the proto-karyotype and stable karyo-
type. Sci. Pap. Coil. Gen. Educ. Univ. Tokyo 10: 225-243.
Segeren, W., Maas, P.J.M. 1971. The genus Canna in northern
South America. Acta bot. neerl. 20: 663-680.
Sivori, E., Nakayama, F., Cigliano, E. 1968. Germination of
achira seed (Canna sp.) approximately 550 years old.
Nature 219: 1269-1270.
106 Cannaceae/Centrolepidaceae
Skutch, A.F. 1930. Unrolling ofleaves of Musa sapientium and
some related plants and their reactions to environmental
aridity. Bot. Gaz. 90: 337-365.
Skvarla, J.J., Rowley, J.R. 1970. The pollen wall of Canna and
its similarity to the germinal apertures of other pollen. Am.
J. Bot. 57: 519-529.
Takhtajan, A.L. 1982. See general references.
Tomlinson, P.B. 1969. Commelinales-Zingiberales. In:
Metcalfe C.R. (ed.) Anatomy of the Monocotyledons, Vol. 3.
Oxford: Clarendon Press.
Trivedi, B.S., Verma, C.L. 1970. Silicified pseudostem of Canna
L. from early Eocene of Deccan Intertrappean beds, M.P.,
India. Curr. Sci. 39: 442-443.
Vogel, S. 1969. Chiropterophilie in der neotropischen Flora.
Neue Mitt. III. Flora 158: 289-323.
Winkler, H. 1930. Cannaceae. In: Engler, A., Prantl, K., Die
natiirlichen Pflanzenfamilien 2nd edn. 15a: 640-654.
Leipzig: W. Engelmann.
Yeo, P.F. 1993. Secondary pollen presentation- form, func-
tion and evolution. Plant Syst. Evol. Suppl. 6. Wien:
Springer.
Centrolepidaceae
D.A. CooKE
Centrolepidaceae End!., Gen. Pl.: 199 (1836), nom. cons.
Small tufted annuals or cushion-forming perenni-
als. Stems with condensed internodes, producing
axillary adventitious roots but never rhizomatous.
Leaves alternate, basal in annuals, imbricate along
the stems in perennials, with open sheathing bases
and linear to subulate blades; uppermost leaf
reduced usually to a veinless sheathing cataphyll.
Inflorescence terminal, usually scapose, con-
densed compound-cymose, capitular within a pair
of bracts or spikelike with distichous bracts, often
containing shorter veinless hyaline secondary
bracts. Flowers minute, unisexual, naked, anemo-
philous, often combined into pseudanthia; male
flower a solitary stamen; filament glabrous; anther
dorsifixed, versatile, bisporangiate unilocular, de-
hiscing by a slit; female flower a solitary carpel;
ovary unilocular, stipitate; ovule 1, pendulous,
atropous; style terminal, filiform, undivided, per-
sistent, with stigmatic papillae along the adaxial
side. Fruit a membranous 1-seeded follicle,
usually deshiscing by an abaxial split; seed albu-
minous; embryo minute, apical; testa membra-
nous, formed from inner integument.
A family of three genera and about 35 species in
Australia, SE Asia, and a single species in south-
ernmost S America.
VEGETATIVE MORPHOLOGY AND ANATOMY.
Members of the family are small, tufted perennial
or annual herbs, which anatomically are simpli-
fied in comparison with Restionaceae. A short and
upright stem is present in the perennial species.
The anatomy of the vegetative parts of the family
has been treated by Cutler (1969). A pericycle is
usually absent from the roots. Narrow vessels
with scalariform perforations are reported to
be present in all vegetative parts. Stomata are
paracytic, with the subsidiary cells not derived
from the guard cell-mother cells, but from adja-
cent epidermal cells (Hamann 1963). Crystals and
silica are absent.
INFLORESCENCE AND FLORAL MORPHOLOGY. The
reproductive units are often pseudanthia, which
 Centrolepidaceae
are composed of unisexual, perianthless flowers
(Fig. 30). Each flower consists of a single, bis-
porangiate stamen, or a single, ascidiate carpel,
which is supplied by a dorsal bundle extending to
the stigma, and a ventral bundle supplying the
single, pendulous atropous ovule. Staminate and
pistillate flowers or bisexual pseudanthia com-
posed of 1-2 staminate and 1-many pistillate
flowers are grouped together in capitate or spicate
inflorescences. Hieronymus (1886) considered the
pseudanthia to be true flowers, a view that is now
rejected. The partial inflorescences are usually
interpreted as cincinni, but the suggestion by
Hamann (1962) that the whole complex inflores-
cence is racemose merits further attention.
Fig. 30A-G. Centrolepidaceae. Centrolepis aristata. A Habit.
B Head with outer primary bract detached. C Pseudanthium
with secondary bracts. D Pistil. E Stamen with secondary
bract. F Seed. G Same, longitudinal section. (Cooke 1992)
107
EMBRYOLOGY. The anthers are bisporangiate; the
inner of the 4 cell layers develops into a glandular
tapetum with binucleate cells. The pollen mother
cells form a single layer around the inner wall
of the pollen sac, just as in Gramineae and
Cyperaceae. The mature pollen grains are 3-celled.
The division of the pollen mother cells is succes-
sive. The ovules are pendulous and atropous,
bitegmic, and weakly crassinucellar. The arche-
spore cell does not produce a parietal cell. The
chalazal megaspore develops into an embryo sac
of the Polygonum type. In most species investi-
gated (Hamann 1975) the nuclei of the antipodals
divide, thus forming binucleate cells which de-
generate during endosperm development.
Endosperm development is Nuclear.
POLLEN MORPHOLOGY. The pollen grains (Fig.
31) are more or less uniform throughout the
family. They are monoaperturate and have a char-
acteristic exine with a scrobiculate and punctate
sculpture; the structure is granular (in contrast to
other Restionales) and an endexine is lacking. The
aperture is usually irregular in outline and a
margo is absent (Ladd 1977; Linder and Ferguson
1985).
KARYOLOGY. Chromosome data exist for a few
Centrolepis species only, and suggest that the base
number is 10 (Hamann 1962).
Fig. 31. Centrolepidaceae. Centrolepis aristata, distal
polar view of pollen grain showing irregularly defined
("centrolepoid") ulcus; interapertural exine scrobiculate. SEM
X2300. (Photo H.P. Linder)
108 Centrolepidaceae
POLLINATION. All species are pollinated by wind.
Self-pollination may also occur, especially in the
low-growing scapeless species.
FRUIT AND SEED. The pericarp is membranous
and usually dehisces by a longitudinal slit on the
dorsal side; this gives an appearance of paired
valves in the fruiting pseudanthia of Gaimardia
and Centrolepis. In Aphelia the pericarp does not
open, and forms part of the disseminule.
The mature seed coat is formed by the inner cell
layer of the inner integument. These cells are rich
in tannins. The outer cell layer of the inner integu-
ment and the complete outer integument degener-
ate during seed development, but the cuticle
between the 2 integuments is well developed. The
endosperm is starchy; the starch grains are com-
pound (Hamann 1962, 1975; Prakash 1969). The
embryo is conical and in an apical position.
DISTRIBUTION AND HABITATS. The centre of di-
versity is southern Australia, with some species in
the tropics extending to Malesia, Indochina, the
Philippines and Hai Nan Dao. Centrolepis extends
to New Zealand, and one Gaimardia species to S
America and the Falkands. Many species are
winter-growing annuals in temperate summer-dry
climates, where they grow in heath, scrub and
woodland communities on soils of low nutrient
status. The perennial species include alpine
herbfield plants such as Gaimardia setacea in Tas-
mania, New Zealand and the mountains ofSE Asia
to New Guinea. None are aquatic.
AFFINITIES. Centrolepidaceae are generally con-
sidered to be most closely related to Restionaceae,
with a simplified structure associated with their
herbaceous and usually anr.ual habit. This is
also supported by the embryological evidence,
although Linder and Ferguson (1985) have di-
rected attention to some subtle differences in
pollen morphology between Centrolepidaceae and
the rest of the Poales. The superficially similar
genera Hydatella and Trithuria, formerly placed
in this family, are now treated as the family
Hydatellaceae.
KEY TO THE GENERA
1. Inflorescence spicate with 6-18 distichous bracts
!.Aphelia
- Inflorescence capitular with 2 or 3 distichous bracts 2 margins. Inflorescences terminal on glabrous
2. Pseudanthia asymmetrical, of 1-many superposed carpels,
stamen solitary or absent 2. Centrolepis
- Pseudanthia symmetrical, of 2 collateral carpels, stamens 2
or absent 3. Gaimardia
1. Aphelia R. Br.
Aphelia R. Br., Prod.: 251 (1810); Cooke, J. Adelaide Bot. Gard.
16: 95-109 (1995), rev.
Small tufted annuals. Hairs unbranched. Leaves
all basal, 1-3-veined, with membranous to hyaline
sheathing bases; ligule absent. Scape terminal,
erect, terete, glabrous, consisting of a single inter-
node. Inflorescence a terminal laterally com-
pressed spike; rachis short, flexuose, bearing
distichous herbaceous bracts enclosing sessile
partial inflorescences or solitary flowers; primary
bracts 1 or 2; distal bracts 6-18, imbricate, 3-
veined, with hyaline margins, all simila, succes-
sively smaller, each becoming inrolled around a
fruit and caducous with it. Male flower with a
short secondary bract; filament capillary. Female
flower 1 in each distal bract. Fruit indehiscent;
seed fusiform. Six spp., endemic to southern
Australia.
2. Centrolepis Labill. Fig. 30, 31
Centrolepis Labill., Nov. Holl. Sp. Pl. 1: 7 (1804); Cooke, J.
Adelaide Bot. Gard. 15: 1-63 (1992), rev. Aust. spp.
Pseudalepyrum Dandy (1932).
Annual or perennial. Hairs unbranched or absent.
Leaves basal with dilated membranous sheathing
bases and 1-veined blades; ligule absent. Inflores-
cence a terminal cymose head, often scapose, en-
closed by 2 primary bracts on a very short rachis.
Secondary bracts 0 or 2 or 3 per pseudanthium.
Pseudanthia sessile, 1-many, each comprising 0-1
male and 1-30 female flowers. Male flower with
capillary filament. Ovaries within each pseu-
danthium superficially connate, alternating in 2
rows on a false axis (gynophore); styles at least
partially free. Fruit compound, of connate 1-
seeded follicles dehiscing abaxially; seed ovoid to
fusiform. Twenty spp., Australia and about six
others, SE Asia, New Guinea and New Zealand.
3. Gaimardia Gaudich.
Gaimardia Gaudich., Ann. Sci. Nat. I, 5: 100 (1825).
Cushion-forming perennials. Hairs lax and
branching or absent. Stems leafy; leaves obscurely
distichous, with imbricate scarious sheaths and 3-
veined blade. Ligule formed by union of the sheath
accrescent scapes. Primary bracts 2 or 3, lower-
most enclosing a pseudanthium of 2 equal, collat-
eral carpels and 2 stamens; next bract empty or
 Centrolepidaceae/Commelinaceae
containing carpels only. Third bract empty. Sec-
ondary bracts absent. Ovaries connate, stipitate;
styles free. Male flower with robust filament. Fruit
compound, the carpels dehiscing abaxially; seed
ovoid-oblong. Four spp., Tasmania, New Guinea,
New Zealand, S America and the Falkland Islands.
Selected Bibliography
Chanda, S. 1966. On the pollen morphology of the
Centrolepidaceae, Restionaceae and Flagellariaceae, with
special reference to taxonomy. Grana Palynol. 6: 355-415.
Cooke, D.A. 1992. A taxonomic revision of Centrolepis
(Centrolepidaceae) in Australia. J. Adelaide Bot. Gard. 15:
1-63.
Cutler, D.F. 1969. Juncales. in: Metcalfe, C.R. (ed.) Anatomy of
the monocotyledons, Vol. IV. Oxford: Clarendon Press.
Hamann, U. 1962. Beitrag zur Embryologie der Centrol-
epidaceen mit Bemerkungen tiber den Bau der Bliiten und
Bllitenstande und die systematische Stellung der Familie.
Ber. Dtsch. Bot. Ges. 75: 153-171.
Hamann, U. 1963. Uber die Entwicklung und den Bau des
Spaltoffnungsapparates der Centrolepidaceen. Bot. Jahrb.
Syst. 82: 316-320.
Hamann, U. 1975. Neue Untersuchungen zur Embryologie
und Systematik der Centrolepidaceen. Bot. Jahrb. Syst.
96:154-191.
Hieronymus, G. 1886. Dber Bllite und Bllitenstand der
Centrolepidaceen. Bot. Jahrb. Syst. 7: 319-330.
Ladd, P.G. 1977. Pollen morphology of some members of the
Restionaceae and related families, with notes on the fossil
record. Grana 16: 1-14.
Linder, H.P., Ferguson, I.K. 1985. On the pollen morphology
and phylogeny of the Restionales and Poales. Grana 24:
65-76.
Prakash, N. 1969. The floral development and embryology of
Centrolepis fascicularis. Phytomorphology 19: 285-291.
109
Commelinaceae
R.B. FADEN
Commelinaceae R. Br., Prod. Fl. Nov.-Hall.: 268 (1810)
("Commelineae"), nom. cons.
Perennial or occasionally annual, terrestrial, or
rarely epiphytic, monoecious, or andromono-
ecious, rarely polygamomonoecious, small to
large herbs, with erect to ascending, diffusely
spreading or stoloniferous, occasionally rhizoma-
tous, rarely scandent shoots. Roots fibrous or
tuberous. Leaves sheathing the stem at the base,
alternate or in pseudowhorls, spirally arranged
or distichous, sheaths closed, eligulate; lamina
simple, entire, often narrowed into a false
petiole, commonly somewhat succulent, ptyxis in-
volute, less commonly convolute or supervolute.
Inflorescences terminal or terminal and axillary,
in some genera all axillary and perforating
the sheaths, commonly a paniclelike thyrse com-
posed of several to many helicoid, cymose
branches (cincinni), sometimes reduced to a
single cincinnus or, rarely, to a single flower,
sometimes enclosed in or closely subtended
by a leafy bract (spathe). Flowers actinomorphic
or zygomorphic, usually bisexual or bisexual and
male, rarely female and bisexual or female, bi-
sexual and male, remaining open for only a few
hours, then deliquescing. Nectaries lacking. Calyx
composed of 3, equal or unequal, free or partly
fused, sepaline or petaline sepals. Corolla com-
posed of 3, equal to dimorphic, free or basally
fused, white or colored petals. Androecium
composed of 6 stamens arranged in 2 whorls,
sometimes all fertile and equal or unequal, often
2-3( -4) reduced to staminodes, occasionally 1-3
(rarely all) lacking, staminodes either alternating
with the stamens or else arranged on one side of
the flower and the stamens on the other, filaments
glabrous or some or all bearded, anthers dehiscing
longitudinally, rarely poricidally. Ovary superior,
bi- or trilocular, locules equal or dorsallocule re-
duced, ovules uniseriate or biseriate, 1-many per
locule. Style terminal on ovary, simple, usually
slender, stigma terminal, simple, small, or en-
larged. Fruit a dehiscent or rarely indehiscent
capsule, berry-like indehiscent capsule, or berry.
Seeds with an embryotega or operculum covering
110 Commelinaceae
the embryo, hilum dorsal, lateral, semilateral, or
terminal.
A mainly tropical and warm temperate family of
about 41 genera and 650 species.
VEGETATIVE MoRPHOLOGY. The overwhelming
majority of Commelinaceae are perennials, in-
cluding almost all of the species that occur in
forest. Perennials of nonforested habitats typically
persist during the dry season as leaf or stem
succulents or by means of underground storage
organs, including rhizomes, root tubers or, rarely,
bulbs. Annuals are common in some genera such
as Aneilema, Commelina, and Tripogandra, but
only Tinantia, among genera comprising more
than one species, is composed entirely of annuals.
Climbers are uncommon and are restricted to
forest. They occur in tropical America, Africa, and
Asia. The only African true climber is Palisota
thollonii, which has been recorded to 15m in
length, and is probably the longest/tallest species
in the family. Other genera with climbers include
Dichorisandra (tropical America) and Porandra
and the three genera of subtribe Streptoliriinae
(Asia). Some African species of Aneilema and
Commelina are sometimes described as scandent
because they scramble through other vegetation.
Tripogandra grandiflora of Mexico and Central
America seems to scramble similarly.
Epiphytes are rare in the family. Both species
of the neotropical Cochliostema can grow as
bromeliadlike tank-epiphytes that hold rainwater.
Cochliostema velutinum can also grow as a creep-
ing epiphyte. Although there are occasional
records of additional epiphytic Commelinaceae,
the only other species that is regularly epiphytic is
Belosynapsis vivipara of peninsular India.
The leaves of Commelinaceae have been charac-
terized as having involute vernation or ptyxis.
This is true for many genera, including Palisota,
Pollia, Tinantia, and Amischotolype. However, in
other genera, including Murdannia, Antheri-
copsis, Cyanotis, and Tripogandra, the leaves are
either simply folded or, more commonly, they
show a convolute or supervolute vernation. Both
involute and convolute vernation may occur
within some genera, such as Tradescantia and
Commelina.
VEGETATIVE ANATOMY. The vegetative anatomy
of the family has been summarized by Tomlinson
(1966, 1969). The stem has a distinctive basic
structure that distinguishes Commelinaceae from
all other monocots. A narrow cortex lacking vas-
cular tissue (except in Cartonema) surrounds a
central cylinder that is enclosed in an endo-
dermislike sheath. Longitudinal vascular bundles
are arranged in a characteristic pattern and con-
nect only at the nodes. The lamina has relatively
few longitudinal veins but a dense system of finer,
transverse veins. Succulence of the lamina is often
manifest in an epidermis comprising more than
half of the thickness of the leaf. In some genera,
water storage is mainly confined to a colorless hy-
podermallayer that is present beneath the adaxial
epidermis or rarely both epidermises.
Needlelike, calcium oxalate crystals (raphides)
are present in rap hide sacs that are arrayed in long
series to form raphide canals (lacking only in
Cartonema). Stomata usually have 4 or 6 (rarely 2)
subsidiary cells that are arranged in a characteris-
tic pattern (Tomlinson 1966, 1969). Three-celled
glandular microhairs are found in all genera
except Cartonema and Triceratella. Macrohairs
are diverse, but are multiseriate only in Palisota.
Short, stiff, apically directed hairs are common on
the lamina margins in many genera. Silica occurs
in a few genera and is confined to the epidermis.
FLORAL AND INFLORESCENCE MORPHOLOGY. The
flowers may be actinomorphic and vertical (Fig.
32) slightly to strongly zygomorphic and typi-
cally horizontal (Fig. 34). Zygomorphy is usually
due primarily to differentiation among the petals,
but it may also be due mainly to differences within
the androecium. Androecial zygomorphy, which
commonly accompanies a zygomorphic corolla,
may be the result of differentiation of the stamens
or stamens and staminodes into anterior and pos-
terior sets, as in Commelina and Aneilema (Fig.
34), or it may be the result of secondary reorien-
tation of the stamens in the open flower, as in
Tripogandra and some Murdannia.
The flowers may be somewhat asymmetric when
the style strongly curves to one side, as in enantio-
stylous species of Murdannia. In Cochliostema the
columnlike structure formed by the fused stamens
always bends to one side of the flower.
Floral dimorphism is common in the family.
In most cases the 2 flower types are bisexual
and male, with the male differing by the partial
or complete abortion of the gynoecium, as in
Commelina and many Aneilema species. Other
differences may include pedicel length (Com-
melina), stamen filament length and curvature
(Aneilema and Palisota), anther orientation
(Aneilema), and anther-sac dehiscence (Palisota)
(Faden 1991). Enantiostyly in Murdannia (Fig. 33)
is another form of floral dimorphism in the family,
as is the occurrence of cleistogamous as well as
 Commelinaceae 111

Fig. 32A-G. Commelinaceae. Cyanotis somaliensis. A Flower-

ing shoot. B Partial inflorescence and flower. C Flower with
front petal and three stamens removed; all stamens fertile and
equal. D Pistil. E Stamen. F Dehisced capsule. G Seed, dorsal
view (above) and ventral view (below). (Faden 1995) Fig. 33A-I. Commelinaceae. A-F Murdannia cryptantha. A
Habit. B Chasmogamous flower, front view; note antepetalous
staminodes. C Cleistogamous flower, slightly opened. D Cleis-
chasmogamous flowers in species of Commelina togamous flower, sepals removed. E Staminode with hastate
antherode. F Pistil. G-1 M. simplex. G Dehisced capsule. H
and Murdannia (Fig. 33B-D) (Uphof 1934; Faden Seed, dorsal view. I Seed, ventral view. (A-F Faden 1993, G-1
1993). Heterostyly has been reported erroneously Faden 1995)
in Aneilema (Ornduff 1974; see Faden 1991).
The flowers are typically pedicellate, trimerous,
and pentacyclic. The sepals and petals are imbri- Cochliostema and Buforrestia, similar in length
cate in bud, with 1 member, the outer sepal or but narrower (Floscopa), not clawed (Rho-
petal, usually overlapping the inner 2 members of palephora and Aneilema section Aneilema), of
the whorl. The calyx and corolla are usually dis- a different shape (Aneilema section Lampro-
tinct, with the sepals green or flushed with red to dithyros), or much smaller and sometimes incon-
violet and the petals white or pink to violet, rarely spicuous, as in Polyspatha and most species of
yellow to orange or red. In a few genera, Palisota, Commelina and Aneilema.
Pollia, and the genera of subtribe Streptoliriinae, The sepals are sometimes fused at the base, but
the sepals are petaline and very similar to the the calyx is rarely tubular (Weldenia). On the
petals. other hand, a clearly tubular corolla occurs in
In most genera the 3 sepals are equal and free, as several genera (Cyanotis, Coleotrype, Weldenia,
are the 3 petals. Not uncommonly, but especially and some species of Tradescantia).
in the genera with zygomorphic flowers, the outer The androecium shows the greatest variation
member or each of these whorls is differentiated among the floral whorls. The stamens may be all
from the inner members. In the calyx this differ- fertile and equal (Fig. 32), e.g., Streptolirion,
ence may be manifested by the outer sepal being Cyanotis, Amischotolype, and Stanfieldiella. The
slightly different in size, shape, or pubescence. antepetalous and antesepalous stamens may be
The sepals are never strongly differentiated slightly unequal in filament length, e.g., species
among themselves. In the corolla, the outer petal of Tradescantia section Cymbispatha or more
may be slightly larger than the inner petals, as in strongly dimorphic (species of Tripogandra). The
112 Commelinaceae

rangements, the middle of the 3 is often morpho-

logically distinct from the other 2.
One or more stamens or staminodes may be
lacking. When only 1 is missing it is usually the
one in front of the outer sepal, but occasionally it
is the outer antepetalous one, e.g., Geogenanthus.
When 3 are absent, they may all be from the same
whorl (species of Callisia and Murdannia) or they
may be 1-sided, as in Dictyospermum and some
Rhopalephora. Female flowers in Callisia repens
completely lack stamens.
Some or all of the filaments may be bearded
(Faden 1992), the presence or absence of filament
hairs often characterizing a genus. The hairs are
mainly moniliform, but a variety of hair types may
be present on the filaments in Aneilema species
(Faden 1991).
The anthers are basifixed or, less commonly,
versatile. Anther dehiscence is usually longitudi-
nal and may be introrse, extrorse, or latrorse. Api-
cal poricidal dehiscence occurs in all or most
species of Dichorisandra and Porandra and in at
least single species of Amischotolype and Coleo-
trype. The anthers in Cyanotis dehisce longitudi-
nally but the pollen is squeezed out basally in each
anther sac, making the anther functionally
poricidal.
The superior ovary is bi- or trilocular with the
Fig. 34A-N. Commelinaceae. Aneilema sebitense. A Habit. B locules 1-many ovulate. The ovules may be uni- or
Thyrse. C Flower, front view, note posterior staminodes with biseriate, with a genus usually showing one or the
bilobed antherodes. D Flower, side view. E Lateral stami- other arrangement. However, both arrangements
node. F Medial staminode. G Androecium, top view, medial occur within Palisota, Pollia, and Murdannia.
staminode omitted. H Pistil, side view. I Capsule attached to The style is simple, unbranched, and usually
cincinnus, side view. J Dorsal capsule valve, dorsal view. K
Dorsal locule seed, dorsal view. L Same, ventral view. M slender. It may be straight or curved and central in
Ventrallocule seed, dorsal view. N Same, ventral view. (Faden the flower or curved out of the floral midplane,
1991) especially in zygomorphic flowers. The presence
of enantostyly in Murdannia has been mentioned
above.
morphologically distinct sets of stamens may also Stigmas vary from very small to slightly capitate,
have a 1-sided arrangement, with the anterior penicillate, triangular, or slightly 3-lobed. Stigma
(or anticous) ones different from the posterior types in the family have been surveyed by Owens
(posticous) ones, e.g., Floscopa, Tinantia, and and Kimmins (1981).
Geogenanthus. The basic inflorescence type in Commelinaceae
Approximately 1/3 of the genera have sta- is a terminal, pedunculate thyrse (Fig. 34) with an
minodes. The staminodes may be all antesepalous indeterminate central axis to which are attached
Palisota, all antepetalous (Murdannia, Antheri- several - many, determinate, stipitate, cymose
copsis, and some species of Tripogandra), all ante- branches (cincinni) that may be alternate, sub-
rior (Cochliostema), or all posterior (Commelina, opposite, or subverticillate in attachment (Faden
Aneilema, Pollia, Polyspatha, Rhopalephora, 1991). At the base of each cincinnus is a bract, the
Tapheocarpa, and some species of Pollia). The cincinnus bract, and opposite each pedicel is a
sterile anthers, or antherodes, may be of a charac- bracteole.
teristic form for a genus, e.g., bilobed in Aneilema Evolutionary trends in the inflorescence largely
(Fig. 34E,F), trilobed or hastate in Murdannia have been in the direction of reduction or loss of
(Fig. 33£), and cruciform in Commelina. When some parts or axes (see Brenan 1966). However,
the staminodes and/or stamens are in 1-sided ar- while the cincinnus bracts have become caducous
 Commelinaceae
or have disappeared in some genera, in other
genera such as Tradescantia they have become
enlarged and foliaceous or spathaceous, and
sometimes lateral inflorescences become closely
associated with the terminal inflorescence form-
ing a compound inflorescence or synflorescence,
as in Aneilema (Faden 1991), Callisia, Stanfiel-
diella, Tripogandra, and other genera. Like the
cincinnus bracts, the bracteoles may have become
caducous or been lost but they have never become
much enlarged.
Change in position of the inflorescence has oc-
curred from terminal or terminal and axillary to
strictly axillary and sometimes perforating the leaf
sheaths, particularly in forest understory genera
(Faden 1988a), e.g., Buforrestia, Coleotrype, and
Amischotolype, or even basal (Geogenanthus). In a
few genera, e.g., Commelina, the inflorescences
are morphologically terminal but become leaf-
opposed and superficially appear lateral due to the
sympodial growth of the shoot.
FRUITS AND SEEDS. The fruits are typically
loculicidally dehiscent capsules, but occasionally
they are indehiscent. In Palisota they are fleshy
berries, and in Pollia and a few species of
Commelina they are indehiscent, crustaceous,
and colorful. In Tradescantia zanonia the sepals
greatly increase in size, envelop the capsule, and
become fleshy and berrylike. Capsules are usu-
ally hi- or trilocular, but in a few species of
Commelina, Aneilema, and Rhopalephora they are
unilocular.
Seeds may be uni- or biseriate in the locules and
range from 1 to about 50 per fruit. Seeds range
from about 0.5-8 mm in length. The hilum varies
from punctiform to linear. A caplike callosity, the
embryotega (or embryostega or operculum), cov-
ers the embryo and is sharply demarcated from
the rest of the testa by a clear line or groove, the
micropylar collar, in all genera except Cartonema
(Grootjen and Bouman 1981; Grootjen 1983). The
embryotega ranges from dorsal to lateral or, in
the case of Cyanotis and Belosynapsis, terminal.
Bruckner (1926) has shown that the combination
of hilum shape and position of the embryotega
can be used to characterize many genera, such as
Commelina (hilum linear, embryotega lateral). A
fleshy aril, usually red, covers the seeds in
Dichorisandra, Amischotolype, and Porandra. The
testa may be smooth, rugose, alveolar, reticulate,
scrobiculate, foveate, foveolate, furrowed, or
ribbed, the pattern often radiating on the dorsal
surface from the embryotega. At germination the
embryotega is lifted off or pushed to one side due
113
to pressure from the developing embryo beneath
it.
A summary of the anatomy of the seeds has been
published by Takhtajan (1985). The ovules of
Commelinaceae are atropous to hemianatropous,
camplyoptropous, or rarely anatropous, bitegmic
and crassinucellate. The testa is sometimes
formed by the inner integument and the silicified
innermost layer of the outer integument. In those
cases, e.g., Stanfieldiella and Cartonema (Grootjen
and Bouman 1981), the remaining layers of the
outer integument form a loose cover around the
seed and may be sloughed off. In many genera
the outer layers of the outer integument may be
more firmly attached and are part of the testa.
The embryotega is surrounded by a micropylar
collar except in Cartonema, where it still can be
identified as a bump. It is covered by both integu-
ments except in Cartonema, in which only the
outer integument covers it.
The embryo is well differentiated except in
Cartonema. It is usually short-cylindric and
dumbbell-shaped in outline. Endosperm is abun-
dant and starchy. Germination is hypogaeal. Seed-
lings of Commelinaceae have been studied by
Tillich (1995), who reports that the most distinc-
tive feature of the family is a root collar, which,
however, is lacking in Cartonema.
KARYOLOGY. Chromosome numbers have been
published for all genera except Triceratella,
Sauvallea, Aetheolirion, Porandra, Elasis, Pseu-
doparis, and Tapheocarpa. The most comprehen-
sive overview is that of Jones and Jopling (1972)
with a contribution by Faden and Suda (1980).
Basic numbers vary from x = 4 (some Gibasis)
to x = 29 (Rhopalephora). Small and midsized
genera (up to 15 spp.) tend to have single basic
numbers, e.g., Stanfieldiella (x = 11), Polyspatha
(x = 14), and Buforrestia (x = 17). In contrast, the
larger genera usually have multiple basic num-
bers, e.g., x = 9, 10, 13-16 in Aneilema (Faden
1991), x = 11-15 in Commelina, and x = 6, 7?,
9-11 in Murdannia.
In general, the chromosomes of subfamily
Cartonematoideae (unknown in Triceratella) and
the tribe Commelineae (sensu Faden and Hunt
1991) are relatively small (Jones and Jopling 1972),
while those of tribe Tradescantieae are medium -
large. Basic number and chromosome size show
very weak correlation: the basic numbers in tribe
Commelineae range from x = 6 to x = 29, while in
tribe Tradescantieae they vary from x = 4 to x =
20. A nombre fondamental of 7 has been shown
for Tradescantia sect. Cymbispatha (as genus
114 Commelinaceae
Cymbispatha by Jones et al. (1981). Bimodal
complements characterize some genera, such as
Anthericopsis, Floscopa, and Tinantia, and also
occur in occasional species of other genera.
The ancestral basic chromosome number for
Commelinaceae or for either large tribe is unclear.
The pattern has been obscured by repeated polyp-
loidy, both paleopolyploidy and neopolyploidy,
and by aneuploid reduction in basic number, at
least within some genera, e.g., Aneilema (Faden
and Suda 1980). Aneuploid reduction in basic
chromosome number by Robertsonian transloca-
tion has been demonstrated for Gibasis by Jones
(1974) and postulated for Tradescantia (including
Zebrina) by Jones (1990).
POLLEN MORPHOLOGY. Pollen is shed as indi-
vidual grains. Pollen grains are sulcate except in
a few species of Tinantia, Tradescantia, and
Tripogandra (Poole and Hunt 1980). Although a
number of exine patterns were recognized by
Poole and Hunt (1980), Faden and Hunt (1991)
combined them into 2 general types, spinulose
and lacking spines. The pollen grains may be di-
morphic in different anthers of the same flower
(see p. 115).
PHYTOCHEMISTRY. The available information
was surveyed by Hegnauer (1963, 1986). Saponins
and alkaloids seem to be absent from the
Commelinaceae, while common flavonols, gly-
coflavones, and phenolic acids are widespread.
Gallic acid was once recorded for Commelina
agraria, a record in need of confirmation. Flower
pigments are based on glycosides of cyanidin and
delphinidin, particularly the latter often stabilized
by complex copigmentation. The presence of
cyanogenesis was proved for Tinantia Jugax ( =
T. erecta). Harris and Hartley (1980) found
unlignified cell walls fluorescent due to the pres-
ence of bound cinnamic acids.
DISTRIBUTION, PHYTOGEOGRAPHY, AND HABI-
TATS. The Commelinaceae consists of about 41
genera and 650 species. The family as a whole is
distributed worldwide but is especially diverse in
the tropics. In north temperate regions the family
is well represented in North America and Asia, but
no species is native to Europe.
Subfamily Cartonematoideae is confined to the
Southern Hemisphere. All seven subtribes of tribe
Tradescantieae are wholly confined either to the
Old World (subtribes Palisotinae, Streptoliriinae,
Cyanotinae, Coleotrypinae) or the New World
(Dichorisandrinae, Thysantheminae, Tradescanti-
inae ), suggesting a long period of isolation. The 14
genera of tribe Commelineae are either wholly Old
World (eight genera) or else they have native spe-
cies in both the Eastern and Western Hemispheres
(six genera). Of the latter six genera (Murdannia,
Buforrestia, Floscopa, Pollia, Aneilema, and Com-
melina), all but Buforrestia, which is confined to
West Africa and northeastern South America, are
widespread, occurring in Australia, Asia, Africa,
and tropical America.
The major centers of diversity for the family
are Mexico (especially Oaxaca and Chiapas) and
northern Central America (especially for the
subtribes Tradescantiinae and Thysantheminae);
tropical Africa (including Madagascar
particularly for tribe Commelineae and subtribes
Palisotinae and Coleotrypinae); and peninsular
India and the foothills of the Himalayas to
Thailand and southwestern China (especially
for Commelineae, and subtribes Cyanotinae and
Streptoliriinae). South America, although poorer
in species than either Africa or Asia, is the center
of diversity for subtribe Dichorisandrinae.
Commelinaceae are ecologically diverse. They
are found chiefly in humid and mesic habitats,
such as forest and grassland. Within the tropics,
where they grow from sea level to 3800 m, species
occur in all but the most xeric and most saline
habitats. Few Commelinaceae are aquatics except
for species of Floscopa.
REPRODUCTIVE BIOLOGY. Flowers lack nectar
and are open for only a few hours during the day.
Andromonoecy is common, with the bisexual and
male flowers sometimes occurring in specific parts
of the inflorescence and/or opening in a predict-
able sequence; but in some genera there is no
pattern to their position or flowering order. Poly-
gamomonoecy and gynomonoecy are rare. Dioecy
is lacking.
The short anthesis allows related species to
occur sympatrically without crossing. This is true
in Aneilema (Faden 1983, 1991) and Stanfieldiella,
and is likely also to be the case in Palisota,
Polyspatha, and other genera.
Commelinaceae flowers are chiefly entomophil-
ous and usually attract a variety of bees and
diptera. Buzz pollination has been reported in
Dichorisandra (Sigrist and Sazima 1991) and
might be predicted in the other genera that have
apical poricidal anther dehiscence (Porandra and
some species of Amischotolype and Coleotrype).
More specialized pollination systems have not
been reported but might be expected in Cochlio-
stema because of its highly modified androecium.
 Commelinaceae
Autogamy is common in many genera. Self-
incompatibility has been found in some Trade-
scantiinae, which would make them obligate
outcrossers (Owens 1981). Callisia repens and
perhaps one other species are probably ane-
mophilous.
Individual flowers are usually brightly colored
and conspicuous to potential pollinators, even in
autogamous species. When the flowers are small,
however, the inflorescences and/or associated
bracts may attract pollinators. The most extreme
case occurs in some populations of Coleotrype
madagascarica in which the uppermost leaves on
the flowering shoot may be bright pink, like bracts
of poinsettia (Faden 1992).
Floral scents are uncommon but, when present,
they may be strong, as in Callisia fragrans and
Cochliostema. In Palisota, floral odors vary from
species to species and may be biologically and
taxonomically significant (Faden 1992).
The position and length of the stamens may af-
fect the species of insects that visit a flower and the
place of deposition of pollen on the insect, as in
Aneilema (Faden 1992). The reciprocal position of
the lower, fertile stamen in male flowers and of the
gynoecium in bisexual flowers in Palisota en-
hances pollen transfer. Mirror-image symmetry of
the stamens and style, or enantiostyly, occurs in
many species of Murdannia.
Floral hairs are common in some genera (Faden
1992). Although they may be present on any of the
floral parts, it is primarily those of the stamen
filaments that are related to pollination. Filament
hairs may attract pollinators when they are
brightly colored, provide footholds for insects,
retain shed or dropped pollen grains within the
flower, and affect the efficiency with which pollen
can be collected (Faden 1992).
Pollen is the only reward for pollinators. Often
staminodes, yellow hairs, or dimorphic stamens
deceptively attract insects away from the main
source of pollen in the flower (Vagel 1978; Faden
1992). Pollen grains may be differentially colored
in different anthers of the same flower. Dimorphic
pollen occurs in individual flowers of Palisota,
Tripogandra, Aneilema, and Tinantia species (Lee
1961; Handlos 1975; Simpson et al. 1986; Faden
1988b, 1991).
Information on seed dispersal in Commelina-
ceae is almost all conjectural or anecdotal. Most
Commelinaceae seeds have no specialized means
of dispersal. The berries of Palisota and berrylike
fruits of Pollia, a few species of Commelina, and
Tradescantia zanonia are probably endozoo-
chorous. The sticky fruits of Rhopalephora are
115
likely to be dispersed externally on birds and
mammals. Ants have been observed carrying
seeds of Commelina erecta (R.B. Faden, unpubl.).
The winged seeds of Aethiolirion are probably
wind-assisted in their descent. Dispersal by water
and/or in mud on animal feet is likely for spe-
cies of Commelina, Murdannia, Cyanotis, and
Floscopa that grow in or beside water. Dimorphic
seeds or propagules with presumably different
means of dispersal are frequent in Commelina and
Aneilema (Faden 1991).
SuBDIVISION. The history of the subdivision of
the Commelinaceae was summarized by Faden
and Hunt (1991). The predominant trend, which
was begun by Meisner (1842), was to divide
the family into two major groups. His tribe
Tradescantieae had 6 fertile stamens while tribe
Commelineae had a reduced number of fertile sta-
mens. Hasskarl (1870) used Meisner's classifica-
tion, as did Clarke (1881), who also separated out
a small, third tribe, Pollieae, for genera with
berries or berrylike fruit.
Bruckner (1926, 1930) raised the two major divi-
sions of the family to subfamily rank and distin-
guished them on the basis of floral symmetry:
subfamily "Tradescantieae", (flowers actinomor-
phic); subfamily "Commelineae" (flowers zygo-
morphic). Subfamily Tradescantieae was divided
into tribes Hexandreae (6 fertile stamens) and
Triandreae (3 fertile stamens), and subfamily
Commelineae was split into tribes Declinatae
(buds bent downward, the 3 posterior stamens less
developed and usually sterile) and Inclinatae
(buds curved inward, the 3 anterior stamens less
developed and usually sterile).
Woodson (1942) and Rohweder (1956), focusing
on the American genera, returned to 2 tribes,
Commelineae and Tradescantieae, but defined
them on the basis of inflorescence rather than
floral characters.
Pichon ( 1946) was the first worker to completely
break away from the tradition of dividing the
family into 2 major components. He recognized
10 tribes, Tradescantieae, Callisieae, Anthericops-
ideae, Commelineae, Geogenantheae, Cochlios-
temateae, Pseudoparideae, Zebrineae, Cyanoteae,
and Dichorisandreae, on morphological charac-
ters. He further separated the genus Cartonema
into its own family Cartonemataceae, based in
part on anatomical grounds.
Brenan (1966) recognized 15 informal "groups,"
using a combination of morphological characters.
He predicted that not all of his groups would
prove to be of equal taxonomic rank.
116 Commelinaceae
Faden and Hunt (1991), which was briefly pre-
viewed by Faden (1985), accepted 2 subfamilies,
subfamily Cartonematoideae, which was divided
into 2 tribes, tribes Cartonemateae [as Cartone-
meae] and Triceratelleae, and subfamily Com-
melinoideae, which also comprised 2 tribes, tribes
Tradescantieae and Commelineae. Within tribe
Tradescantieae, 7 subtribes were accepted, Pali-
sotinae, Streptoliriinae, Cyanotinae, Coleotry-
pinae, Dichorisandrinae, Thyrsantheminae, and
Tradescantiinae. This classification employed
anatomical, palynological, and cytological charac-
ters in addition to morphological ones, but it did
not include a cladistic analysis.
Evans (1995) studied the cladistic relationships
among the genera of Commelinaceae using mor-
phological characters and sequence data from the
chloroplast gene rbcL. He also made a combined
analysis of the two data sets. Cartonema was
found to be basal in the family. Tribe Com-
melineae was shown to be monophyletic, as were
subtribes Cyanotinae and Coleotrypinae. Subtribe
Thyrsantheminae was polyphyletic, and tribe
Dichorisandrinae was polyphyletic in some anal-
yses and monophyletic in others. With the
inclusion of the genus Elasis from subtribe
Thyrsantheminae, subtribe Tradescantiinae was
monophyletic. The positions of the subtribes
Streptoliriinae and Palisotinae varied, with
Palisota, the sole genus of subtribe Palisotinae,
sometimes emerging as the sister group to the en-
tire family except for Cartonema. From the above
it is evident that molecular studies need to be ex-
panded to more genera and more genes, and that
additional nonmolecular characters need to be
studied systematically.
AFFINITIES. Modern ideas about the relation-
ships of the Commelinaceae vary greatly, depend-
ing upon the kinds of data employed. Using
characters other than DNA sequence data, the
family is generally combined with Mayacaceae,
Xyridaceae, Rapateaceae, and sometimes Erio-
caulaceae as the order Commelinales (see
Dahlgren and Clifford 1982 for a review of classifi-
cations of the monocotyledons up to that time).
The most comprehensive such analysis, using 101
morphological, anatomical, embryological, and
phytochemical characters and all of the families
of Dahlgren et al. (1985), yielded an order Com-
melinaceae comprising the same five families
(Stevenson and Loconte 1995).
In contrast to the above, analyses using DNA
sequence data for the plastid gene rbcL (Chase et
al. 1993, 1995, 1995a; Duvall et al. 1993) place the
Pontederiaceae, Philydraceae, and Haemodora-
ceae as the families closest to Commelinaceae.
Combining morphological characters and rbcL
sequence data produced results identical with or
very similar to the rbcL phylogenies alone (Chase
et al. 1995b; Kellogg and Linder 1995), showing
the Pontederiaceae as the sister group to the
Commelinaceae.
KEY TO THE SUBFAMILIES, TRIBES, SUBTRIBES,
AND GENERA
1. Rap hide canals absent or next to the veins of the lamina;
glandular microhairs lacking; internodes and/or foliage
glandular-pubescent; flowers yellow, actinomorphic.
Subfam. Cartonematoideae 2
- Raphide canals present and mostly between the veins of
the lamina; glandular microhairs almost invariably
present; flowers pink or bule to white, rarely yellow to
orange, actinomorphic or zygomorphic. Subfam.
Commelinoideae 3
2. Rap hide canals lacking; cincinni 1-flowered, the inflores-
cence seemingly a spiciform raceme; stomata with 2 or 4
subsidiary cells; perennials or annuals; Australia. Tribe
Cartonemateae 1. Cartonema
- Raphide canals next to the veins; cincinni several-
flowered, the inflorescence composed of leaf-opposed
cincinni; stomata with 2 subsidiary cells; annuals; Africa.
Tribe Triceratelleae 2. Triceratella
3. Stomata with 4( -2) subsidiary cells, or if 6, then the termi-
nal pair equal to or larger than the second lateral pair
(Geogenanthus and Streptoliriinae ); pollen exine lacking
spines (except Tripogandra), mostly with a cerebroid tec-
tum; flowers mainly actinomorphic; filament hairs (when
present) moniliform (except Tripogandra species). Tribe
Tradescantieae 4
- Stomata with 6 subsidiary cells, the terminal pair smaller
than the second lateral pair; pollen exine spinulose, tectum
perforate; flowers actinomorphic or zygomorphic; fila-
ment hairs (when present) usually not moniliform. Tribe
Commelineae 26
4. Cincinni elongate or contracted, never fused in pairs 5
- Cincinni contracted, fused in bifacial pairs (rarely free,
stipitate, 2-several in a pseudo-umbel, with the cincinnus
axis reflexed at junction with stipe: Gibasis). Subtribe
Tradescantiinae 23
5. Fruit a berry; sepals petaloid; pollen-bearing stamens 3,
antepetalous. Subtribe Palisotinae 3. Palisota
- Fruit capsular; sepals sepaloid; pollen-bearing stamens
1-6, if 3, then not antepetalous 6
6. Plants scandent (except Spatholirion ornatum); inflores-
cences thyrses, the lowermost 1-2 cincinni subtended
by spathaceous bracts; flowers bisexual and male or,
occasionally, all bisexual. Subtribe Streptoliriinae 7
- Plants rarely scandent; inflorescences various, if
thyrsiform, then lower cincinni not subtended by
spathaceous bracts; flowers usually all bisexual, rarely
bisexual and male 9
7. Stamens 6, equal; capsules ovoid or ellipsoid, less than
2 em long; seeds not winged 8
- Stamens 3 or 5, when 5, unequal; capsules linear, 8-16cm
long; seeds winged 6. Aetheolirion
8. Inflorescences axillary, perforating, also sometimes leaf-
opposed; all cincinni bracteate 4. Streptolirion
 Commelinaceae 117

- Inflorescences all leaf-opposed; upper cincinni ebracteate 21. Petals free, blue to purple; cincinni 1-several per shoot;
5_ Spatholirion shoots usually elongate, rarely rosettelike 22
9. Seeds with a terminal embryotega; filaments and style - Petals united into a long tube, white or rarely blue;
usually with a subterminal swelling ( Cyanotis ); anthers cincinni numerous, aggregated towards the shoot apex,
usually dehiscing by basal pores (Cyan otis); bracteoles rosette plants 19. Weldenia
conspicuous, herbaceous; plants succulent. Subtribe 22. Cincinni stipitate; plants procumbent; roots not tuberous
Cyanotinae 10 20. Elasis
- Seeds with a dorsal to lateral embryotega; filaments and - Cincinni sessile; plants erect; roots tuberous
style lacking a swelling; anther dehiscence various but not 21. Matudanthus
by basal pores; bracteoles and plants various 11 23. Cincinni stipitate, free, 2-several in a pseudo-umbel
10. Petals free; filaments not swollen; anther dehiscence 22. Gibasis
longitudinal 7. Belosynapsis - Cincinni sessile, fused in back-to-back pairs, the pairs
- Petals united into a tube; filaments usually swollen near variously arranged 24
the apex; anther dehiscence functionally by basal pores; 24. Cincinni subtended by or enclosed in spathaceous or folia-
cincinni compact 8. Cyanotis ceous bracts 23. Tradescantia
11. Inflorescences all lateral and perforating the sheath, - Cincinni subtended by small bracts 25
cincinni usually congested and contracted, bracts 25. Flowers actinomorphic; stamens 0-6, equal or unequal,
and bracteoles herbaceous; paleotropical. Subtribe but never alternating long and short 24. Callisia
Coleotrypinae 12 - Flowers zygomorphic; stamens 6, the antesepalous short,
- Inflorescences terminal or terminal and lateral, rarely the antepetalous long 25. Tripogandra
all lateral and/or perforating the sheaths; bracts and 26. Flowers solitary, terminal, becoming leaf-opposed; fruit
bracteoles various; neotropical and New World temperate deeply buried 40. Tapheocarpa
14 - Flowers in inflorescences variously arranged; fruit usually
12. Petals free; seeds arillate 13 aerial 27
- Petals united into a tube; seeds exarillate 9. Coleotrype 27. Cincinni enclosed in or closely subtended by spathaceous
13. Plants of various habits but not scandent; anther bracts 28
dehiscence usually longitudinal 10. Amischotolype - Cincinni not enclosed in or closely subtended by
- Plants scandent; anther dehiscence poricidal spathaceous bracts 30
11. Porandra 28. Inflorescences all axillary and perforating the sheaths;
14. Ovules biseriate (except an undescribed genus); petals fertile stamens 6; fruiting sepals accrescent, more than
sometimes fringed; anther dehiscence sometimes pori- lOmm long; petals not clawed 29. Buforrestia
cidal (Dichorisandra); chromosomes large, basic number - Inflorescences terminal or leaf-opposed, rarely some also
x = 19. Subtribe Dichorisandrinae 15 axillary and perforating; fertile stamens 3; fruiting sepals
- Ovules uniseriate; petals never fringed; anther dehiscence marcescent, less than 7 mm long; upper petals clawed 29
longitudinal; chromosomes small to medium-sized, 29. Spathes distichously arranged on an axis, never leaf-
basic number various but not x = 19. Subtribe opposed; flowers white; capsules bilocular, locules 1-
Thyrsantheminae 18 seeded 34. Polyspatha
15. Petals not fringed; anther dehiscence poricidal or - Spathes solitary and becoming leaf-opposed or irregularly
occasionally longitudinal; seeds usually arillate 16 clustered, never distichously arranged on an axis; flowers
- Petals fringed; anther dehiscence always longitudinal; mostly blue, uncommonly other colors, rarely white;
seeds exarillate 17 capsules bi- or triblocular, locules 1-2-seeded
16. Filaments much longer than the anthers; anthers about as 35. Commelina
wide as long, dehiscence longitudinal the whole length of 30. Fruits indehiscent, trilocular 31
the anther 12. Siderasis - Fruits dehiscent, bi- or trilocular 32
- Filaments equal to or shorter than the anthers; anthers 31. Fruits spherical or ellipsoid, lustrous, metallic blue, black
much longer than wide, dehiscence usually poricidal, oc- or lead-colored; seeds usually biseriate; fertile stamens
casionally by short longitudinal slits 13. Dichorisandra 3 or 6 36. Pollia
17. Plants tank or rarely creeping epiphytes, rarely terrestrial; - Fruits cylindric-trigonous, probably brown at maturity;
leaves sword-shaped; inflorescences borne among the seeds uniseriate; fertile stamen 1 33. Pseudoparis
leaves; fertile stamens 3; anthers enclosed within hoodlike 32. Stamens 6, equal, fertile 27. Stanfieldiella
expanded connectives, not visible without dissection - Stamens 6 or fewer, all fertile and unequal, or some
14. Cochliostema staminodial or lacking 33
- Plants rhizomatous, neither tank nor creeping in habit, 33. Fertile stamen 1 33. Pseudoparis
terrestrial; leaves elliptic to ovate or orbicular; inflores- - Fertile stamens 2-6 34
cences borne below the leaves, commonly basal; fertile 34. Fertile stamens 5-6 35
stamens 5-6; anthers not enclosed, although often some - Fertile stamens 2-3 36
surrounded by dense hairs, plainly visible 35. Upper and lower stamens nearly equal, some or all
15. Geogenanthus filament bases fused; capsule bilocular, locules 1-seeded
18. Inflorescences terminal, thyrsiform and/or cincinni 28. Floscopa
elongate 19 - Upper stamens much shorter than the lower, filaments
- Inflorescences terminal or terminal and axillary, cincinnus free; capsules trilocular, locules 1-several-seeded
axis contracted (although cincinni sometimes stipitate)21 32. Tricarpelema
19. Flowers actinomorphic; stamens equal 20 36. Capsules bilocular or unequally trilocular, bivalved;
- Flowers zygomorphic; stamens unequal 16. Tinantia bracteoles usually perfoliate 37
20. Inflorescences thyrsiform 17. Thyrsanthemum - Capsules equally trilocular, trivalved; bracteoles not
- Inflorescences umbelliform 18. Gibasoides perfoliate 38
118 Commelinaceae
37. Capsules glabrous or pubescent, rarely with hooked hairs;
locules often more than 1-seeded; inflorescence usually
thyrsiform 37. Aneilema
- Capsules covered with hooked hairs; locules 1-seeded or
empty; inflorescence corymblike 38. Rhopalephora
38. Stamens antesepalous, alternating with the staminodes 39
- Stamens on one side of the flower, staminodes above or
lacking 40
39. Rosette plants; inflorescences subscapose; filaments all
glabrous; antherodes unlobed 31. Anthericopsis
- Plants of various habits, rarely rosettes with subscapose
inflorescences; some or all filaments usually bearded,
rarely all glabrous; antherodes 3-lobed or hastate
30. Murdannia
40. Stamens 3, staminodes 3; locules usually 2-several-seeded,
rarely 1-seeded 32. Tricarpelema
- Stamens 3, staminodes 0; locules always 1-seeded
35. Dictyospermum
GENERA OF CoMMELINACEAE
I. Subfamily Cartonematoideae (Pichon) Faden
ex G. Tucker (1989).
Subfam. Cartonemoideae Faden in Dahlgren, Clifford & Yeo
(1985), non rite pub!.
Glandular microhairs lacking; raphide canals
lacking or next to the veins; stomata with 2 or
4 subsidiary cells; internodes and/or foliage
glandular-pubescent; flowers actinomorphic,
yellow, petals free; stamens 6, all fertile, equal or
subequal; fruits capsular, trilocular, dehiscent,
shorter than the sepals; seeds exarillate, with a
punctiform hilum and dorsal embryotega.
1. Tribe Cartonemateae (Pichon) Faden &
D.R. Hunt (1991}.
Cartonemataceae Pichon (1946).
Commelinaceae "Group II" Brenan (1966).
Perennials; raphide canals lacking; stomata with
2-4 subsidiary cells; inflorescence a seemingly
spiciform raceme, but individual flowers actually
1-flowered cincinni and the inflorescence there-
fore a thyrse (see Brenan 1966); seeds uniseriate.
Chromosomes small, x = 12 (or 6?). Only one
genus: Cartonema R. Br.
1. Cartonema R. Br.
Cartonema R. Br., Prod.: 271 (1810); Conran, Fl. Australia, vol.
40 (in press).
Perennial or annual herbs, sometimes arising
from tubers; leaves spirally arranged, lamina
linear, sessile, glandular-pubescent; inflores-
cences terminal and axillary, spiciform or
racemiform, but each flower representing a re-
duced, 1-flowered cincinnus; flowers bisexual,
actinomorphic, subsessile (above the bracteole);
sepals subequal, free, glandular-pubescent; petals
equal, free, not clawed, yellow (rarely pink);
stamens 6, equal, filaments glabrous, anthers with
a narrow connective, dehiscence longitudinal;
ovary trilocular, locules 2-several-ovulate; cap-
sules trilocular, trivalved; seeds uniseriate, 2-
several/locule, hilum punctiform, embryotega
dorsal. 2n = 24. Eleven spp., Australia, with 1 sp.
apparently extending to Trangan Island (SW of
New Guinea), Indonesia.
2. Tribe Triceratelleae Faden & Hunt (1991).
Commelinaceae "Group IX" Brenan (1966).
Annuals with sympodial growth; raphide canals
next to the veins; stomata with 2 subsidiary cells;
inflorescence composed of single, leaf-opposed
cincinni, cincinni elongate, several-flowered;
flowers yellow, actinomorphic; seeds numerous,
biseriate. Chromosomes unknown. Only one
genus, Triceratella.
2. Triceratella Brenan
Triceratella Brenan in Kirkia 1: 14 (1961).
Annual herbs; leaves linear, lamina sessile, more
or less glandular-pubescent; inflorescences single,
leaf-opposed, elongate cincinni; flowers bisexual,
subsessile; sepals longer than the petals, subequal,
free, glandular-pubescent; petals subequal, free,
not clawed; stamens 6, equal or subequal, fila-
ments pubescent, anthers with a narrow connec-
tive, dehiscence longitudinal; ovary sessile, locules
with numerous ovules, ovules biseriate; capsules
trilocular, trivalved; seeds biseriate, numerous,
more or less hemispherical, hilum punctiform,
embryotega dorsal. 2n = ? Only one sp., T.
drummondii Brenan, Zimbabwe.
II. Subfamily Commelinoideae Faden &
D.R. Hunt (1991).
Glandular microhairs almost invariably present;
raphide canals present and mostly between
the veins; internodes and foliage almost never
glandular-pubescent; flowers variously colored
but very rarely both yellow and actinomorphic;
stamens, capsules, and seeds various.
 Commelinaceae
3. Tribe Tradescantieae Meisner (1842).
Tribe Hexandreae Bruckner (1926) p. p.
Commelinaceae "Group XI" Brenan (1966).
An exceedingly diverse assemblage recognized as
a group only by the characters given in the key;
silica sometimes present in epidermal cells. Chro-
mosomes very variable, but large in many genera,
less often small to medium-sized, x 5 4-20. Cos-
mopolitan: 25 genera/285 spp.
3a. Subtribe Palisotinae Faden & D.R. Hunt
(1991).
Tribe Triandreae Bruckner (1926) (p. p.).
Commelinaceae "Group III," Brenan (1966) (p. p.).
Rhizomatous perennials; inflorescences thyrses,
cincinni not paired; flowers bisexual and male;
sepals petaline; petals free, not clawed; pollen-
bearing stamens 3, antepetalous, filaments usually
glabrous; staminodes 2-3, antesepalous, lacking
anthers, filaments bearded; fruit a berry; seeds
uni- or biseriate, exarillate, hilum punctiform,
embryotega dorsal. Chromosomes medium-sized,
x = 20. Only one genus, Palisota.
3. Palisota Rchb.
Palisota Rchb. ex Endlicher, Gen. Pl.: 125 (1836).
Perennial herbs, generally medium- to large-sized,
mostly rosettes or herbaceous shrubs, leaves alter-
nate or in pseudowhorls, lamina petiolate; inflo-
rescences terminal, terminal and axillary, or all
axillary, pedunculate, thyrsiform; flowers slightly
zygomorphic, bisexual and male, pedicellate;
sepals subequal; petals subequal; staminodes
2-3, subequal or the posterior one different or
lacking; stamens 3, 2 posterior shorter, pollen ster-
ile, anterior stamen longer, pollen fertile; fruits
berries, trilocular, indehiscent, orange to red or
blue to black, locules 1-6-seeded. 2n = 40, ca. 60,
80. About 18 spp., Africa, in forest understory and
disturbed situations, usually growing in shade.
3b. Subtribe Streptoliriinae Faden & D.R. Hunt
(1991).
Tribe Hexandreae Bruckner (1926) (p. p.).
Commelinaceae "Group IV," Brenan (1966).
Climbing or erect herbs with lamina petiolate;
inflorescences leaf-opposed or axillary thyrses,
lowermost 1-2 cincinni usually subtended by a
119
spathaceous bract; flowers bisexual and male or
all bisexual; sepals free, equal or subequal; petals
equal, free, not clawed, narrower than the sepals;
stamens, 6, equal, or 5 with the posterior 3 shorter,
or 3 and all posterior, some or all filaments
bearded; capsule trilocular, dehiscent; seeds uni-
or biseriate, exarillate, hilum linear, embryotega
dorsal to semidorsal. Chromosomes (unknown in
Aetheolirion) large, x = 5, 6.
4. Streptolirion Edgew.
Streptolirion Edgew. in Proc. Linn. Soc. London 1: 254 (1845).
Annual (or sometimes perennial?), scan dent to
erect herbs; ptyxis involute; inflorescences axillary
or leaf-opposed thyrses, upper cincinni bracteate;
flowers bisexual (lower cincinni) and male (upper
cincinni), subsessile; sepals equal, petaline; sta-
mens 6, free, fertile, filaments bearded, anthers
with a broad connective, dehiscence latrorse; cap-
sule trilocular, trivalved, locules 2-seeded; seeds
uniseriate, hilum linear, embryotega semilateral.
2n = 10, 12, 48. One or 2 spp., Himalayas to Japan.
5. Spatholirion Ridl.
Spatholirion Rid!. in J. Bot. 34: 329 (1896).
Perennial, scandent to erect herbs; ptyxis super-
v?lu~e; ~nflorescences leaf-opposed thyrses, upper
cmcmm ebracteate; flowers bisexual (bracteate
cincinni) or male (ebracteate cincinni), subsessile;
sepals equal, petaline; stamens 6, free, fertile, fila-
ments bearded, anthers with a narrow connective
dehiscence latrorse; capsule trilocular, trivalved:
locules several-seeded; seeds (uni-? or) biseriate,
hilum linear, embryotega dorsal. 2n = 20. Three
spp., Thailand to China.
6. Aetheolirion Forman
Aetheolirion Forman in Kew Bull. 16: 209 (1962).
Scandent herbs; inflorescences leaf-opposed thyr-
ses, upper cincinni bracteate; flowers subses-
sile, bisexual (lower cincinni) and male (upper
cincinni); sepals subequal, sepaline; stamens 3 or
5, 3 posterior with shorter, bearded filaments and
2 anterior (only in male flowers) with longer gla-
brous filaments, connective narrow; ovary with
locules 6-8-ovulate; capsules trilocular, trivalved;
seeds biseriate, winged, hilum linear, embryotega
dorsal. 2n = ? Only one sp., A. stenolobium
Forman, Thailand.
120 Commelinaceae
3c. Subtribe Cyanotinae (Pichon) Faden &
D.R. Hunt (1991).
Tribe Cyanoteae Pichon (1946).
Tribe Hexandreae Bruckner (1926) (p. p.).
Commelinaceae "Group VI," Brenan (1966).
Mostly perennial herbs, frequently with under-
ground storage organs; leaves mainly distichous,
lamina sessile, succulent, with a hypodermis
beneath the upper epidermis; inflorescences
mostly consisting of single or aggregated cincinni
subtended by small or more or less foliaceous
bracts; bracteoles large, herbaceous; flowers
bisexual, actinomorphic; petals equal, free or
united below into a tube; stamens 6, equal,
fertile, filaments bearded, sometimes swollen
subterminally; ovary with locules biovulate,
style sometimes bearded; capsules trilocular,
trivalved, locules (1-)2-seeded; seeds uniseriate,
exarillate, hilum punctiform, embryotega termi-
nal. Chromosomes small to medium-sized, x = 8,
9?, 10-14.
7. Belosynapsis Hassk.
Belosynapsis Hassk. in Flora 54: 259 (1871).
Perennial herbs, sometimes epiphytic; inflores-
cences terminal and axillary, consisting of 1-2
cincinni; bracts small or large, persistent; cincinni
!-several flowered; flowers bisexual, subsessile
or pedicellate; sepals equal; petals free, not
clawed; stamens 6, free, filaments not swollen
apically, anther dehiscence longitudinal; style
not swollen apically. 2n = 36, 40, 52. Four spp.,
Madagascar, and India to New Guinea.
8. Cyanotis D. Don Fig. 32
Cyanotis D. Don, Prod, Fl. Nep.: 45 (1825).
Amischophacelus R.S. Rao & Kammathy (1966).
Perennial or occasionally annual herbs; inflores-
cences terminal and/or axillary, each cincinnus or
group of cincinni usually subtended by a folia-
ceous bract, rarely the inflorescences enclosed
in the leaf sheaths; flowers subsessile; sepals
subequal; petals united below into a tube; stamens
6, filaments usually with a subapical swelling, an-
thers with longitudinal dehiscence but pollen usu-
ally released from base of anther sacs; style usually
with a subapical swelling. 2n = 16, 20, 22, 24, 26,
28(?), 48, 52, 72, 78. Fifty spp., paleotropics, grassy,
rocky or other open habitats, rarely aquatic
situations, or in forest.
3d. Subtribe Coleotrypinae Faden & D.R. Hunt
(1991).
Tribe Hexandreae Bruckner (1926) (p. p.).
Commelinaceae "Group 1," p. p., and "Group V," Brenan
(1966).
Perennial, mostly decumbent, occasionally scan-
dent herbs; leaves spirally arranged, lamina
petiolate, leaf epidermal cells with silica; inflores-
cences axillary and perforating the sheaths, sessile
or subsessile; cincinni usually congested and con-
tracted; bracts and sometimes bracteoles herba-
ceous; flowers usually all bisexual, actinomorphic;
petals free or fused basally into a tube; stamens 6,
fertile, equal or unequal; capsules trilocular,
trivalved, locules 1-2-seeded; seeds uniseriate,
arillate, or exarillate, with a linear hilum and lat-
eral embryotega. Chromosomes (unknown m
Porandra) large, x = 9, and possibly others.
9. Coleotrype C.B. Clarke
Coleotrype C.B. Clarke in DC., Monogr. Phan. 3: 238 (1881).
Perennial herbs; ptyxis involute; inflorescences
sessile, consisting of a few congested and con-
tracted cincinni each subtended by a large, ovate
bract; flowers sessile; sepals subequal, sepaline;
petals equal or unequal, united below into a nar-
row tube; stamens equal or unequal, filaments
adnate to the corolla tube, sometimes connate,
bearded or glabrous, anthers dehiscing longitudi-
nally or by apical pores; capsules hidden among
the bracts and sepals. 2n = 36, 42. Nine spp.,
Africa, Madagascar, in forest understory.
10. Amischotolype Hassk.
Amischotolype Hassk. in Flora 46: 391 (1863).
Perennial herbs; ptyxis involute; inflorescences
very compact and globose or more lax, composed
of 2-several cincinni; flowers subsessile to shortly
pedicellate; sepals subequal, sepaline or colored;
petals free, equal, not clawed; stamens 6, equal or
subequal, filaments bearded or subglabrous, an-
thers dehiscing longitudinally or by apical pores;
seeds embedded in a red aril, hilum linear,
embryotega lateral. 2n = 18, 20(?), 30(?), 36. Fif-
teen spp., Central Africa and India to New Guinea,
in forest understory.
11. Porandra Hong
Porandra Hong in Acta Phytotax. Sin. 12: 462 (1974).
 Commelinaceae
Perennial scandent herbs; inflorescences sub-
sessile, very compact, globose; flowers bisexual,
subsessile; sepals equal, sepaline; petals free, equal,
not clawed; stamens 6, free, equal, filaments
bearded, anthers dehiscing by apical pores; seeds
arillate. 2n = ? Three spp., China to Thailand, in
forests. Questionably distinct from Amischotolype.
3e. Subtribe Dichorisandrinae (Pichon) Faden &
D.R. Hunt (1991).
Tribes Dichorisandreae, Geogenantheae, and Cochlioste-
mateae Pichon (1946).
Tribe Inclinatae Bruckner (1926).
Tribe Hexandreae Bruckner (1926) (p. p.).
Commelinaceae "Group I p. p., XIII p. p., and XIV, XV,"
Brenan (1966).
Perennials of diverse habit; ptyxis involute; inflo-
rescences thyrses or sometimes reduced to single
cincinni, terminal and/or axillary, sometimes
basal; flowers bisexual or bisexual and male, zygo-
morphic or actinomorphic; sepals free, sepaline;
petals free, equal to somewhat unequal, not
clawed, margins sometimes fringed; stamens
(3- )5-6, equal or unequal, when 3, all posterior,
anther dehiscence longitudinal or by apical pores;
staminodes 0-3; ovary trilocular; fruit capsular,
dehiscent; seeds arillate or exarillate, usually
biseriate, hilum slightly elongate to linear,
embryotega dorsal to lateral. Chromosomes large,
X= 19.
12. Siderasis Raf.
Siderasis Raf., Fl. Tellur. 3: 67 (1837).
Perennial herbs, sometimes climbing; inflores-
cence terminal, consisting of a single cincinnus or
thyrsiform; flowers bisexual, actinomorphic, pedi-
cellate; sepals equal, sepaline; petals free, equal,
glabrous; stamens 6, free, equal, filaments gla-
brous, anther dehiscence latrorse; capsules
trilocular, locules unequal, dorsal locule 0-1-
seeded, ventrallocules more than 2-seeded; seeds
biseriate, exarillate. 2n = 38. Two or 3 spp., Brazil,
in forest understory.
13. Dichorisandra Mikan
Dichorisandra Mikan, Del. Fl. et Faun. Bras. 1: tab. 3 (1820).
Perennial herbs of diverse habit; roots usually
with distal tubers; inflorescences terminal, axillary
or basal, thyrsiform or reduced; flowers bisexual
and male, usually slightly zygomorphic; sepals
121
subequal; petals free, subequal, glabrous; stamens
5-6, equal or unequal, filaments short, glabrous,
anthers large, thick, dehiscing by terminal pores
or longitudinal slits; capsules trilocular, trivalved;
seeds biseriate, embedded in a red (rarely white)
aril. 2n = 38, 76. Twenty-five spp., neotropics, in
forest understory.
14. Cochliostema Lem.
Cochliostema Lem. in Illus. Hort. 6, Misc.: 70, tab. 217 (1859);
Read, Baileya 13: 8-15 (1965).
Tank or creeping epiphytes (rarely terrestrial);
inflorescences axillary thyrses with cincinni
subtended by large, colored bracts; flowers
zygomorphic, fragrant; sepals subequal; petals
unequal, margins fringed; staminodes 3, anterior,
medial one vestigial, lateral ones densely bearded,
lacking an anther; stamens 3, posterior, filaments
fused, with a dense dorsal tuft of yellow hairs,
anthers enclosed within a columnlike structure
composed of the broadly expanded connectives of
the lateral stamens, the connectives each termi-
nally drawn out into a narrow tube with a terminal
pore; seeds exarillate, biseriate, hilum linear,
embryotega dorsal. 2n = 38. Two spp., Nicaragua
to Ecuador, in forest.
15. Geogenanthus Ule
Geogenanthus Ule in Feddes Repert. 11: 524 (1913).
Perennial herbs; roots with distal tubers; shoots
determinate, produced from subterranean rhi-
zomes; inflorescences basal, perforating bladeless
sheaths, consisting of 1-2( -4) cincinni; flowers
slightly zygomorphic; sepals equal; petals equal or
subequal, margin fringed; stamens 5-6, posterior
3 shorter, filaments usually bearded, anterior sta-
mens 2-3, filaments usually glabrous, anther de-
hiscence introrse; ovary with locules 4-6-ovulate;
seeds with hilum linear, embryotega lateral.
2n = 38. Five spp., western S America, in forest
understory.
3f. Subtribe Thyrsantheminae Faden &
D.R. Hunt (1991).
Tribe Hexandreae p. p. and Declinatae p. p. Bruckner (1926).
Commelinaceae "Group I" p. p. and "Group XII" p. p. Brenan
(1966).
Inflorescences thyrsiform or reduced sometimes
to single cincinni, terminal and axillary, rarely all
122 Commelinaceae
axillary; cincinni never fused in pairs; flowers acti-
nomorphic or zygomorphic; petals free or fused
into a tube; stamens 6, fertile, equal or unequal,
anther dehiscence longitudinal; ovary trilocular;
fruit capsular, trilocular, trivalved; seeds exaril-
late, hilum linear to punctiform, embryotega
dorsal to lateral. Chromosomes (unknown in
Elasis) small to medium-sized, x = 9-17.
16. Tinantia Scheidw.
Tinantia Scheidw. in Otto and Dietrich, Allg. Gartenz. 7: 365
(1839).
Annuals herbs; ptyxis involute; inflorescences
paniculiform or umbelliform thyrses or composed
of 1-2 cincinni; cincinni elongate; flowers zygo-
morphic; sepals free; petals free, slightly to
strongly unequal, not clawed; stamens 6, fertile,
polymorphic, filaments fused basally, posterior 3
stamens shorter with densely bearded filaments,
anterior 3 with longer filaments, the lateral
ones bearded, the medial one glabrous; seeds
uniseriate, 2-several per locule, hilum linear,
embryotega lateral. 2n = 26, 28, 32, 34, 64, 66-68,
ca. 128. Thirteen spp., Texas to neotropics.
17. Thyrsanthemum Pichon
Thyrsanthemum Pichon in Not. Syst. (Paris) 12: 224 (1946);
Hunt, Kew Bull. 31: 407-410 (1976), rev.
Erect or decumbent, perennial herbs; roots tuber-
ous; ptyxis supervolute; inflorescence a thyrse
or panicle of thyrses, cincinni several- to many-
flowered; flowers bisexual, actinomorphic, sub-
sessile to shortly pedicellate; sepals free, subequal;
petals free, equal, not clawed; stamens 6, free,
equal, filaments bearded; seeds uniseriate, 2 per
locule, hilum linear, embryotega lateral or semi-
lateral. 2n = 28, 30, 32. Three spp., Mexico.
18. Gibasoides D.R. Hunt
Gibasoides D.R. Hunt in Kew Bull. 33: 331 (1978).
Perennial herbs; roots tuberous; ptyxis super-
volute; inflorescence terminal, umbelliform, glan-
dular-pubescent, consisting of numerous elongate
cincinni, each with a long cincinnus peduncle not
geniculate at the apex, and an elongate cincinnus
axis with numerous flowers; flowers actinomor-
phic, pedicellate; sepals subequal, free; petals free,
equal, not clawed; stamens 6, free, subequal,
filaments bearded; seeds uniseriate, 2 per locule,
hilum linear, embryotega lateral. 2n = 32, 48. Only
one sp., G. laxiflora (C.B. Clarke) D.R. Hunt,
Mexico.
19. Weldenia Schult.f.
Weldenia Schult.f. in Flora 12: 3, tab. 1A (1829).
Rosette perennial herbs; shoot vertical, subterra-
nean; roots tuberous; ptyxis supervolute; inflores-
cence a terminal, sessile, congested thyrse in the
center of the rosette, consisting of numerous
sessile cincinni, the outer ones bracteate, the inner
ebracteate; flowers actinomorphic, pedicellate; se-
pals united into a tube; petals united into a long,
slender tube, with equal, spreading lobes; stamens
6, equal, epipetalous, filaments glabrous; seeds
uniseriate, several per locule, hilum punctiform,
embryotega lateral to semilateral. 2n = 18, 20, 24.
Only one sp., W. candida Schult. f., mountains of
Mexico and Guatemala.
20. Elasis D.R. Hunt
Elasis D.R. Hunt in Kew Bull. 33: 332 (1978).
Procumbent or scandent perennial herbs rooting
at the nodes; roots thin, fibrous; ptyxis super-
volute; inflorescence consisting of up to 5 simple
cincinni, terminal, and solitary from the upper
axils, forming a loose cluster; flowers actinomor-
phic; sepals subequal, free, glandular-pubescent;
petals free, equal, not clawed; stamens 6, subequal,
filaments bearded; seeds uniseriate, 2 per locule;
hilum linear, embryotega dorsal. 2n = ? Only one
sp., E. hirsuta (Kunth) D.R. Hunt, high elevations,
Ecuador.
21. Matudanthus D.R. Hunt
Matudanthus D.R. Hunt in Kew Bull. 33: 333 (1978).
Perennial herbs with annual, unbranched or
sparsely branched shoots; roots tuberous; ptyxis
supervolute; inflorescences composed of ter-
minal and axillary, solitary, sessile, contracted, 1-
3-flowered cincinni; flowers actinomorphic,
long-pedicelled; sepals equal; petals equal, not
clawed; stamens 6, subequal, filaments bearded;
seeds uniseriate, 2 per locule. 2n = 32. Only
one sp., M. nanus (Martens & Gal.) D.R. Hunt,
Mexico.
 Commelinaceae
3g. Subtribe Tradescantiinae Rohweder {1956).
Tribes Hexandreae p. p. and Triandreae p. p. BrUckner (1926}.
Tribes Callisieae p. p. and Zebrineae p. p. Pichon (1946}.
Subtribe Gibasinae Rohweder (1956).
Commelinaceae "Groups XI, XII, and XIII" Brenan (1966}.
Perennials or annuals; inflorescences composed of
pairs of sessile, contracted cincinni fused back to
back or of pairs or pseudo-umbels of stipitate
cincinni; flowers actinomorphic or zygomorphic;
sepals free, subequal; petals free or fused basally,
subequal, clawed or not; stamens 0-6, when 6, all
equal or alternately unequal with the antesepalous
stamens always fertile, and the antepetalous ones
fertile or staminodial, anther connectives broad,
dehiscence longitudinal; fruits capsular, locules
1-2-seeded; seeds exarillate, uniseriate, hilum
punctiform to linear, embryotega dorsal to lateral.
Chromosomes mostly large (occasionally small to
medium-sized), x = 4-13.
22. Gibasis Raf.
Gibasis Raf., Fl. Tellur. 2: 16 (1837}, Hunt, Kew Bull. 41: 107-
129 (1986}, rev.
Annual or perennial herbs; roots fibrous or tuber-
ous; ptyxis involute or supervolute; inflorescences
terminal, composed of pairs or pseudo-umbels of
stipitate cincinni, cincinnus axis sharply angled at
junction with cincinnus peduncle; flowers actino-
morphic; petals free, equal, not clawed; stamens 6,
equal, fertile, filaments bearded; capsules trilo-
cular, trivalved; seeds uniseriate, 2 per locule,
hilum elongate-punctiform to linear, embryotega
dorsal. 2n = 10, 12, 15, 16, 20, 22, 30, 32, 48. Eleven
spp., neotropics, mainly in Mexico.
23. Tradescantia L.
Tradescantia L., Sp. Pl.: 288 (1753}; Hunt, Kew Bull. 30: 443-
458 (1975} (Setcreasea and Separotheca reduced}, ibid. 35:
437-442 (1980} (sectional class.}, ibid. 41: 401-405 (1986}
( Campelia, Rhoeo and Zebrina reduced).
Campelia L.C. Rich. (1808}.
Zebrina Schnizl. (1849}.
Rhoeo Hance (1853).
Setcreasea K. Schum. & Sydow (1901}.
Cymbispatha Pichon (1946}.
Separotheca Waterfall (1959}.
Perennial (rarely annual) herbs of various habits;
ptyxis involute or supervolute; inflorescences ter-
minal and/or axillary, composed of pairs of sessile
cincinni subtended by spathaceous or foliaceous
bracts; flowers bisexual, actinomorphic; petals
123
free or fused basally, not clawed (rarely clawed);
stamens 6, equal or slightly unequal, filaments
bearded or glabrous; capsules trilocular, trivalved;
seeds 1-2 per locule, hilum oblong to linear,
embryotega dorsal to lateral. 2n = 12, 14, 16, 18,
21-30,32,35,36,38,40,47,48,50,60,64,67,70,72,
74, 76, 90, 92, 108-110, 114, 130, 132, 140, 144.
About 70 spp., New World.
24. Callisia Loefl.
Callisia Loefl., Iter Hispan.: 305 (1758); Moore, Baileya 6: 135-
147 (1958}; Hunt, Kew Bull. 41: 407-412 (1986}.
Hadrodemas H.E. Moore (1963}.
Leptorhoeo C.B. Clarke & W.B. Hemsley (1880).
Cuthbertia Small (1903}.
Perennial or annual herbs; roots thin, rarely tuber-
ous; ptyxis supervolute or conduplicate; inflores-
cences terminal and/or axillary, composed of pairs
of sessile cincinni subtended by small bracts
usually less than 1 em long, often aggregated into
compound inflorescences; flowers actinomorphic,
bisexual, rarely bisexual and female, subsessile or
pedicellate; petals free, not clawed; stamens equal
or subequal, 6 or 3-0, filaments glabrous or
bearded; capsules tri-or bilocular, tri-or bivalved,
locules 1-2-seeded; seeds with hilum punctiform,
embryotega dorsal. 2n = 12, 14, 16, 24, 28, 32, 36,
48, 56, 72. About 20 spp., New World.
25. Tripogandra Raf.
Tripogandra Raf., Fl. Tellur. 2: 16 (1837}; Handlos, Rhodora
77: 213-333 (1975}, rev.
Perennial or annual herbs; roots fibrous; ptyxis
supervolute; inflorescences composed of pairs of
sessile cincinni subtended by small bracts; flowers
bisexual, moderately zygomorphic; petals free, not
clawed; stamens 6, dimorphic, the 3 antesepalous
shorter and fertile, the 3 antepetalous longer and
fertile or staminodial, filaments of 1 whorl, rarely
both, bearded, rarely both glabrous; capsules
trivalved, trilocular, locules 1-2-seeded; seeds
with hilum punctiform to linear, embryotega
dorsal. 2n = 16, 24, 28, 30, 32, 39, 42, 48, 50, 52, 60,
62, 64, 65, 76. Twenty-two spp., neotropics.
3h. Subtribe Uncertain
26. Sauvallea Wright
Sauvallea Wright in Sauv., Fl. Cuba: 156 (1873).
Apparently a decumbent perennial; leaves disti-
chous, lamina sessile to subpetiolate; inflores-
124 Commelinaceae
cence terminal, cons1stmg of a solitary spathe
enclosing a single flower; flowers bisexual; sepals
3, subequal; petals 3, subequal, not clawed; sta-
mens 6, equal, filaments bearded, anthers with a
broad connective, dehiscence longitudinal; ovary
biloc-ular, locules 2-ovulate; capsules and seeds
unknown. 2n = ? Only one sp., S. blainii Wight,
Cuba.
The spathaceous bracts subtending solitary
flowers make the placement of this genus uncer-
tain. It appears to be a reduced member of either
the Thyrsantheminae or Tradescantiinae.
4. Tribe Commelineae Meisner (1842).
Tribe Pollieae C.B. Clarke (1881) (p. p.).
Tribes Hexandreae, Triandreae and Declinatae Bruckner
(1926) (all p. p.).
Tribes Anthericopsideae (p. p.) and Pseudoparideae Pichon
(1946).
Commelinaceae "Group I," (p. p.), "Group III" except
Palisota, and "Groups VII, VIII, and X," Brenan (1966).
As a tribe, characterized by the features given in
the key; silica never present in epidermal cells.
Chromosomes (unknown in Pseudoparis and
Tricarpelema) usually small, occasionally
medium-sized, x = 6-17, 29, several genera with
multiple basic numbers. Cosmopolitan, 13 genera/
348 species.
27. Stanfieldiella Brenan
Stanfieldiella Brenan in Kew Bull. 14: 283 (1960).
Perennial herbs; leaves spirally arranged, lamina
petiolate, ptyxis involute; inflorescences terminal
and/or axillary thyrses or reduced to a single
cincinnus; flowers actinomorphic, pedicellate; se-
pals free, subequal, sepaline, usually glandular-
pubescent; petals free, equal, not clawed; stamens
6, subequal, filaments glabrous; capsules trilo-
cular, trivalved; seeds uniseriate, 2-10 per locule,
hilum linear, embryotega lateral. 2n = 22, 24(?),
44. Four spp., Africa, in forest understory, often in
disturbed places or in shaded cultivation.
28. Floscopa Lour.
Floscopa Lour., Fl. Cochinch.: 192 (1790).
Perennial or annual herbs; leaves spirally ar-
ranged, lamina petiolate or sessile, ptyxis involute;
inflorescences terminal and axillary thyrses (or
reduced to !-several cincinni), commonly form-
ing compound inflorescences; flowers zygomor-
phic, shortly pedicellate; sepals free, subequal,
sepaline, usually glandular-pubescent; petals free,
unequal, not clawed, posterior petals broader than
the anterior petal; stamens 6, fertile, posterior 3
different in form from the anterior 3, filaments
fused basally, glabrous; capsules bilocular, bi-
valved, locules 1-seeded; seeds usually ribbed or
tuberculate, hilum linear, embryotega lateral. 2n
= 12, 16, 18, 22, 24, 30, 36, 42, 54. Twenty spp.,
pan tropical, mostly in aquatic situations, either in
forest or open habitats.
29. Buforrestia C.B. Clarke
Buforrestia C.B. Clarke in D.C., Monogr. Phan. 3: 233, tab. 6, 7
(1881); Brenan, Kew Bull. 14: 280-286 (1960).
Perennial herbs; roots fibrous; leaves spirally
arranged, lamina petiolate, ptyxis involute; inflo-
rescences axillary, perforating the sheaths, con-
tracted or elongate, composed of 1-5, sessile,
contracted cincinni; flowers zygomorphic, pedi-
cellate; sepals free, subequal, greenish white,
accrescent in fruit; petals free, unequal, posterior
petals narrower than the anterior petal; stamens
6, fertile, polymorphic, posterior 3 shorter
than the anterior 3, filaments glabrous; capsules
trilocular, trivalved, locules ca. 4-10-seeded; seeds
uniseriate, hilum linear, embryotega lateral. 2n =
34. Three spp., in forest understory, 2 in W and
Central Africa, 1 in northeastern S America.
30. Murdannia Royle Fig. 33
Murdannia Royle, Illus. Bot. Himal.: 403, tab. 95 (1839);
Brenan, Kew Bull. 7: 179-208 (1952); Faden & Inman in van
der Maesen, L.J.G., van der Burgt, X.M., van Medenbach de
Rooy, J.M. (eds) The biodiversity of African plants, pp. 464-
471. Dordrecht: Kluwer. (1996) (leaf anat.).
Perennial or annual herbs; roots fibrous or tuber-
ous; leaves spirally arranged, lamina sessile, ptyxis
supervolute; inflorescences terminal and/or axil-
lary thyrses or variously reduced; flowers actino-
morphic to somewhat zygomorphic, pedicellate;
sepals free, subequal, sepaline; petals free,
subequal, not clawed; filaments generally free; fer-
tile stamens 2-3, antesepalous, filaments glabrous
or bearded; staminodes 3( -4), rarely lacking,
antepetalous, (when 4, one antesepalous),
antherodes 3-lobed or hastate; capsules (bi-)
trilocular, (bi-) trivalved, locules 1-many-seeded;
seeds uni- or biseriate, hilum slightly elongate to
linear, embryotega dorsal to semilateral. 2n = 12,
18,20,22,24,30,32,36,40,42,44,60,64,80.Fifty
 Commelinaceae
spp., pan tropical and war~ temperate, ~n a va_riety
of open, mesic, or occasiOnally aquatiC habitats,
rarely in forests.
31. Anthericopsis Engl.
Anthericopsis Engl., Pflanzenw. Ost-Afr., C: 139 (1898); Faden
& Inman. (1996), cit. under 30. Murdannia (leaf anat.).
Rosette perennial herbs dormant in the dry sea-
son; roots with distal tubers; leaves spirally
arranged, lamina sessile, ptyxis supervolute; inflo-
rescences scapose or subscapose, terminal and
axillary, composed of 1-2 cincinni at the scape
summit; flowers actinomorphic, pedicellate; se-
pals free, equal, sepaline; petals free, equal, not
clawed; stamens 3, antesepalous, free, equal, fila-
ments glabrous; staminodes, 3, antepetalous, free,
equal, filaments glabrous, antherode~ u~lobed;
capsules trilocular, trivalved; seeds umsenate, 8.-
12 per locule, hilum linear, embryotega semi-
lateral. 2n = 14. Only one sp., A. sepalosa (C.B.
Clarke) Engl., eastern Africa, in grassland and
bushland.
32. Tricarpelema J.K. Morton
Tricarpelema J.K. Morton in J. Linn. Soc., Bot. 59: 436 (1966).
Perennial herbs; roots fibrous; leaves usually spi-
rally arranged (rarely distichous), lamina usually
petiolate (rarely sessile), ptyxis usually involute;
inflorescences terminal and axillary thyrses;
flowers zygomorphic, pedicellate; sepals free,
subequal, sepaline; petals free, the posterior 2
slightly to strongly differentiated from the ante-
rior one, clawed or not clawed; stamens 6, poste-
rior 3 (sometimes staminodial?) usually shorter
than anterior 3, filaments usually glabrous; cap-
sules trilocular, trivalved, locules 1-9-seeded;
seeds uniseriate, hilum linear, embryotega lateral
to semidorsal. 2n = ca. 46. Eight spp., one in
Central Africa and 7 from NE India to the
Philippines and Borneo, usually in the forest
understory. .
A single African species, as yet undescnbed,
from Cameroon and Gabon, is tentatively in-
cluded here. It differs from the Asian species in
having succulent, distichous leaves with a sessile
lamina and in growing in exposed rocky areas.
33. Pseudoparis H. Perrier
Pseudoparis H. Perrier, Not. Syst. (Paris) 5: 176 (1936), and in
Humbert, H. (ed.), Flore de Madagascar, Commelinacees,
Imprimerie Officielle, Tananarive (1938).
125
Perennial herbs; roots tuberous; leaves spirally ar-
ranged, lamina petiolate or subpetiolate; inflores-
cences terminal or axillary thyrses; flowers
pedicellate; sepals subequal, sepaline; petals free,
subequal, not clawed; staminodes 3, posterior,
filaments glabrous, antherodes 2-3-lobed, stamen
1, anterior, antepetalous, filament glabrous; ovary
3-locular, locules 4-8-ovulate; fruit dehiscent (or
indehiscent?), trilocular, locules 1-6-seeded; seeds
uniseriate, hilum linear. 2n = ? Two to 3 spp.,
Madagascar.
34. Polyspatha Benth.
Polyspatha Benth. in Hook., Niger. Fl.: 543 (1849).
Perennial stoloniferous herbs; roots fibrous;
leaves spirally arranged, lamina petiolate, ptyxis
involute; inflorescences terminal and axillary,
each inflorescence a modified thyrse with disti-
chous, sessile spathes on an elongate axis, each
spathe enclosing 1(-3) sessile, contracted cin-
cinnus; flowers zygomorphic, shortly pedicellate;
sepals free, subequal, sepaline; 2 posterior petals
clawed, anterior petal smaller, not clawed; fila-
ments fused basally, glabrous; staminodes 3, pos-
terior; stamens 3, anterior; capsules bilocular,
bivalved, locules 1-seeded; seeds ribbed, hilum
linear, embryotega semilateral. 2n = 28. Three
spp., Africa, in forest understory and disturbed,
shady situations.
35. Dictyospermum Wight
Dictyospermum Wight, Icon. Pl. Ind. Orient. 6: 29, tab. 2069-
2071 (1853).
Perennial herbs; roots fibrous or tuberous; leaves
spirally arranged, lamina petiolate; ptyxis invo-
lute; inflorescences thyrses, terminal and axillary,
bracts and bracteoles caducous; flowers slightly
zygomorphic; sepals free, sepaline; petals free,
equal, not clawed; stamens 3, 1-sided, attached in
front of the outer sepal and inner petals, filaments
free or shortly fused basally, glabrous; staminodes
O( -2); capsules trilocular, trivalved, locules 1-
seeded; seeds with hilum linear, embryotega lat-
eral. 2n = 28. About five spp., India and Sri Lanka
to New Guinea, in forest understory.
36. Pollia Thunb.
Pollia Thunb., Gen. Nov. 1: 11 (1781).
Aclisia Hassk. (1870).
126 Commelinaceae
Stoloniferous or rhizomatous perennial herbs;
roots fibrous; leaves spirally arranged, lamina
petiolate, ptyxis involute; inflorescences thyrses,
terminal; flowers actinomorphic to slightly
zygomorphic, pedicellate; sepals free, subequal,
sepaline or petaline; petals free, equal or posterior
2 slightly differentiated from the anterior one,
sometimes shortly clawed; stamens 6, equal, fertile,
or posterior 3 shorter than the anterior 3 and either
fertile or staminodial, filaments free, glabrous;
fruits berrylike, hard and crustaceous, indehiscent,
trilocular, usually lustrous metallic blue; seeds
(uni- or) biseriate, (1-)3-8 per locule, hilum ±
punctiform, embryotega dorsal. 2n = 10(?), 30, 32,
38(?). Seventeen spp., pantropical, and warm tem-
perate, in forest understory and moist, shaded dis-
turbed places; known in the Western Hemisphere
from a single collection from Panama.
37. Aneilema R. Br. Fig. 34
Aneilema R. Br., Prod.: 270 (1810); Morton, J. Linn. Soc.,
Bot. 59:431-478 (1966), rev.; Faden, Smithson. Contrib. Bot.
1-166 (1991), rev.
Perennial or annual herbs; roots fibrous or tuber-
ous; ptyxis usually involute; inflorescences ter-
minal and axillary (rarely all axillary) thyrses
(sometimes reduced to a single cincinnus); flowers
strongly zygomorphic, pedicellate; sepals free, un-
equal, sepaline; petals free, posterior 2-clawed, an-
terior petal not clawed; staminodes 2-3 posterior,
shorter than the stamens, filaments glabrous,
antherodes bilobed; stamens 3, anterior, filaments
free or basally fused, glabrous or 2 bearded;
capsules (uni- ), bi- or trilocular, bivalved, (rarely
indehiscent), locules unequal, 1-6-seeded; seeds
uniseriate, hilum linear, embryotega lateral.
2n = 18,20,26,28,30,32,36,40(?),52,60,64, 78,
90, ca. 104. Sixty-four spp., pantropical but mainly
tropical African, of various habitats, but mainly
bushland, woodland, and grassland, occasionally
forest.
38. Rhopalephora Hassk.
Rhopalephora Hassk. in Bot. Zeit. 222: 58 (1864); Faden,
Phytologia 37: 479-481 (1977), rev.
Perennial herbs; roots fibrous; leaves distichous or
spirally arranged, lamina usually petiolate, ptyxis
involute; inflorescences terminal and axillary,
corymblike thyrses composed of several to many
elongate cymes; flowers zygomorphic, pedicel-
late; sepals free, sepaline; petals free, subequal,
posterior 2-clawed, anterior one not clawed;
staminodes (0- )2-3, posterior, filaments glabrous,
antherodes bilobed; stamens anterior, filaments
fused basally, glabrous; capsules stipitate, 1- or 3-
locular, bivalved, "sticky" -pubescent, locules 0-
1( -2)-seeded; seeds with hilum linear, embryotega
lateral. 2n = 58. Four spp., Madagascar, and India
to Fiji.
39. Commelina L.
Commelina L., Sp. Pl.: 40 (1753).
Phaeosphaerion Hassk. (1866).
Commelinopsis Pichon (1946).
Perennial or annual herbs; roots usually fibrous;
leaves usually distichous, ptyxis involute or
supervolute; inflorescences terminal and leaf-
opposed, composed of 1-2 cincinni enclosed in a
folded spathe; flowers strongly zygomorphic,
pedicellate; sepals free or anterior 2 fused; petals
free, unequal, posterior 2 larger and clawed, ante-
rior petal usually reduced; filaments free, gla-
brous; staminodes 2-3, posterior, antherodes
usually 4-6-lobed; stamens 3, anterior, longer
than staminodes; capsules (uni-) bi- or trilocular,
bi- or trivalved, locules 0-2-seeded; seeds
uniseriate, hilum linear, embryotega lateral. 2n =
16(?), 18(?), 22, 24, 26, 28, 30, 32(?), 36(?), 40(?), 42,
44-46, 48, 52, 54(?), 56, 58(?), 60, 61, 64, 66, 70(?),
72(?), 75, 76(?), 80(?), 84, c. 86, 88, 90, 98(?), 104,
120, 150, ca. 180. About 170 spp., cosmopolitan, of
diverse habitats, but seldom of forests.
40. Tapheocarpa Conran
Tapheocarpa Conran in Austral. Syst. Bot. 7: 585 (1994).
Perennial, creeping aquatic herbs; leaves appar-
ently distichous, lamina sessile; flowers solitary,
terminal, becoming leaf-opposed, ebracteate, bi-
sexual, slightly zygomorphic, pedicellate; sepals 2
fused, 1 free, sepaline; petals free, subequal, free,
not clawed; staminodes 3, posterior, filaments gla-
brous, antherodes cruciform; stamens 3, anterior,
filaments glabrous; ovary trilocular, the dorsallo-
cule abortive, the ventral !-ovulate; fruit indehis-
cent, borne on an elongate pedicel, deeply buried,
echinate; seeds one per ventrallocule. 2n = ? Only
one sp., T. calandrinoides (F. Muell.) Conran
Australia (Queensland), in ephemeral swamps
and billabongs.
Selected Bibliography
Brenan, J.P.M. 1966. The classification of Commelinaceae. J.
Linn. Soc. Bot. 59: 349-370.
 Commelinaceae
Bruckner, G. 1926. Beitrage zur Anatomie, Morphologie und
Systematik der Commelinaceae. Bot. Jahrb. Syst. 61, Beibl.
137: 1-70.
Bruckner, G. 1930. Commelinaceae. In: Engler, A. (ed.) Die
natiirlichen Pflanzenfamilien, 2nd edn, 15a. W. Engelmann
Leipzig, pp. 159-181.
Chase, M.W. et al. 1993. See general references.
Chase, M.W. et al. 1995. See general references.
Chase, M. W. et al. 1995a. See general references.
Clarke, C.B. 1881. Commelinaceae. In: De Candolle, A., De
Candolle, C. (eds.) Monographiae Phanerogamarum, T. 3,
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Dahlgren, R.M.T., Clifford, H.T. 1982. See general references.
Dahlgren, R.M.T., Clifford, H.T., Yeo, P.F. 1985. See general
references.
Duvall, M.R., Clegg, M.T., Chase, M.W., Clark, W.D., Kress,
W.J., Hills, H.G., Eguiarte, L.E., Smith, J.P., Gaut, B.S.,
Zimmer, E.A., Learn, G.H. Jr. 1993. Phylogenetic hypotheses
for the monocotyledons constructed from rbcL sequence
data. Ann. Mo. Bot. Gard. 80: 607-619.
Evans, T.M. 1995. A phylogenetic analysis of the Com-
melinaceae based on morphological and molecular data. Ph
D Dissertation, University of Wisconsin, Madison.
Faden, R.B. 1983. Isolating mechanisms among five sympatric
species of Aneilema R. Br. (Commelinaceae) in Kenya.
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Faden, R.B. 1985. Commelinaceae. In: Dahlgren, R.M.T.,
Clifford, H.T., Yeo, P.F. (eds.) The families of monocotyle-
dons. pp. 381-387. Berlin: Springer.
Faden, R.B. 1988a. Vegetative and reproductive features of
forest and nonforest genera of African Commelinaceae.
Monogr. Syst. Bot. Mo. Bot. Gard. 25: 521-526.
Faden, R.B. 1988b. Reproductive biology of Palisota Reichb.,
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Faden, R.B. 1991. The morphology and taxonomy of Aneilema
R. Brown (Commelinaceae). Smithson. Contrib. Bot. 76: 1-
166.
Faden, R.B. 1992. Floral attraction and floral hairs in the
Commelinaceae. Ann. Mo. Bot. Gard. 79: 46-52.
Faden, R.B. 1993. Murdannia cryptantha (Commelinaceae), a
new species with cleistogamous flowers from Australia and
Papua New Guinea. Novon 3: 133-136.
Faden, R.B. 1995. Commelinaceae. In: Thulin, M. (ed.) Flora of
Somalia, Vol. 4, pp. 79-94. Royal Botanic Gardens, Kew.
Faden, R.B., Hunt, D.R. 1991. The classification of the
Commelinaceae. Taxon 40: 19-31.
Faden, R.B., Suda, Y. 1980. Cytotaxonomy of Commelinaceae:
chromosome numbers of some African and Asiatic species.
Bot. J. Linn. Soc. 81: 301-325.
Grootjen, C.J. 1983. Development of ovule and seed in
Cartonema spicatum R. Br. (Cartonemataceae). Aust. J. Bot.
31: 297-305.
Grootjen, C.J., Bouman, F. 1981. Development of ovule and
seed in Stanfieldiella imperforata (Commelinaceae). Acta.
Bot. Neerl. 30: 265-275.
Hamann, U. 1961. Merkmalsbestand und Verwandtschaft-
beziehungen der Farinosae. Ein Beitrag zum System der
Monokotyledonen. Willdenowia 2: 639-768.
Hamann, U. 1962. Weiteres tiber Merkmalsbestand und
Verwandtschaftsbeziehungen der Farinosae. Willdenowia
3: 169-207.
Handlos, W.L. 1975. The taxonomy of Tripogandra
(Commelinaceae). Rhodora 77: 213-333.
Harris, P.J., Hartley, R.D. 1980. See general references.
127
Hasskarl, J.K. 1870. Commelinaceae Indicae, Imprimis
Archipelagi Indica. Wien: M. Slazer.
Hegnauer, R. 1963, 1986. See general references.
Jones, K. 1974. Chromosome evolution by Robertsonian
translocation in Gibasis (Commelinaceae). Chromosoma
(Berl.) 45: 353-368.
Jones, K. 1990. Robertsonian change in allies of Zebrina
(Commelinaceae). Plant Syst. Evol. 172: 263-271.
Jones, K., Jopling, C. 1972. Chromosomes and the classifica-
tion of the Commelinaceae. Bot. J. Linn. Soc. 64: 129-162.
Jones, K., Kenton, A., Hunt, D.R. 1981. Contributions to the
cytotaxonomy of the Commelinaceae. Chromosome evolu-
tion in Tradescantia section Cymbispatha. Bot. J. Linn. Soc.
83: 157-188.
Kellogg, E.A., Linder, H.P. 1995. Phylogeny of Poales. In:
Rudall, P.J., Cribb, P.J., Cutler, D.P., Humphries, C.J. (eds.)
Monocotyledons: systematics and eolution. Royal Botanic
Gardens, Kew, pp. 511-542.
Lee, R.E. 1961. Pollen dimorphism in Tripogandra grandi-
flora. Baileya 9: 53-56.
Meisner, C.F. 1842. CCLXI Commelinaceae. In: Plantarum
Vascularium Genera, Vol. 1. Leipzig: Weidmann, pp. 406-
407.
Ornduff, R. 1974. Heterostyly in South African plants: a con-
spectus. J. S. Afr. Bot. 40: 169-187.
Owens, S.J. 1981. Self-incompatibility in the Commelinaceae.
Ann. Bot 47: 567-581.
Owens, S.J., Kimmins, P.M. 1981. Stigma morphology in
Commelinaceae. Ann. Bot. 47: 771-783.
Pichon, M. 1946. Surles commelinacees. Not. Syst. (Paris) 12:
217-242.
Poole, M.M., Hunt, D.R. 1980. Pollen morphology and the
taxonomy of the Commelinaceae: an exploratory survey.
American Commelinaceae: VIII. Kew Bull. 34: 639-660.
Rohweder, 0. 1956. Die Farinosae in der Vegetation von
El Salvador. Abh. Auslandsk. 61 Reihe C Naturwiss. 18: 1-
197.
Rohweder, 0. 1963. Anatomische und histogenetische
Untersuchungen an Laubsprossen und Bliiten der
Commelinaceen. Bot. Jahrb. 82: 1-99.
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por vibra<;ao em duas especies simpatricas de Dichori-
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de Botanica Goiana, 20-26 Janeiro 1991, Universidade
Federal de Goias, Sociedade Botanica do Brasil, Goiania,
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Humphries, C.J. (eds.) Monocotyledons: systematics nd
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128 Commelinaceae/Costaceae

auf andere Commelinaceen. Beitr. Bioi. Pflanz. 36: 325- Costaceae

389.
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154. Costaceae (Meisn.) Nakai, J. Jpn. Bot. 17: 203 (1941}.

Perennial, rhizomatous, non-aromatic herbs, ter-

restrial or more rarely epiphytic. Indumentum
of uniseriate, multicellular or unicellular hairs.
Stems terete, most often unbranched, leafy, some-
times spirally contorted; when branched, then
secondary branches breaking through the leaf
sheaths (Tapeinochilus). Leaves spirally arranged,
sheaths closed, tubular; ligule present; petiole
short; pulvinus absent; lamina narrowly to
broadly elliptic, rolled up from one side to the
other in bud. Inflorescence a strobilaceous spike,
terminal on the leafy shoot or on a separate,
short, leafless shoot, or flowers solitary, axillary
(Monocostus). Bracts imbricate, with a linear,
nectariferous callus below the tip, each subtending
1-2 flowers; bracteoles tubular or folded. Flowers
epigynous, perfect, zygomorphic; calyx tubular,
shortly 2-3-lobed; corolla 3-lobed, the lobes ba-
sally fused, imbricate in bud, unequal; stamen 1,
often petaloid, bearing 2 2-sporangiate thecae,
dehiscing introrsely by longitudinal slits, often
appendaged at the apex. Labellum petaloid,
staminodial, opposite to the stamen, as long as or
much longer than the corolla, ± 3-lobed, often
with crisped margin; basal part of stamen and
labellum united into a papillate tube. Gynoecium
3( -2)-carpellate; ovary inferior, 3( -2)-locular; pla-
centation axile; ovules numerous, anatropous,
crassinucellar; septal nectaries 2, immersed in the
apical part of the ovary; style filiform, commonly
enclosed between the thecae; stigma Wet, with or
without a dorsal appendage. Fruit a 3(-2)-locular
capsule crowned by the persistent calyx, dehiscing
loculicidally, rarely explosively, or indehiscent
and irregularly breaking when old. Seeds numer-
ous, aril white to yellow; operculum next to the
prominent radicle; embryo straight; endosperm
poorly developed; perisperm abundant, with
copious starch in simple grains.
A pantropical family comprising 4 genera and
about 100 species.

VEGETATIVE MoRPHOLOGY. The rhizome is

subterranean, sympodial, glabrous or sometimes
 Costaceae 129

hairy; mostly fleshy, thick, but in some species

thin and long-creeping. The roots are filiform and
fleshy, rarely terminating in fleshy tubers. There
is much variation in stem length from almost
acaulescent herbs to tall plants reaching up to 8 m.
With the exception of few species, the stems are
unbranched. Bulbils have been found in the leaf
axils and lower bracts of some species. The veg-
etative parts are often densely hairy. The leaf
sheaths are tubular and closed, not open as in the
Zingiberaceae. Leaf size strongly decreases to-
wards the base of the stem, where only bladeless
sheaths are developed; apically the leaves gradu-
ally change into bracts. The leaves are apparently
monostichously arranged, with unusual diver-
gence angles (Fig. 35). After much debate in the
literature of this "spirostichous monostichy", von
Veh (1931) demonstrated that developmentally
the phyllotaxy starts from distichy, the common
pattern in Zingiberales, and that the apparent
monostichy results from asymmetric growth of
the shoot apex.
The indumentum of the vegetative parts is com-
posed of uniseriate, multicellular hairs, base
never sunken (Tomlinson 1969), or unicellular
hairs (Maas 1972).

ANATOMY. The main features have been sum-

marised by Tomlinson (1969). The differences
from Zingiberaceae include the absence of oil
cells; the occurrence of uniseriate, multicellular
hairs; a different arrangement of the vascular
bundles in the leaves and the stem; and the ab-
sence of air canals. Rap hide sacs are absent. Silica
bodies are present as solitary, stellate or druselike
bodies in internal cells next to the vascular
bundles, but never in the epidermis. Tannin cells
seem to be lacking. Starch is stored copiously in
the rhizomes; the starch grains are often large and
irregular, not flattened.
Fig. 35A-H. Costaceae. Costus megalobractea. A Habit,
INFLORESCENCE AND FLORAL MORPHOLOGY. The
showing spirodistichous phyllotaxy. B Inflorescence. C-F C.
inflorescence terminates either a leafy or a lateral, lucanusianus. C Flower. D Longitudinal section of ovary. E
leafless shoot; both forms can be found on one Petaloid stamen, anther crested. F Bilamellate stigma, lateral
plant. In Monocostus, however, the flowers occur and front view. G, H C. cuspidatus. G Capsule, discharging
singly in the leaf axils. The inflorescence is a glo- seed packets. H Seeds sticking together by their arils.
(Takhtajan 1982)
bose or cylindric conelike spike (Fig. 35B), due to
the densely imbricate bracts that fully cover the
main axis. The bracts are persistent and either
coriaceous, chartaceous or herbaceous. Apically calyx are of utmost importance for identification.
the bracts may be provided with a more or The corolla varies from white through yellow to
less well-developed foliar appendage. In many orange or red and consists of a basal tube of about
species extrafloral nectaries localised on the bracts the same length as the calyx. The corolla lobes are
(nectarial "callus") support ants as protective imbricate and the dorsal one is usually larger than
agents (Schemske 1980). The shape and size of the the lateral ones and cucculate. The labellum is the
130 Costaceae

most conspicuous part of the flower, except in (1981). In the Old World, Costus is predominantly
Tapeinochilos, where it is small and inconspicu- pollinated by several genera of bees. Traplining
ous. Most authors, including Troll (1928), regard anthophorid bees were recorded pollinating two
the labellum as formed by all 5 staminodes, based species of Costus in Asia (Sarawak) by Kato (1996).
on the occurrence of 5 vascular strands, a view Also sunbird pollination of C. speciosus was re-
confirmed by developmental studies (Kirchoff ported from Indonesia.
1988). The colours of the labellum are the same
as those found in the corolla. The filament is flat, SEED ANATOMY. The seeds of Costus are 1-2mm
often ± petaloid, and somewhat succulent; the long and thus are the smallest of all Zingiberales.
anther is attached to the middle of the filament or They are provided with a white or yellow aril. An
below, or, in Tapeinocheilos, apically and with a operculum and a hypostase are well developed.
connective crest (Fig. 35E); the anther is dithecal. The mature seed coat of Costus is completely
The style is filiform and placed close to the fila- formed by the outer integument and the principal
ment, enclosed by the thecae. Two types of stig- mechanical layer is the endotesta, the cells of
mas occur (Maas 1972): (1) bilamellate stigmas, which are filled with silica. The tegmen is almost
consisting of 2 ciliate structures with a dorsal, hi- entirely degenerated. Costus agrees with nearly all
lobed appendage (Fig. 35F); this type is restricted other members of the Zingiberales in the posses-
to Costus subgen. Costus; (2) cupshaped stigmas sion of an aril, operculum and micropylar collar
with ciliate margin. (Grootjen and Bouman 1981). The perisperm rep-
resents the principal storage tissue and contains
EMBRYOLOGY. The anther wall comprises a secre- copious starch composed of simple concentric
tory tapetum of binucleate cells. The embryo sac starch grains. The endosperm is less abundant and
develops according to the Polygonum type; the free of starch. The embryo is straight and nearly as
synergids are hooked. Endosperm development is long as the perisperm (H. Huber, unpubl. observ.).
of the Helobial type. Embryogeny conforms to the The seedling structure is unique among Zingi-
Caryophyllad type (Davis 1966; Johri et al. 1992). berales; the cotyledonary sheath is subdivided
into a basal sheath and a bladelike distal part
PoLLEN MoRPHOLOGY. The pollen grains are that bears a basipetally directed, conelike out-
ellipsoidal and very large (80-210 f!m). They are growth (Tillich 1995).
distinctly aperturate: disulcate or mixed sulcate-
porate (Dimerocostus and Monocostus), panto- DISPERSAL. After anthesis, the tightly imbricate
porate (Costus) or spiraperturate (Tapeinochilus) bracts of several species of Costus spread out, ex-
(Punt 1968). The grains are resistant to acetolysis, posing the white capsule on the red inner side of
in contrast to the pollen of most other members of the bracts, thus probably attracting birds. In
Zingiberales, and their wall structure is quite un- some species of Costus subgen. Cadalvena and in
usual and has been interpreted as neotenic (Stone Monocostus uniflorus the fruiting pedicel elon-
et al. 1981). gates, exposing the fruit beyond the bract. The
ripe seeds hold together by their well-developed
KARYOLOGY. Apart from some old uncertain aril and are then explosively discharged as one
counts, 2n = 18 or 36 have been established in packet. Dispersal by water currents may also
all species studied of Costus; in Dimerocostus occur.
strobilaceus 2n = 28 were counted (see compila-
tion in Maas 1972). PHYTOCHEMISTRY. Nine species of Costus and
one of Tapeinochilos have been studied and found
PoLLINATION. Pollination by large euglossine to contain common flavonoids. Oil cells are lack-
bees is known from species with short and wide ing and no ethereal oils have been found. Rhi-
floral tubes, in which the large lip is flat and white zomes from several Costus species were found to
to yellow, often striped with red (Dimerocostus, contain steroid saponins and sapogenins. From
Costus subg. Costus sect. Costus, Maas 1972, 1977). the seeds benzoquinones and fatty oils have been
Species with narrow, thick-walled floral tubes isolated (Hegnauer 1986).
with yellow, orange or red lips are pollinated by
hummingbirds (Costus sect. Ornithophilus, Stiles DISTRIBUTION. The family is pantropic, as is the
1978). The sharing of a small pollinating largest genus, Costus. Monocostus and Dimero-
euglossine bee by two synchronously flowering costus are neotropical while Tapeinochilos is re-
species of Costus was reported by Schemske stricted to New Guinea and adjacent parts of
 Costaceae 131

Indonesia and Australia. Most species are compo- locular, longitudinally dehiscent. Seeds angular
nents of lowland tropical rainforests and rarely with short, lobed aril. n = ? About 15-20 spp., E
occur in mountain forests up to 2000 m. They are Malesia to NE Australia.
often found in clearings as they are somewhat
light-requiring. Several species are bound to
wet habitats such as riverbanks and freshwater 2. Costus L. Fig. 35
swamps. Costus L., Sp. Pl.: 2 (1753); Loesener, E.-P. ed. 2, 15a: 628-640
(1930); Maas, Fl. Neotr. 8: 1-140 (1972), 18: 162-218 (1977)
AFFINITIES. The Costaceae are closely related to (Neotrop. spp.).
the Zingiberaceae of which until recently they
have been regarded as a subfamily, or a tribe. Low to tall herbs. Ligule short or long, truncate or
Seed characters indicate a close affinity with bilobed. Inflorescence on a leafy shoot or on a
Zingiberaceae-Alpinieae. Mainly in view of the basal leafless shoot, conelike. Bracts densely
anatomical and morphological differences, the imbricate, sometimes apically appendaged.
family status of Costaceae seems justified and is Bracteoles tubular or folded. Calyx longer or
now accepted by most recent authors. shorter than the bracts. Corolla and labellum vari-
ously coloured. Labellum large and spreading, or
EcoNOMIC IMPORTANCE. The starch present in small and tubular. Stamens with the anther at-
the rhizome of Costus speciosus is used locally in tached to the middle of the filament, rarely to the
India, it is not aromatic. Fruits of Dimerocostus base. Basal part of stamen and labellum joined
are said to be edible. Some species are commonly into a papillate tube. Ovary 3-locular, ovules 2-
grown in greenhouses. On a whole the family is of seriate. Capsule white, ellipsoid or globose, often
no or little economic importance. trigonous. Seeds glossy, black or brown; arillarge,
lacerate, rarely short and cushionlike. 2n = 18, 36.
KEY TO THE GENERA About 90 spp., pantropical, most in tropical
America.
1. Labellum conspicuous; filament petaloid; connective crest
petaloid, distinctly exceeding the anther; bracts coriaceous
to herbaceous; leafy shoot not branched 2 3. Dimerocostus Kuntze
- Labellum inconspicuous, small; filament not petaloid; con-
nective crest small; bracts coriaceous to woody; leafy shoots Dimerocostus Kuntze, Rev. Gen. 2:687 (1891); Maas, Fl. Neotr.
often branched. Malesia and NE Australia 1. Tapeinochilus 8: 20-27 (1972).
2. Flowers solitary in the upper leaf axils. Peru 4. Monocostus
- Flowers in a conelike inflorescence 3
3. Ovary 2-locular; bracteole tubular. S America
Very tall herbs. Leaves large, usually long
3. Dimerocostus acuminate; ligule very short. Inflorescence often
- Ovary 3-locular; bracteole tubular or folded. Pan tropical strongly elongate, terminal on a leafy stem. Bracts
2. Costus coriaceous, often sheathing, sometimes with folia-
ceous appendages. Bracteole tubular, 2-keeled or
2-winged. Calyx exceeding the bracts. Corolla and
Genera of Costaceae labellum white or yellow. Labellum very large,
spreading. Stamen with anther attached to the
1. Tapeinochilos Miq. middle of the filament. Stigma cup-shaped. Ovary
Tapeinochilos Miq., Ann. Mus. Lugd.-Bat. 4: 101 (1868),
2-locular; ovules 2-seriate. Capsule 2-locular, tar-
Loesener in E.-P., Natiirl. Pflanzenfamil. ed. 2, 15a: 638-640 dily dehiscent. Seeds with small, cushionlike aril.
(1930). 2n = 28. Two spp., central and western S America.

Tall herbs, often branched in upper part. Inflores-

cence cone-like, terminal on a short, leafless
shoot. Bracts reddish, coriaceous to almost
woody, acuminate. Flowers yellowish or brown-
ish, hardly exceeding the bracts. Calyx coriaceous
to almost woody. Corolla tube with short lobes.
Labellum flanked by lateral teeth. Stamen with
broad filament, anther terminal with an apical
connective crest. Ovary 2( -3)-locular. Style at base
enlarged; stigma flattened triangular. Capsule 2-
4. Monocostus K. Schum.
Monocostus K. Schum., Pflanzenreich IV 46:427 (1904); Maas,
Fl. Neotr. 8: 17-20 (1972).
Low herbs. Ligule short. Flowers solitary in the
upper leaf axils. Bracteole tubular. Calyx longer
than the bracts. Corolla yellow, lobes shorter than
tube. Labellum large and spreading. Stamen with
the anther attached at the middle of filament.
132 Costaceae
Stigma cup-shaped. Ovary 2-locular; ovules 1-
seriate. Capsule strongly elongate, longitudinally
flattened, 2-locular, longitudinally dehiscent.
Seeds black, aril small. 2n = 28. Only one sp., M.
uniflorus (Poepp. ex Petersen) Maas, endemic to E
Peru.
Selected Bibliography
Davis, G.L. 1966. See general references.
Grootjen, C.J., Bouman, F. 1981. Development of the ovule and
seed in Costus cuspidatus (N. E. Br.) Maas (Zingiberaceae),
with special reference to the formation of the operculum.
Bot. J. Linn. Soc. 83: 27-39.
Hegnauer, R. 1986. See general references.
Holttum, R.E. 1950. The Zingiberaceae of the Malay Peninsula.
Gard. Bull. (Singapore) 13: 240-245.
Johri, B.M. et a!. 1992. See general references.
Kato, M. 1996. Plant-pollinator interactions in the understory
of a lowland mixed Dipterocarp forest in Sarawak. Am. J.
Bot. 83: 732-7 43.
Kirchoff, B.K. 1988. Inflorescence and flower development in
Costus scaber. Can. J. Bot. 66: 339-345.
Kress, W.J. 1990. The phylogeny and classification of the
Zingiberales. Ann. Mo. Bot. Gard. 77: 698-721.
Loesener, L.E.T. 1930. Zingiberaceae. Unterfam. II. Costoideae
In: Engler, A., Prantl, K., Pflanzenfamilien, 2nd edn., 15a.
Leipzig: W. Engelmann, pp. 628-640.
Maas, P.J.M. 1972. Flora Neotropica, Monogr. 8. Costoideae
(Zingiberaceae). New York: Hafner.
Maas, P.J.M. 1977. Flora Neotropica, Mongr. 18. Renealimia
(Zingiberaceae-Zingiberoideae), Costoideae (Additions).
New York: The New York Botanical Garden.
Pijl, L. van der 1954. Xylocopa and flowers in the tropics I-III.
Proc. K. Ned. Akad. Wet. C 57: 413-423, 541-562.
Punt, W. 1968. Pollen morphology of the American species of
the subfamily Costoideae (Zingiberaceae). Rev. Palaeobot.
Palynol. 7: 31-43.
Schemske, D.W. 1980. The evolutionary significance of
extrafloral nectar production by Costus woodsonii
(Zingiberaceae): an experimental analysis of ant protection.
J. Ecol. 68: 959-967.
Schemske, D.W. 1981. Floral convergence and pollinator shar-
ing in two bee-pollinated, tropical herbs. Ecology 62: 946-
954.
Schumann, K. 1904. Subfamilia II. Costoideae K. Schum. In:
Engler, A. (ed.) Das Pflanzenreich, IV.46. Leipzig, W. Engel-
mann, pp. 377-437.
Stiles, F.G. 1978. Temporal organization of flowering among
the hummingbird food plants of a tropical wet forest.
Biotropica 10: 194-210.
Stone, D.E., Sellers, S.C., Kress, J.W. 1981. Ontogenetic
and evolutionary implications of a neotenous exine in
Tapeinochilos (Zingiberales: Costaceae) pollen. Am. J. Bot.
68: 49-63.
Takhtajan, A.L. 1982. See general references.
Tillich, H.-J. 1995. Seedlings and systematics of the monocoty-
ledons. In: Rudall, P.J., Cribb, P.J., Cutler, D.F., Humphries,
C.J. (eds.) Monocotyledons: systematics and evolution, Vol.
1. London: Royal Botanic Gardens Kew, pp. 303-352.
Tomlinson, P.B. 1969. See general references.
Troll, W. 1928. Organisation und Gestalt im Bereich der Bliite.
Berlin, Heidelberg, New York: Springer.
Veh, R. von 1931. Untersuchungen und Betrachtungen zum
Blattstelungsproblem. Flora 125: 83-154.
 Cymodoceaceae 133

Cymodoceaceae inner having thin, non-lignified walls (Amphi-

J. Kuo and A.J. McCoMB
Cymodoceaceae N. Taylor, N. Am. Fl. 17:31 (1909), nom. cons.
bolis, Thalassodendron). The cortex parenchyma
may store starch (e.g. Halodule). An exodermis
may be present, and the epidermis may become
lignified (e.g. Amphibolis and Thalassodendron).
In addition to the more or less horizontal rhi-
zomes, there are erect stems bearing leaves; these
stems are long in Amphibolis and Thalasso-
dendron. Their anatomy is similar to that of the

Perennial marine herbs with creeping, monopodi-

ally or sympodially branched rhizomes. Roots of-
ten branched, with few to many root hairs. Leaves
alternate, distichously arranged, with distinct
blade, sheathing base and a ligule between sheath
and blade; sheath after being shed leaving an open
or closed circular scar on stem; blade linear, flat
or terete; tannin cells numerous; stomata absent.
Squamules present in each leaf axil. Inflorescence
cymose or flowers solitary. Flowers dioecious,
naked, enclosed by leaflike bracts; male flowers
subsessile or pedicellate, with 2 tetrasporangiate
stamens; female flowers of 2 free ovaries, each
with a long stylulus which is unbranched or
divided into 2-3 filiform stigmatic stylodia;
carpels 1-ovulate; usually only 1 carpel develop-
ing into fruit. Fruit indehiscent; seeds germinat-
ing on parent plant in Amphibolis and
Thalassodendron.
A family of 5 genera and 16 species of seagrasses,
largely in tropical and subtropical waters of
the Indo-West Pacific and Caribbean, but extend-
ing into warm-temperate Australia and the
Mediterranean.

VEGETATIVE MORPHOLOGY AND ANATOMY. The

rhizomes are cylindrical, herbaceous and mo-
nopodially branched in Cymodocea (Fig. 36A),
Halodule and Syringodium, but woody and
sympodially branched in Amphibolis and
Thalassodendron (Fig. 36D) (Tomlinson 1974).
Each rhizome internode has a central stele with 1
or more large, central protoxylem elements. The
stele is surrounded by a conspicuous endodermis
which may become suberised (Halodule) and may
have several layers of thickened, lignified cells
(Amphibolis, Thalassodendron). The numbers of
cortical vascular bundles differ between genera,
and some bundles may be associated with fibrous
strands (e.g. Cymodocea; Kuo 1983a). Cortex pa-
renchyma cells are thin-walled and the cortex
Fig. 36A-D. Cymodoceaceae. A-C Cymodocea nodosa. A Pis-
contains large lacunae (e.g. Syringodium). Alter- tillate plant with young fruits. B Staminate plant. C Anther. D
natively, the cortex is divided into 2 zones, cells of Thalassodendron cilia tum, plant with pistillate inflorescence.
the outer having thick, lignified walls, those of the (Takhtajan 1982)
134 Cymodoceaceae
rhizome, except that the numbers of fibre strands
may be reduced (e.g. Cymodocea).
From one to three branched or unbranched
roots are produced at every node (Halodule,
Syringodium, Cymodocea and Amphibolis) or
at the node where rhizome branching takes
place (Thalassodendron). Roots may become wiry
(Amphibolis) or woody (Thalassodendron) when
mature. Root hairs may be either extensive
(Syringodium, Cymodocea, Halodule) or infre-
quent (Amphibolis, Thalassodendron) (Kuo 1983a,
1993b).
Bacterial colonies are usually present in the
rhizosphere and may be involved in nitrogen fixa-
tion and nutrient uptake by the roots (Kuo 1993b).
Root epidermal cells are thin-walled and lignified,
with the exception of Cymodocea. The exodermis
may become suberised and the cortex may have a
system of large gas lacunae (Syringodium), or de-
velop distinct outer and inner zones (Amphibolis,
Thalassodendron). A central stele is surrounded
by an endodermis with a distinct Casparian strip
and suberised walls (Kuo 1993b; Barnabas 1994).
The xylem is normally represented by thin-walled,
unlignified cells with small lumens; sieve tube
number and position vary between genera (Kuo
1983a).
There are 3 kinds of leaf: (1) ligulate foliage
leaves with open sheaths; (2) prophylls or rhizome
scales; (3) bracts, leaves with reduced blades asso-
ciated with flowers (Tomlinson 1982). The foliage
leaves are distichously arranged on stems with
short internodes (Syringodium, Halodule,
Cymodocea), or on tall erect stems (Amphibolis,
Thalassodendron). The leaf blade may be terete
(Syringodium), linear and narrow (Cymodocea,
Halodule), or short and broad (Amphibolis,
Thalassodendron). The apex of the blade is
rounded, emarginate or dentate. There are three
(e.g. Halodule) to more than 20 (Thalassodendron,
Amphibolis) longitudinal, parallel vascular bun-
dles, connected periodically by transverse veins
(Tomlinson 1982; Kuo 1983a).
The cells of the leaf epidermis are either uni-
formly chlorenchymatous and contain tannin
(Amphibolis, Thalassodendron), or possess
large, tannin-containing cells, the other cells
being chlorenchymatous (Syringodium, Halodule,
Cymodocea). The cuticle of the leaf is porous
(Amphibolis) or electron-transparent (all other
genera). Mesophyll cells are thin-walled, and
contain some chloroplasts. Gas lacunae are
well developed in linear, narrow or terete leaves
( Cymodocea, Halodule, Syringodium ), and
present in the broad leaves of Amphibolis and
Thalassodendron (Kuo 1983a, 1993a; Larkum et al.
1989b).
Within longitudinal veins the structure of the
bundle-sheath cells varies with genera, the cells
with thick walls and possessing suberised lamellae
in Amphibolis and Thalassodendron (Ducker
et al. 1977; Barnabas 1983), with thin walls and
suberised lamellae in Syringodium and Halodule
or without obvious structural specialisation in
Cymodocea (Kuo 1983a, 1993a; Barnabas 1994).
Normal, thin-walled sieve tubes occur in all genera
except Halodule, which has nacreous-walled sieve
tubes. All have monocotyledonous plastids with
protein crystalloids (Kuo 1983b, 1993a). The
xylem is represented by 2 to 4 small lumens,
with unlignified, thin-walled tracheids. Vascular
parenchyma cells do not have wall ingrowths.
Fibrous strands are only associated with the hypo-
dermis adjacent to the longitudinal vascular
bundles in Cymodocea. Apoplastic tracers move
freely from the medium into the epidermis and
mesophyll tissues but the movement is restricted
by suberised lamella in Thalassodondron leaves
(Barnabas 1988, 1989).
Leaf sheaths are clearly differentiated from the
blades, and enclose young, developing leaves.
Their anatomy is similar to that of the leaf blades,
except that they lack chloroplasts and have more
pronounced gas lacunae. There are 2 to several
squamules at each leaf axil.
INFLORESCENCE STRUCTURE. The dioecious flow-
ers are usually solitary, and terminal on erect
shoots (Halodule, Cymodocea, Fig. 36A) or
their branches (Amphibolis, Thalassodendron),
but they form a distinct cymose inflorescence in
Syringodium (Tomlinson and Posluszny 1978).
In Amphibolis (McConchie et al. 1982a) and
Thalassodendron (Isaac 1969; Kay 1971; Kuo and
Kirkman 1987) each inflorescence consists of 2
flowers and is surrounded by a sheathlike bract.
The 2 morphologically identical male flowers have
different developmental stages (Kuo and Kirkman
1987). In general, the flowers of this family have
been considered as lacking perianth parts, but the
bract subtending the anthers has been referred to
as a perianth part for Amphibolis (McConchie et al.
1982a), and a perianth ridge for Syringodium
(Tomlinson and Posluszny 1978). Details of
floral development in S. filiforme are given by
Tomlinson and Posluszny (1978).
FLORAL STRUCTURE. Interpretation of floral
morphology and structure in the Cymodoceaceae
presents particular difficulties (Kuo and McComb
 Cymodoceaceae
1989). Some workers regard the flower as a single
unit supported by special bracts, while Markgraf
(1936) and others considered each anther or pistil
to be a separate flower forming a partial inflores-
cence. Each male flower is either subsessile or
stalked, the stalk elongating at anthesis. There are
2 anthers, which are laterally fused and bear 2 or
3 apical appendages on each lobe (Amphibolis,
Thalassodendron) (Isaac 1969; Kay 1971;
McConchie et al. 1982a; Kuo and Kirkman 1987).
The anthers of Halodule are unequal in height.
Anthers are longitudinally dehiscent.
The female flower is sessile or with a short
pedicel, and has 2 free carpels, each with either a
simple long stylulus (Halodule) or a stylulus with 2
stout (Syringodium) or 2 to 3 slender (Amphibolis,
Thalassodendron) stigmatic stylodia. The stigmas
are exposed (in intertidal species) during anthe-
sis, but eventually break off after pollination
(McConchie and Knox 1989). The stigma in
Thalassodendron bears irregular ridges and hol-
lows and does not have a pellicle (Pettitt 1976), but
that in Amphibolis is non-papillate (McConchie et
al. 1982a). Both carpels can develop into fruits, but
there is frequent abortion of one or both.
FLORAL ANATOMY. The vascularisation of there-
productive system is similar in all members of the
family. The main vein is derived from the central
axial bundle and extends into the base of the
flower. In Amphibolis (McConchie et al. 1982a),
two additional cortical bundles terminate in the
pedicel, while in Syringodium (Tomlinson and
Posluszny 1978), 4-6 cortical bundles become lat-
eral bundles of the bracts. The main vein may
divide at the base of the flower, one branch be-
coming the main bundle of the bract; the other
divides again, and supplies anther or pistil. In the
anther, each bundle may or may not lead into
the terminal appendage of the anther, while in the
pistil, it divides again, giving rise to a dorsal group
of veins that encircle the integuments, so supply-
ing the ovary. A single, central bundle forms the
stylar trace that leads to the adaxial wall of the
ovary, and terminates in the base of the style or
passes into the style and central stigmatic branch,
where it terminates.
EMBRYOLOGY. The ovule has been described as
atropous, pendulous; the embryo sac has not been
investigated. Endosperm development is Nuclear.
In Thalassodendron, the innermost bracts con-
tinue to develop after anthesis or fertilisation, and
eventually enclose the carpels to form a false fruit
(Isaac 1969; Kay 1971; Kuo and Kirkman 1987).
135
A feature known as the grappling apparatus in
Amphibolis is initiated from the ovary wall late
in development of the flower, but its initials are
present before fertilisation. There are 4 fleshy, her-
baceous pericarp lobes which become hard and
have fine bristles united at the base (Black 1913;
McConchie et al. 1982a). It lacks vascular tissue.
Tepper (1882) regards the grappling apparatus
as tepals, while Tomlinson and Posluszny (1978)
refer them to sepals.
POLLEN MORPHOLOGY. Pollen grains are filiform
and trinucleate. The wall lacks an exine but the
surface usually has a longitudinal ridge pattern
with pits (McConchie et al. 1982b; McConchie and
Knox 1989). Pollen grains are tightly coiled within
the anther, irregularly coiled in Halodule and
Thalassodendron and coiled like a spring in
Amphibolis (Ducker et al. 1978). Pollen grains have
fine tips which are usually unforked, but may
fork as in Amphibolis (McConchie and Knox
1989). The pollen grains are 1000[lm long in
Halodule and 3000 to 5000 [lm in Amphibolis
(Pettitt and Jermy 1975; Ducker et al. 1978; Pettitt
et al. 1984). The development of pollen in
Amphibolis and Thalassodendron has been de-
scribed in detail (Ducker et al. 1978; Pettitt et al.
1980b, 1984; Pettitt 1981). The pollen morphology
of Cymodocea and Syringodium has not been
described.
KARYOLOGY. Chromosome counts appear diffi-
cult to obtain from these seagrasses; all available
numbers are based on root tip materials, and
counts are variable. For Cymodocea species 2n =
14, 28 and 30; Syringodium has 2n = 20; Halodule
2n = 16 and 44, and den Hartog et al. (1979)
suggested that populations of H. wrightii in
Curayao are tetroploid. Chromosome counts of
Amphibolis and Thalassodendron have not been
reported (den Hartog et al. 1979, 1987; Uchiyama
1993).
POLLINATION. Pollination is underwater (Ducker
and Knox 1976; Pettitt et al. 1980a). At the time of
anthesis the female flower remains hidden under
the bracts, except for the stigmas, the pollen ad-
heres tenaciously to the stigmas, but pollen germi-
nation proceeds near but never at the point of
pollen contact (Yamashita 1976).
Pollination biology in seagrasses has been ex-
tensively studied (see Pettitt et al. 1980b, 1983;
Pettitt 1984; Cox 1988, 1993; Cox and Knox 1989;
McConchie and Knox 1989; Cox et al. 1991). Cox
and his coworkers discussed 2- and 3-dimensional
136 Cymodoceaceae

pollination systems in seagrasses (Cox and Knox Much of the expansion of meadows occurs
1989), where pollen grains having a hydrophobic through the growth of rhizomes. Vegetative
surface are not transported through the water sin- propagules can also develop on the erect stems of
gly, but in rafts of "search vehicles" to increase Amphibolis (Kuo et al. 1987).
probability of encountering a stigma.
Entry of the pollen tube into the embryo sac PHYTOCHEMISTRY. Sulphated phenolic com-
has rarely been observed in any seagrass. In pounds are present in all genera in the family and
Amphibolis the pollen tube grows through and may play a role in the adaptation of the seagrasses
between the cells of the nucellus before entering a to their marine habitats (Zapata and McMillan
synergid, laterally from below. The pollen tube in 1979; McMillan et al. 1981). Drew (1983) found
Cymodocea enters through a micropyle and grows that the distribution of sugars and cyclitols corre-
directly towards a synergid (Bornet 1864). lates with generic limits.
Isoenzyme patterns in Thalassodendron and
FRUIT AND SEED. After loss of the fleshy Amphibolis are more similar to each other than to
exocarp, Cymodocea, Halodule and Syringodium other genera (Halodule, Cymodocea and Syringo-
produce indehiscent 1-seeded fruits with a stony dium) in the family (McMillan 1982). The isoen-
endocarp of various colours. Seeds of these zyme patterns in the two sympatric species of
seagrasses have seed coats whose structures vary Cymodocea (C. serrulata and C. rotundata) differ
with genera (McMillan and Bragg 1987). The em- markedly but those of Amphibolis antarctica
bryo has an enlarged hypocotyl and stores nutri- and A. grif.fithii are nearly identical (McMillan
ent reserves, mainly starch (Kuo 1983a). The seeds 1980a, 1981, 1982; McMillan et al. 1981). Further-
of Amphibolis and Thalassodendron lack seed more, Halodule and Syringodium in the Gulf of
coats and germinate while attached to the parent Mexico-Caribbean have different isoenzymes
plants, and numerous transfer cells are present at from those of species in the same genera in the
the interface between seed and parent tissue (Kuo Indo-Pacific, but species populations of various
and Kirkman 1990). origins within each ocean system show little
isoenzyme variation, suggesting that genetic
GERMINATION AND EsTABLISHMENT. Seeds of divergence is a consequence of spatial isolation
the temperate Cymodocea nodosa have a distinct (McMillan 1980a).
dormancy period of 7-8 months, and germination DNA techniques did not disclose genetic
commences when the temperature exceeds 18°C; variability in a dioecious species of Amphibolis
the rate rises at higher temperatures (Caye and antarctica (Waycott et al. submitted).
Meinzies 1985). In the tropical species C.
rotundata, germination can occur throughout the CLASSIFICATION. The genera fall into two groups,
year. Seeds of Halodule are dormant possibly one containing Cymodocea, Syringodium and
for many years, while those of Syringodium have Halodule, the other Amphibolis and Thalas-
a short dormancy. Seeds of Halodule and Syringo- sodendron (Tomlinson 1982). The latter group
dium continue are viable for more than 3 years is distinguished from the former by the pos-
after seeds have been released from parent plants. session of long erect stems and viviparous
For Halodule, dormancy involves both seed coat reproduction.
and embryo, but for Syringodium it probably in-
volves only the seed coat (McMillan 1983). EVOLUTION AND AFFINITIES. den Hartog (1970)
Seeds of Thalassodendron and Amphibolis ger- proposed that seagrasses might have evolved from
minate on the parent plant immediately after woody, salt-tolerant maritime plants by direct
fertilisation, and develop into seedlings with leafy passage from land into salt water. He further con-
shoots but without roots; they remain attached sidered that Amphibolis and Thalassodendron,
for about 7 to 12 months (McConchie et al. 1982a; which have erect, woody stems and viviparous
Kuo and Kirkman 1987). When the seedlings be- reproduction, are the most primitive of the
come free, the grappling apparatus of Amphibolis recent seagrass genera, a topic that merits further
serves to anchor the seedling to a substrate and consideration.
roots develop (Tepper 1882; Black 1913; Ostenfeld On basis of the analysis of phylogenetic relation-
1916). The seedling of Thalassodendron, detached ships and sexual conditions, Waycott and Les
from the parent plant, is eventually released from (1966) disagreed with the conclusion of Cox and
the enveloping bract of the false fruit, and the root Humphries (1993) that the three seagrass families
develops (Issac 1969; Kuo and Kirkman 1987). Zosteraceae, Posidoniaceae and Cymodoceaceae
 Cymodoceaceae
share a common ancestor with noodle-shaped
pollen. The molecular studies of Les et al. (1997)
indicate a close relationship between Cymo-
doceaceae, Posidoniaceae, and Ruppiaceae.
GEOGRAPHIC DISTRIBUTION. Members of the
family are largely tropical and subtropical, apart
from Amphibolis and Thalassodendron pachy-
rhizum from warm-temperate, southern and SW
Australia respectively, and Cymodocea nodosa
from the Mediterranean. There is a rich centre for
the family in the Indo-West Pacific including Aus-
tralia (all five genera), and a secondary centre in
the Caribbean (two genera). Most species have a
wide distribution, but T. pachyrhizum and C.
angustata are restricted to certain areas of W
Australia.
The family is wholly marine and generally sub-
tidal, in water from 0.5 to 40 m deep; some species
(including C. serrulata and T. ciliatum) extend
into the intertidal. The plants often form extensive
meadows of single or mixed species on sandy or
muddy substrates. T. pachyrhizum usually grows
on rocks, and S. isoetifolium is normally found
among other species, and does not form meadows.
Vast meadows, mainly of Amphibolis, occur
at Shark Bay, W Australia, where stabilisation of
sediments by seagrasses, and production of cal-
careous skeletal material from epiphytes and in-
fauna has produced striking changes in submarine
geology, and the effective isolation from the open
ocean of large areas of water (Walker 1985). The
plants are of great importance to nutrient cycling
and detrital food webs in nearshore environments.
There is also direct grazing, notably by turtles and
dugongs (Phillips and McRoy 1981; Larkum et al.
1989).
PARASITES. In Halodule (H. uninervis, H.
beaudettei, H. wrightii and H. piniforlia) the
parasitic fungus Plasmodiophora diplantherae
(Plasmodiophoraceae) causes marked swelling of
internodes, often giving the stem the appearance 1. Syringodium Kiitz.
of a string of beads. Syringodium Kiitz. in Hohen., "Meeralgen" 9, No. 426 (1860);
PALAEOBOTANY. Thalassocharis has been de-
scribed from the Upper Cretaceous of Limburg
(The Netherlands) (Voigt and Domke 1955).
Thalassodendron auricula-leporis has been re-
ported from the late Middle Eocene in Florida,
USA (Lumbert et al. 1984). The occurrence of this
genus as a fossil in America, where it does not now
live, is in line with the concept of den Hartog
(1970) that the present-day area of this genus is
relictual. The genus Cymodocea is represented in
137
the fossil record by various fossil and possibly
extant species.
EcoNOMIC IMPORTANCE. The significance of the
group is substantial, especially in areas where
large meadows occur. It lies in the stabilisation of
sediments, in nutrient cycling, in supporting graz-
ing dugong and turtles, and, probably most sig-
nificant of all, in the support of food webs through
the production of organic detritus. These food
webs presumably underlay the important fisheries
of regions where seagrasses dominate submarine
plant biomass.
KEY TO THE GENERA
1. Rhizome herbaceous with monopodia! branching; root
fleshy; erect stems short, bearing leaves; leaf blade shed,
leaving persistent leaf sheath; anthers stalked; mainly tropi-
cal and subtropical 2
- Rhizome stiff, woody, with sympodial branching; roots stiff
or wiry; erect stems elongated, bearing terminal leaf clus-
ters; leaf blade and leaf sheath shed together; anthers
sessile; mainly temperate, subtropical 4
2. Leaf blade distinct terete; many flowers in conspicuous
cymose inflorescences 1. Syringodium
- Leaf blade linear, fiat; inflorescence 1-fiowered or 2 flowers
resembling a single flower 3
3. Leaf blade narrow ( <3 mm wide) with 3longitudinal vascu-
lar bundles; roots unbranched; anthers not attached at the
same height; stylulus unbranched 2. Halodule
- Leaf blade broad (usually >3mm wide) with 7-17longitu-
dinal bundles; roots branched; anthers attached at the same
height; stylulus with 2 stylodia 3. Cymodocea
4. Stems and roots arising at intervals of 4 rhizome inter-
nodes; stems unbranched or little branched; leaf blade mar-
gin and apex coarsely denticulate; flowers surrounded by 4
bracts; each anther crowned with 1 appendage; stylulus with
2 stylodia; fruit lacking comblike outgrowths but with an
enlarged inner bract 5. Thalassodendron
- Stems arising at irregular intervals, branching roots at every
node; stems branched; leaf blade margin smooth, leaf apex
bidentate; flowers not enclosed by bracts; each anther
crowned with 2-3 appendages; stylulus with 3 stylodia; fruit
with 4-lobed, comblike outgrowths 4. Amphibolis
den Hartog, Verh. K. Ned. Akad. Wet. II, 59 (1): 176-186
(1970).
Rhizome monopodially branched, herbaceous,
creeping, with 1 or more branched roots and
short, erect, unbranched shoots at each node.
Leaves 2 or 3 on each erect shoot; blade terete,
with 1-several peripheral vascular bundles and
several large air lacunae. Leaf scars open, circular;
leaf sheath open; 2 squamules at each node. Dioe-
cious; inflorescences with bracts terminating erect
138 Cymodoceaceae

shoots; branching racemose below, cymose above; 4. Amphibolis Agardh

male flowers with anthers at the same level, dor-
sally fused in their lower parts; female flowers
sessile; ovaries extending into a short stylulus and
2 stout stylodia. Two spp., one in the Indo-West
Pacific, the other in the Caribbean.
2. Halodule Endl.
Halodule End!., Gen. Pl. suppl. 1: 1368 (1841); den Hartog,
Verh. K. Ned. Akad. Wet. II, 59 (1): 148-160 (1970), rev.
Rhizome monopodially branched, herbaceous,
with 2 cortical vascular bundles in each internode,
bearing one or more roots and a short erect shoot
at each node. Leaves 1-4 on each erect shoot;
blade linear with dentate, rounded or emarginate
apex; 3 longitudinal vascular bundles; leaf sheath
auriculate, ligulate, shed after leaf blade leaving a
circular scar on the stem; squamules 2. Dioecious;
inflorescences solitary, terminal on a short erect
shoot, subtended by the terminal leaf. Male flower
with anthers attached at different levels. Female
flower subsessile; with 2 free carpels each with a
long stylulus. Fruit with stony pericarp, ovoid or
somewhat compressed, with a short subapical
beak. Six very similar spp., widely distributed
along all tropical sea coasts.
Amphibolis Agardh, Species Algarum 1: 474 (1823); den
Hartog, Verh. K. Ned. Akad. Wet. II, 59 (1): 198-212 (1970).
Rhizomes sympodially branched, woody, creep-
ing, bearing 1-2-branched, wiry roots at each
node and erect stems irregularly at intervals of few
internodes. Erect stems becoming much branched
sympodially, bearing terminal tufts of leaves.
Leaf sheaths compressed, ligulate and auriculate,
amplexicaul, leaving closed circular scars when
shed; blade linear, flattened, with 8-21 longitu-
dinal veins; squamules present in the axil of each
leaf. Dioecious; inflorescence solitary, terminal on
short lateral leafy branches. Male flower shortly
stalked, subtended by a bract; anthers joined on a
common stalk. Female flower subsessile; the
styluli dividing into 4loriform stylodia. Fruit with
1 ripe ovary subtended by 4 spreading, toothed
lobes as extensions of a pericarp acting as a grap-
pling apparatus; seed germinating immediately on
parent plant and remaining there for several
months. Two sympatric spp., restricted to tem-
perate Australia.
5. Thalassodendron den Hartog Fig. 36D
Thalassodendron den Hartog, Verh. K. Ned. Akad. Wet. II, 59
(1): 186 (1970).

3. Cymodocea Konig Fig. 36A-C

Perennials with sympodially branched, robust
creeping rhizomes. Rhizomes bearing 1-2 erect,
Cymodocea Konig, Ann. Bot. (Konig & Sims, Lond.) 2: 96
strong, woody, unbranched (or little branched)
(1805); den Hartog, Verh. K. Ned. Akad. Wet. II, 59 (1): 160-
176 (1970), rev. stems at every 4th internode; roots arising at the
nodes preceding the stem-bearing nodes. Leaves
Rhizome monopodially branched, herbaceous, in clusters at stem tips. Leaf sheath compressed,
the nodes each bearing 1-5 branched roots and a auriculate, ligulate, amplexicaul; blade linear, fiat
short erect shoot bearing 2-7 leaves. Leaf blade apex almost truncate, denticulate, with many
linear, with 7-17 parallel veins. Leaf sheath com- parallel veins; squamules present in each leaf
pressed biauriculate, ligulate, shed after the leaf axil. Dioecious; inflorescence solitary, terminal on
blade, leaving an open or a closed circular scar on short lateral shoots, enclosed by 4 leafy bracts.
the stem; squamules 10 or more in 2 groups at Anthers with a terminal appendage. Fruit consist-
each node. Dioecious; inflorescences terminal ing of 1 fertilised ovary and a fleshy innermost
with erect lateral shoots, naked. Male flowers bract; seedling often remaining attached to the
with anthers fused at the same level, each with a parent plant after germination, then free-floating
short apical process. Female flower sessile or for a time. Two spp., one widely but discontinu-
shortly stalked, each ovary with a short stylulus ously distributed in the tropical Indo-West Pa-
extending into 2 long, slender stylodia. Fruit with cific, the other confined to temperate W Australia.
stony pericarp, smooth or somewhat warty with a
beak, semicircular to semiovate or elliptical in
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 Cyperaceae 141

Cyperaceae uppermost glumes), or only 1 to few upper

(larger) glumes fertile, and then flower often en-
P. GOETGHEBEUR closed by the wings of the next glume. Flowers
bisexual or (functionally) unisexual, and then
spikelets bisexual or unisexual, rarely dioeci-
ous. Hypogynous scales 0-(1-)3-6( -many), often
bristle-like and retrorsely scabrid, more rarely
laminar, sometimes with highly elaborated forms,
shorter or longer than the fruit, usually deciduous
Cyperaceae Juss., Gen. Pl.: 26 (1789), nom. cons. with the fruit, rarely persistent on the rachilla.
Stamens usually 3 (1 medio-anterior, 2 lateral),
Small to tall perennial or annual herbs, rarely but not rarely reduced to 1 or 2, or 4-6 and even
dwarf shrubs, shrubs, or lianas, terrestrial or more numerouso; filaments sometimes strongly
helophytic, rarely aquatic, epiphytic, or epilithic; elongating after anthesis, rarely connate; anther
the perennial species tufted, rhizomatous, stolo- basifixed, introrse, elliptic or oblong to linear,
niferous, bulbiferous, more rarely with a caudex. base often ± sagittate, connective often developed
Culms central, more rarely lateral, scapose or 1- into a ± pronounced crest. Pistil superior, usually
few-noded, more rarely many-noded, base some- 3-carpellate, unilocular, ± trigonous, with 1
times bulbously thickened, (sub )trigonous to medio-anterior rib and 2 lateral ribs; style long,
(sub)terete in cross-section, more rarely com- more rarely short, with 3 stigmatic branches, these
pressed, winged or 4-5-6-angular, rarely hollow sometimes with particular papillae; pistil some-
or septate, smooth or scabrid, rarely viscid, papil- times 2-carpellate, and then either dorsiventrally
lose, or hairy. Leaves all (sub)basal or 1-few or more rarely laterally compressed, rarely with a
cauline, rarely all cauline, (spiro)tristichous, more more elevated number of stigmatic branches. Style
rarely (spiro )distichous, lower leaves often (rarely base distinct from the ovary or not, often con-
all leaves) reduced to a sheath; sheaths usually spicuously thickened and persistent, glabrous,
closed, very rarely open, sometimes succulent scabrid, or variously hairy. Ovule single, basal,
or spongy; ligule present or absent, contraligule anatropous. Fruit an achene s.l. (nutlet), rarely
rarely developed; blade linear, rarely oblong, en- drupaceous, usually ovate, elliptic, or obovate in
siform or terete, rarely pseudopetiolate. Inflo- outline, and (sub)trigonous to (dorsiventrally or,
rescence terminal, more rarely pseudolateral, more rarely, laterally) lenticular in cross-section,
paniculate, corymbose, spicate, anthelate, or capi- surface smooth, punctate, reticulate or rugose,
tate (often in combinations), more rarely reduced sometimes extremely differentiated, glabrous or
to a single spikelet, open or (partly) condensed, finely hairy, sometimes seated on different types
with few to many spikelets, branching from the of cuplike structures. Seed with thin testa, free
axil of normal bracts, rarely from the axil of from the fruit wall, raphe and chalaza usually con-
prophylls; prophylls present or absent; bracts spicuous, embryo small, basal, surrounded by
leaflike or not, sheathing or not, lower primary abundant mealy or oily endosperm.
bract sometimes culmlike and erect, spikelet A cosmopolitan family with greatest diversity
bracts often small and glumelike. Spikelets with in humid and semihumid tropics, but also often
1-many glumes, in cross-section rounded, 4-5- dominant in temperate and cold temperate re-
angular, laterally or rarely dorsiventrally com- gions of the world, comprising 104 genera and
pressed. Rachilla persistent or deciduous, rarely over 5000 species, of which 2000 belong to the
disarticulating into 1-flowered parts, rachilla genus Carex.
internodes short or elongated, sometimes winged,
or thickened and curved round the ripening VEGETATIVE MORPHOLOGY. An excellent, com-
fruit(s). Lateral spikelets usually prophyllate (with prehensive, and well-illustrated description can be
a basal, 2-keeled, usually empty glume), this found in Haines and Lye (1983: 7-26).
prophyll rarely fertile, rarely utriculiform, and Plants of this family are often seen as rather
then enclosing the rachilla and its own flower. uniform and grasslike. Contrary to this opinion,
Glumes spirally arranged or distichous, rarely distorted by superficial knowledge of only a few
pentastichous, all glumes (sub)similar or of in- weedy species, Cyperaceae in fact show some very
creasing length towards the (often upper) female strange habits, admittedly often in rare species of
or bisexual flowers, deciduous or persistent; all restricted localities (Fig. 37). Many species are
glumes fertile (often except for the lowermost and indeed more or less grass-like, but often with a
142 Cyperaceae
Fig. 37A-K. Cyperaceae. Plant habits and bases. A, B Ephem-
erals (often named annuals) without rhizomes, Fuirena
leptostachya and Lipocarpha nana. C Tufted perennial,
Schoenoxiphium sparteum. D Perennial with thick rhizomes
(surculi), Carex bequaertii. E Fine underground stolons,
Kyllinga sphaerocephala. F Branching stolon, Remirea
maritima. G Overground runners, Isolepis fluitans. H Creep-
ing rhizome, Cyperus haspan subsp. platycaulis. I Stolons and
bulbs, new plant offset from bulb, Cyperus usitatus. J Stolons
and tubers, new plant directly from tuber, Cyperus bequaertii.
K Tuber set on a rhizome, Cyperus boreo-chrysocephalus.
(Haines and Lye 1983)
yellowish or brownish tinge, in many annuals and
in forest dwellers often partly reddish or purplish
colored. They range from tiny ephemerals a few
em high to tall robust forbs. There are a few lianas
with scabrid stems and leaves, and dwarf shrubs
or caudex-building species with false trunks
formed mainly by leaf-sheath remnants and
adventitious roots.
The majority of the species are perennials, while
annuals are restricted to a limited number of
genera and clearly represent highly specialized
offspring. A few of these specialists are even
recognized on the generic level, e.g., Alinula,
Courtoisina, Queenslandiella, Trianoptiles, and
Volkiella. Annuals are conspicuously absent from
the Mapanioideae and the Caricoideae, rare in the
Sclerioideae (only in Diplacrum and Scleria),
and frequent in the Cyperoideae.
Basal (mostly underground) parts of the
Cyperaceae come in a noticeable variability,
whose typology is not yet well established, partly
due to confused terminology. Haines and Lye
(1983: 8-10, Fig. 2) have presented an excellent
survey.
Perennial Cyperaceae may be tufted or even
tussocky; they may develop a short or elongated
rhizome (sometimes with bulbously thickened
internodes), subterranean stolons (sometimes
ending in a tuber or in a bulb), or superficial run-
ners. Both monopodia! and sympodial growth are
known from these organs.
The basal nodes, or many nodes from the rhi-
zomes, stolons, or runners bear many adventi-
tious roots. Some perennial species, in particular
forest dwellers, often develop thick, tough, long-
living stilt roots. Thick roots can also be seen
in many helophytes and tussock-forming
species. Swollen and markedly hairy lateral
rootlets (dauciform roots) are reported from
Carex, Cladium, Cyathochaeta, Gahnia, Kobresia,
Lepidosperma, and Schoenus (Davies et al. 1973;
Lamont 1978). These peculiar rootlets are usually
found as laterals on superficial roots. Their mer-
istematic potential is persistent, sometimes lead-
ing to successive swellings. The radial expansion is
 Cyperaceae
most marked in the epidermis. Young dauciform
roots do not exhibit endophytes.
Each stem normally grows into a culm with a
terminal inflorescence (culms central), although
the stem of some species has a continuous growth,
and flowering culms develop from the axil of some
lower leaves (culms lateral). Nonflowering culms
are rare but are normal, e.g., in Eleocharis, where
such culms bear a clearly reduced single terminal
spikelet (Eiten 1976b, Fig. 203).
Stems are mostly unbranched with a basal ro-
sette ofleaves. Cauline leaves are known in genera
such as Bolboschoenus, Carex, Didymiandrum,
Fuirena, Pycreus, Rhynchospora, and Scleria.
Branched stems are much rarer and known, e.g.,
in Bulbostylis leucostachya, Caustis (probably re-
duced inflorescence branches), Microdracoides,
lianoid Scleria, and in many floating herbs
(probably of rhizomatous nature).
Branching is from axillary buds. The lateral
branches often bear an adaxial prophyll. From the
axillary bud of this prophyll a new branch of sec-
ond order may develop, again branching from the
prophyll, etc. In this way, in many genera fascicles
of branches of decreasing length emerge from
closed leaf sheaths (Haines 1967; Goetghebeur
1986).
In some scattered species of various genera, par-
ticularly of the Sclerioideae, stems develop into a
basal "caudex", i.e., a perennial, ± woody part
formed by a relatively thin stem surrounded by a
thick layer of leaf-sheath remnants, often inter-
mingled with numerous adventitious roots.
Leaf arrangement is usually (spiro)tristichous.
More rarely, it is distichous, predominantly or
exclusively so in genera such as Chrysitrix,
Coleochloa, Costularia, Fimbristylis, Machaerina,
Nemum, and Oreobolus. In these species the culms
are often ensiform or at least compressed. In some
species of Fuirena with pentangular culms the
leaves are equally pentastichously arranged.
In some species of Carex, Didymiandrum,
Hypolytrum, and Scleria, the tristichous cauline
leaves can be arranged in false whorls.
A useful typology for culm shapes is found in
Metcalfe (1971: Fig. 8).
The leaves usually consist of a closed, tubular
sheath and a linear blade. Open sheaths are very
rare (Coleochloa and Oreobolus, both with disti-
chous leaves). Winged leaf sheaths are known in
some species of, e.g., Fuirena and Scleria. A ligule
can be found on the transition of leaf sheath and
blade in many scattered genera. It is present
in many Scirpeae (but not in Phylloscirpus),
all Dulichieae, and many Caricoideae (not in
143
Cymophyllus), but is conspicuously absent in all
Mapanioideae. On top of the leaf sheath, on the
opposite side of the leaf blade, a contraligule is
present as a ± triangular outgrowth in many
Sclerioideae. This organ is diagnostically valuable
in many species of Scleria.
Leaf blades are usually ± linear and flat,
but ancipitous to terete in (some species of)
Chrysitrix, Fimbristylis, Lepidosperma, Macha-
erina, Neesenbeckia, and Nemum. Plicate leaves
are known in Capitularina, less conspicuously in
some species of, e.g., Hypolytrum and Mapania.
The leaf blade can be (nearly) completely
reduced, as in Actinoschoenus, Androtrichum,
Chorizandra, and Eleocharis, or in some species of
many scattered genera, or in the lower leaves of
many other species.
Nonlinear, broader leaves are typical for many
forest dwellers in the Mapanioideae and Sclerio-
ideae, but also in species of Carex, Cyperus,
Pleurostachys, and Rhynchospora. In many of
these species the broad leaf blade is conspicuously
narrowed at the base into a pseudopetiole. Several
species of Scleria are known to show pseudo-
premorse leaf blades, where a continuation of the
sheath wings along the blade abruptly stops before
reaching the top of that blade. Another type of
pseudopetiolate blade is described for Oreobolus,
where the blade has a narrow basal part and a
slightly to conspicuously wider upper part. This
phenomenon is also observed in other high-
altitude species of further genera such as in
Carpha, Isolepis, and Schoenus, and even more
conspicuously in some psammicolous species like
Cyperus chordorrhizus.
Finally, the leaf blade seems asymmetrically de-
veloped (with a flat basal part, but partly laterally
flattened near the top) in species of Amphiscirpus,
Fimbristylis, and Trichophorum. This phenom-
enon, particularly conspicuous in some erect in-
volucra! bracts, needs more detailed study.
Unicellular hairs are of almost ubiquitous
occurrence in Cyperaceae in the form of prickle
hairs, consisting of an inflated base and ending in
a short sharp barb pointed towards the top of the
blade. Longer, softer hairs are of scattered occur-
rence. Generally hairs do not represent useful di-
agnostic characters.
VEGETATIVE ANATOMY. The comprehensive,
well-illustrated treatment by Metcalfe (1971) cov-
ers all aspects of the vegetative anatomy of the
family and its taxonomic significance. This data
set is upgraded by the extensive descriptions in
a DELTA database by Bruhl (1995). Among the
144 Cyperaceae
taxonomically relevant observations, the follow-
ing may merit more detailed consideration.
Silica bodies are of almost universal occurrence
in the Cyperaceae. Usually, these silica deposits
are based on the inner periclinal walls of the epi-
dermal cells near the veins and consist of one or
more conical bodies. Sometimes, the top of the
cone is flattened, representing a kind of "table
mountain", or the top is surrounded by minor
cones, the "satellites" (Ragonese et al. 1984;
Goetghebeur and Van den Borre 1989). Several
uncommon types are described and illustrated
by Metcalfe (1971: 13-17, 543-545). Differences
between silica bodies in c3 and c4 plants are
described by Lanning and Eleuterius (1989).
Cyperaceous stomata are usually paracytic and
arranged in longitudinal lines parallel to the long
axis of the leaf. Tetracytic stomata are known from
many Mapanioideae (and in some other genera
are irregularly scattered), but also in some
Schoeneae, Cryptangieae, and Trilepideae.
In longitudinal sections of leaves, a number of
axially elongated, moniliform chlorenchyma cells
have been reported by Pfeiffer ( 1927) for many
genera of the Schoeneae. This was not confirmed
by Metcalfe (1971), probably due to his study of
mainly transversal sections.
The structure of ensiform leaves was studied
and discussed by Chermezon (1929). Fisher (1971)
investigated the unusual feature of inverted vascu-
lar bundles in the leaves of Cladium.
Air cavities are characters for which Metcalfe
(1971) stated that these "are of diagnostic value at
the species level". The following examples may
illustrate that they are of systematic value also
at higher levels: (1) The four genera of the
Bisboeckelereae possess large air cavities in the
leaf blade, but these are absent in Scleria (incl.
Acriulus), thus confirming the proposed position
of Diplacrum as distinct from Scleria; (2) Carpha
(incl. Asterochaete) possesses large air cavities,
Trianoptiles none at all, also confirming the ge-
neric distinction of the latter; (3) all species inves-
tigated of the Caricoideae are homogeneous in
this respect, confirming the homogeneity of this
group.
An interesting feature, recently observed by
Porembski and Barthlott (1995), is the presence of
a velamen radicum in several Cyperaceae, notably
dwarf shrubs growing on tropical inselbergs. This
velamen is a remarkable ecological adaptation
that seems to have evolved independently, e.g., in
Bulbostylis and Cryptangieae.
One of the important anatomical characteristics
of the Cyperaceae is the presence in several groups
of a green sheath (with Kranz cells) around the
vascular bundles (Kranz anatomy), already exten-
sively studied by Rikli (1895). These Kranz types
were found to be correlated with a number of
biochemical (C4 photosynthesis), as well as other
anatomical and (ultra)structural phenomena,
hence the denomination Kranz syndrome.
These anatomical types are opposed to the gen-
eral eucyperoid anatomy type, in which the Kranz
sheath is absent; these plants correspond to the
usual C3 photosynthetic pathway. In these species
the vascular bundles are surrounded by a
mestome sheath (the nonchlorophyllous outer-
most layer of the vascular bundle), and a rather
inconspicuous, equally nonchlorophyllous sheath
of parenchyma cells. The vascular bundles and
their sheaths are usually surrounded by radially
arranged and elongated mesophyll cells.
Most of the Kranz Cyperaceae are unusual in
having a mestome sheath of sclerified cells be-
tween the Kranz sheath and the parenchyma
sheath. Ultrastructural differences were reviewed
by Ueno et al. (1988) and Ueno and Samejima
(1989).
To the present, four different types of Kranz
anatomy are known, which are easy to distinguish
by the different number and position of the vari-
ous sheaths surrounding the vascular bundles.
1. Chlorocyperus type: the vascular bundle is sur-
rounded by a Kranz sheath and a mestome sheath.
The parenchyma sheath is lacking. Observed
in Cyperus (partly), Remirea, Sphaerocyperus,
Alinula, Pycreus, Queenslandiella, Kyllinga,
Volkiella, and Lipocarpha (Raynal 1973). A re-
markable structural variant (Kranzkette) was de-
scribed by Li and Jones (1994) for Cyperus
japonicus, where large vascular bundles are linked
by 2 chains of much smaller ones along both sides
of large air cavities.
2. Fimbristylis type: the vascular bundle is sur-
rounded by a Kranz sheath, a mestome sheath, and
a third sheath of small parenchyma cells. Observed
in Fimbristylis, Crosslandia, Bulbostylis, Abild-
gaardia, Nemum, and Nelmesia (Raynal1973). Re-
cently, however, one species of Fimbristylis and
one of Abildgaardia were recorded as having C3
photosynthesis (Bruhl1995: 128).
3. Rhynchospora type: the vascular bundle is sur-
rounded by a Kranz sheath and a second incom-
plete sheath of parenchymatous cells. The
mestome sheath is lacking, therefore this type cor-
responds to the more classic Kranz anatomy of
 Cyperaceae
many other families. Observed in a few species of
Rhynchospora.
4. Eleocharis type (or sub-Fimbristylis type): the
vascular bundle is surrounded by a Kranz sheath,
a mestome sheath, and a third incomplete sheath
of parenchymatous cells. Observed in a few spe-
cies of Eleocharis and Rhynchospora.
A multiple origin of Kranz anatomy even within
genera is evident, and in Rhynchospora, infra-
generic taxa have different types (Ueno and
Koyama 1987). In Eleocharis a transition from
eucyperoid anatomy over intermediate types to
the Eleocharis type and even a Chlorocyperus-like
type could be observed (Ueno et al. 1989;
Guaglianone and Ueno 1991; Bruhl 1995).
The inner Kranz sheath of many Cyperaceae is
probably not homologous to the Kranz sheath in
most other families. Here, it could have originated
from distal xylem and phloem parenchyma cells,
as demonstrated by the presence in some
eucyperoid species of Cyperus of a complete
parenchyma sheath surrounding the xylem and
phloem (Brown 1975; Carolin et al. 1977).
bract
bract
prophyll
terminal .. -
spikelet .-·
of inflorescence
145
INFLORESCENCE AND FLORAL MORPHOLOGY.
Three structural levels are here considered, i.e.,
the general inflorescence type, the spikelet, and
the flower.
Basically, the inflorescence consists of a panicle
(of spikelets!; Raynal1971), but is often modified
into a corymb or an anthela, capitately or spicately
contracted, or reduced to a single spikelet (Fig.
38). Partial inflorescences of the first, second,
third, and higher order can be present and may
correspond to any of these types, in many differ-
ent combinations.
In some species, several combinations can be
observed, but some genera or even a few tribes are
remarkably homogeneous as to their inflorescence
type, e.g., Eleocharis with its unispiculate inflores-
cence, or the tribe Trilepideae with a panicle of
Fig. 38A-E. Cyperaceae. Inflorescence types. A Anthela of
Fimbristylis type: the inflorescence is borne on a naked culm.
B Staged panicle of Schoenoxiphium type. C Plan common to
A and B. D Terminal head. E Apparently lateral head. Inflores-
cence branches consecutively numbered. (Haines and Lye
1983)
involucrate
head
146 Cyperaceae
dense spikes, each comprising many small
spikelets.
The primary bracts are often described as in-
volucra} bracts, the lowermost bracts are usually
foliar (sheathing or not) and the uppermost very
small and bracteate, but, not rarely, all primary
bracts can be foliar (Cyperus involucratus) or all
bracteate (Cyperus papyrus). The bracts of the ul-
timate branchlets (spikelet bracts) are often barely
distinct from glumes.
Many sciaphilous species have developed large
foliar primary bracts, sometimes extremely so.
In Mapania sylvatica all leaves are reduced to a
sheath, and only the 3 lowermost primary bracts
are large and foliar. The lowermost primary bract
can be upright, continuing the culm, and very
much culmlike, the (usually condensed or re-
duced) inflorescence is bent over 90°, suggesting a
lateral position. Such pseudolateral inflorescences
occur scattered in many genera.
Fig. 39A-D. Cyperaceae. Lateral and apparently terminal
flowers. A Spikelet of Rhynchospora corymbosa. It contains a
bisexual flower subtended by bract 4 and enwrapped by glume
5, and two staminate flowers above. B Bisexual spikelet of
Scleria distans. A pistillate flower is subtended by glume 2,
above of which follow several staminate flowers. C Pistillate
spikelet of Scleria hildebrandtii. It contains 4 empty glumes
and remnants of an originally staminate part of the spikelet
above the pistillate flower. D Pistillate spikelet of Scleria
nyasensis, the solitary pistillate flower apparently terminal.
Glumes consecutively numbered. (Haines and Lye 1983)
--~~
A
Branching in the floral region usually starts
from the axil of the primary bracts (normal
branching pattern), but in many genera secondary
and higher-order branches develop from the axil
of the prophylls (prophyllar branching pattern).
These prophylls (cladoprophylls) can be well de-
veloped, but often they are :::+::: reduced or even
absent, often so in capitate inflorescences or inflo-
rescences with long-sheathing bracts. Prophylls of
branches from short-sheathed or open bracts are
often ::!:: tubular and connate around their axis,
not rarely provided with an adaxial swelling (e.g.,
Rhynchospora trichodes), conspicuously so in spe-
cies with anthelate inflorescences ("opening" the
inflorescence before anthesis). Some unusual
forms are observed, e.g., in Cyperus odoratus,
where the tubular cladoprophyll is clearly 2-
keeled and these keels are prolonged into 2
bladelike outgrowths. In the Caricoideae these
(often utriculiform) cladoprophylls can bear a
female flower, and in some Dulichieae the open
cladoprophyll can bear a complete bisexual flower
with a perianth (Meert and Goetghebeur 1979;
Kukkonen 1986). Sometimes, closely related gen-
era can be distinguished by the presence or ab-
sence of cladoprophylls (Morelotia vs. Gahnia).
The spikelet is often the inflorescence unit that
is visible to the naked eye (Eiten 1976a). Essen-
tially, the spikelet consists of an open axis
(rachilla) bearing 1-many glumes, 1-many of
which subtend a sessile flower (Fig. 39). A spikelet
7~
~:(§Jv ' '
1@2,
~ 5 2
3 ·" 3 ··· \
4
 Cyperaceae
can be terminal, i.e., on the apical end of a
branched axis, which bears at least 1 other lateral
spikelet. A lateral spikelet is subtended by a bract
on an axis that usually bears a terminal spikelet.
The first glume of a lateral spikelet is often an
adaxial prophyll, i.e., a modified glume, which is
usually 2-keeled, bifid, smaller (to very reduced or
even absent), and empty. A lateral spikelet of, e.g.,
Cyperus is often incorrectly described as having
the two lowermost scales empty. The lowermost
scale is the spikelet bract and is not at all empty: its
axillary bud has developed into the spikelet, the
next scale is the adaxial first glume or prophyll,
and this first foliar organ of the spikelet rachilla is
indeed usually empty (Raynal1971, Fig. 8).
In a pedicelled lateral spikelet, the prophyll,
when present, is separated from the next glume by
a long rachilla internode (epipodium).
In the Dulichieae the spikelet prophyll is fertile,
i.e., bearing a normal bisexual flower with peri-
anth. In the Caricoideae the spikelet prophyll of
the female or bisexual spikelets is also fertile and
bears a female flower; in female spikelets it is
the only glume present, in bisexual spikelets more
glumes are present and each of these bears a
male flower. This prophyll has developed into the
utricle, usually completely enclosing the female
flower (except for the stigmatic branches) and
(part of) the spikelet rachilla.
The lowermost and uppermost glumes of many-
glumed and many-flowered spikelets are often
empty or provided with poorly or incompletely
developed flowers. In many-flowered spikelets all
glumes are usually similar, but in few-flowered
spikelets the glume length often increases towards
the first flower-bearing glume. On the other hand,
the length of the mucro in mucronate glumes usu-
ally decreases towards the top of the spikelet.
The rachilla internodes are usually short in
many-flowered spikelets with spirally arranged
glumes, but elongated between or around the fer-
tile flower(s) in spikelets with distichous glumes.
The internodes elongate somewhat during the
fruiting stage or develop into a markedly zig-
zaging structure (which has led to sympodial
interpretations of the rachilla in genera such as
Schoen us). The rachilla at maturity breaks up into
many 1-flowered parts in Cyperus odoratus and
Dulichium.
Glumes are arranged spirally or distichously,
rarely pentastichously (Fuirena). The spiral ar-
rangement is probably a form of spirotristichy,
and in many spikelets with apparently distichous
glumes these are really spirodistichous as well.
The distichous position is not always clearly real-
147
ized and often difficult to assess in few-glumed
and few-flowered spikelets. Furthermore, several
species are known where in terminal spikelets the
glumes are spirally set, but distichous in lateral
spikelets, such as Ficinia fascicularis, Machaerina
anceps, and Rhynchospora pubera.
In a few highly specialized groups, the spikelets
are extremely reduced, with subsequent loss of
glumes, flowers, and prophylls, leading to 1-
flowered spikelets. These spikelets tend to aggre-
gate into false spikelets, structures that deceivingly
imitate the real thing, as in Carex, Diplacrum,
Lipocarpha, and Trilepis. Atavistic forms, ob-
served in Ascolepis, and rachilla remnants in spe-
cies of several genera provide strong arguments
for a morphologically correct interpretation.
Scirpodendron is one of the very few Cyperaceae
where the spikelet rachilla apparently is termi-
nated by a flower (see also Chrysitrix). The termi-
nal position is here clearly demonstrated by the
absence of the paired lateral keeled scales, which
always accompany the lower lateral flowers. The
spikelet structure in Chrysitrix (with 1 terminal,
rarely with a 2nd lateral flower, or with a split
rachilla) is highly unusual for Cyperaceae, and the
homology is not yet clearly established. The in-
terpretation presented should be considered as
tentative, although it was already defended by
Bentham (1883).
The cyperaceous flower is basically a bisexual
trimerous structure with 3 + 3 perianth parts, 3
(+ 3?) stamens, and a 3-carpellate pistil with a
single, basal, anatropous ovule, but numerous and
sometimes very strange variations are known. The
standard monocotlike position of the perianth
and stamens around the pistil was confirmed in an
ontogenetic study by Bruhl (1991).
In the Mapanioideae the regular trimerous orga-
nization is often not realized and 2 basal lateral
perianth scales are often larger, are keeled, and
bear a stamen. Such flowers are often described as
a kind of a spikelet with reduced male flowers
(scale + stamen) and a central, terminal female
flower. Here this interpretation is not accepted.
Perianth structure is extremely variable, ranging
from few to many large bristles (stiff to more deli-
cate), flat hyaline scales, or elaborate plates to
complete reduction. The perianth parts are some-
times basally ::±: connate, as in Cyathochaeta,
Eleocharis and Schoenoplectus. Where only 3 peri-
anth parts are developed, these are usually found
alternating with the stamens, i.e., opposite the flat
sides of a trigonous ovary.
The number of stamens varies from 0 (in female
flowers) to 1-2-3-6 or more, with an extremely
148 Cyperaceae
elevated number in Chrysitrix (a genus whose
floral structure is very aberrant from other
Cyperaceae and needs more attention). Com-
monly, 3 stamens are developed, and these are
situated opposite the 3 ribs of a trigonous ovary.
The filaments may be connate at the base or over
their whole length. The anthers are often sur-
mounted by a mucro-like appendage of the con-
nective, which sometimes is extremely elongated
(Chrysitrix, Scleria tepuiensis ).
The pistil is usually 3-carpellate with a :±:
trigonous ovary, a single style, and 3 stigmatic
branches. In many genera, 2-carpellate pistils can
be observed as well, 4-carpellate pistils are rather
exceptional, as in Tetraria and a few relatives. In
Mapania the pistil is 2-3-carpellate (2-carpellate
in 4-scaled flowers, 3-carpellate in 6-scaled
flowers, except for the 6-scaled and 2-carpellate M.
foxworthii). The number of stigmatic branches
can double to 4 (Tetraria, Paramapania), 6
(Neesenbeckia), or 8 (Evandra), or even triple to
9 (Tetraria).
Bicarpellate pistils can be compressed dorsiven-
trally (usually) or laterally (exceptional). The first
form is scattered over the whole family, not rarely
intermingled with 3-carpellate pistils in the same
species, on the same plant, and even in the same
spikelet. It is assumed that this form has devel-
oped through the inhibition of the development
of the anterior carpel, apparently through a rela-
tively easily controlled mechanism. The laterally
compressed bicarpellate pistil has a very restricted
occurrence in Pycreus, Queenslandiella, and
Kyllinga. For the development of this form, a new
organization of the carpels and their vascular
bundles is needed (Blaser 1941) and, in my opin-
ion, constitutes a clear synapomorphy for these
three genera.
Similarly, laterally compressed ovaries are re-
ported in Exochogyne and Rhynchospora rubra.
However, in these few-flowered spikelets an ob-
servation of the actual position is difficult and sec-
ondary displacement is known to occur rather
frequently. This problem needs a careful study of
floral ontogenetic processes.
POLLEN MORPHOLOGY AND MALE GAMETO-
PHYTE. The pollen grains are generally cuneiform
and have an indistinctly marked aperture on the
wider end and 3, mostly smaller, lateral ones.
Ellipsoid to spherical grains with a single aperture
are found, though not exclusively, in the
Mapanioideae. Size varies from 16-66~-tm.
The cuneiform shape is due to the arrangement
of the grains in the pollen sac, where they are
packed in 1 tubular layer. The pollen grains of the
Cyperaceae are degraded tetrads, in which 3 of the
4 nuclei degenerate, while the wall of the pollen
mother cells develops into an exine. The 3 vestigial
microspores are described as being separated by
walls (Shah 1962) or not (Schnarf 1931). They re-
main adpressed to the cell wall inside the pollen
mother cell at the tapering end of the grain. The
grains are 2- or 3-celled at the dispersal stage. Al-
though the exine is built on the wall of the pollen
mother cell, exine structure is stated by Erdtman
(1952) to be similar to that of other angiosper-
mous pollen grains. The terms pseudomonads or
cryptotetrads have been suggested for these pecu-
liar pollen grains.
Erdtman (1952) suspected that the monoporate
mapanioid pollen grains may not belong to this
category, but Koyama (cited in Cronquist 1968:
341-342) affirmed their tetrad nature.
EMBRYOLOGY, SEED, AND FRUIT. The single ovule
is basal, anatropous, bitegmic, and crassinucellar.
At least in some genera, an obturator is formed by
a number of funicular cells near the micropyle,
which enlarge and show dense cytoplasm.
The single macrospore develops into an embryo
sac of the regular Polygonum type. The polar nu-
clei can fuse to form the secondary embryo sac
nucleus. The pollen tube enters the embryo sac
through a degenerating synergid. The develop-
ment of the abundant endosperm is of the Nuclear
type (Tejavathi 1987). The development of the
embryo conforms to the }uncus variation of the
Onagrad type (with early cutting off of dermato-
gen initials).
Testa and tegmen are coalescent and form a thin
and delicate seed coat. The ovary wall usually
develops into a :±: 3-layered fruit wall: the cells of
the outer epidermis contain silica bodies (often
cones), the cells of the next 2-3layers develop into
longitudinal fibers, the cells of the innermost
layers develop into transversal fibers (Tejavathi
1987), although in some species shorter sclereids
are observed (Makde and Untawale 1989).
Starting with Van der Veken's (1965) study of
embryos in Cyperoideae, followed by many papers
of J. Raynal ( 1968-1973) and Haines and Lye
(1971, 1976, 1983), who used these data to support
their taxonomic decisions, the extraordinary taxo-
nomical value of embryographical data became
evident.
Many genera are characterized by a single em-
bryo type (and often their own particular form of
that type), although sometimes 2 closely related
types are known in one genus. Some tribes can be
 Cyperaceae
distinguished solely by the embryo type, e.g.,
Cypereae by the Cyperus or Ficinia type, Tri-
lepideae by the Trilepis type, Fuireneae by the
Schoenoplectus or Bolboschoenus type. Very
poorly differentiated embryos {Juncus type,
poorly developed forms of the Carex type) are
found in many Mapanioideae and Cryptangieae.
The Carex type of embryo is assumed to repre-
sent the primitive form, characterized by a basal
root cap (micropyle-oriented), lateral first leaf pri-
mordium, poorly differentiated coleoptile, trans-
versal germination slit, and poorly differentiated
cotyl (this "cotyledon" is perhaps not homologous
to the cotyledon of dicotyledons, Jacques-Felix
1988). Through differential growth of the coleop-
tile and the cotyledon or parts thereof, many dif-
ferent forms can be obtained, and Goetghebeur
(1986) was able to distinguish 16 types.
KARYOLOGY (in collaboration with Margaret
Johnson). Published data often include approxi-
mate numbers, and also in view of the frequent
misidentifications in this family, many chromo-
some number records should be considered as
doubtful and unreliable. Furthermore, the (near)
absence of data in the subfamilies Mapanioideae
and Sclerioideae is noticeably, as well as the spo-
radic counts in the tribe Schoeneae (except for
Rhynchospora). On the other hand, many genera
and species of the other Cyperoideae and also
Carex are fairly well documented. These differ-
ences are probably related to the abundant fructi-
fication, the easy germination, and the sometimes
weedy character of many species of the latter
groups.
Chromosome records in Cyperaceae range from
2n = 10 in Eleocharis and Fimbristylis to over 200
in Carex and Cyperus with practically every basic
number in between (Fedorov 1969; Goldblatt
1981, 1984; Love and Love 1975). Numbers differ
not only between genera, but even within some
species. Some workers have reported basic num-
bers for certain groups of species, with some hav-
ing almost consecutive numbers in independent
aneuploid series (Davies 1956). Nevertheless, a
basic chromosome number of x = 5 is often sug-
gested, based on the counts of 2n = 10, 20, 40, 50,
60, etc. in several genera (common numbers in
Abildgaardia, Bulbostylis, Courtoisina, Eleocharis,
Fimbristylis, Kyllinga, Lipocarpha, Pycreus, more
rarely found in Blysmus, Bolboschoenus, Carex,
Cyperus, Eriophorum, Isolepis, Schoenoplectus,
and Scirpus).
The unusually large range of numbers is mainly
due to the presence of diffuse centromeres which
149
enable an increase in chromosome number by
chromosome fragmentation (agmatoploidy). The
fragments are able to stabilize within the karyo-
type because the centric activity is found along
the entire length of each chromosome (Braselton
1971). Hakansson (1954, 1958) first proposed the
existence of diffuse centromeres in Eleocharis, al-
though Battaglia (1954) and Sanyal and Sharma
(1972) found only chromosomes with localized
centromeres in Eleocharis uniglumis and Indian
Eleocharis. Bir et al. (1993), however, showed that
both types coexist in Eleocharis. Both types are
also thought to occur in some other genera includ-
ing Carex, Cyperus, and Bulbostylis.
Although there are scattered reports of diffuse
centromeres from various genera in other fami-
lies, their frequent occurrence in the Juncaceae
provides an important argument in support of
a close relationship between Cyperaceae and
Juncaceae.
In spite of the many high chromosome numbers
found in the family, recorded instances of poly-
ploidy are few. Natural hybrids between species
are common, often being vigorous and, in some
cases, more successful than one or both parents
(Cayouette and Morisset 1985).
PoLLINATION. Most, if not all, Cyperaceae are
proterogynous. Due to the fact that the spikelets
are often many-flowered or, if few-flowered, then
often crowded, female (receptive stigmas) and
male phases (pollen-shedding anthers) of flower-
ing are present in a single spikelet or inflores-
cence. In spikelets, flowering is usually acropetal,
as expected.
The Cyperaceae are clearly adapted to anemo-
phily, as is evidenced by their small, inconspicu-
ous flowers and hidden or reduced perianth, the
long stigmatic branches, the filaments elongating
considerably during anthesis, and anthers shed-
ding abundant pollen. Nevertheless, scattered spe-
cies in various genera are known to be visited by
pollen-gathering insects or have traits indicating
pollination by insects (and other invertebrates).
These include in the Mapanioideae species of
Hypolytrum (white flowering), Mapania (fra-
grance, red pigments), in the Cyperoideae species
of Ascolepis (several species with white, yellow, or
orange daisylike inflorescences), Bolboschoenus,
Cyperus, Eleocharis, and Ficinia (condensed head
with yellow bracts), Rhynchospora sect. Dichro-
mena (many species with white bracts, excellently
described and illustrated by Thomas 1984), in the
Caricoideae the genera Carex and Cymophyllus.
No records are known for Sclerioideae. Many of
150 Cyperaceae
these species are wind-pollinated as well, and even
self-pollination definitely occurs.
Entomophily is probably often linked to the for-
est habitat, where wind velocity is much reduced
and other pollen vectors are needed (Lorougnon
1973). The morphological adaptations for ento-
mophily seem to have arisen in several parallel
lines, and even in a small group like Rhynchospora
sect. Dichromena, white bracts have evolved
independently at least 3 times (Thomas 1984).
DISPERSAL. The cyperaceous fruit is usually an
achene, which is sometimes corky thickened, or
rarely drupelike. Often, however, it is more or less
closely surrounded by other, persistent organs
such as a hardened style base, a perianth (often
bristlelike), elongated filaments, 1 to few glumes,
parts of the spikelet axis, or even complete spike-
lets. Sexual diaspores are thought to be adapted to
dispersal by several means:
1. Seawater and freshwater: fruit with (partly)
corky thickened fruit wall (Oxycaryum, Cyperus
gardneri); fruit clasped by a thick corky rachilla
internode (Remirea); fruit surrounded by an in-
flated utricle (Carex ); fruit wall with air cavities
(Cladium); presence of long or widened perianth
bristles holding an air bubble (Eleocharis, Fuirena).
2. Birds: attraction through presence of abundant
food (Cyperus), sometimes by a display of con-
trasting colors (Gahnia) or by aril imitation
(Scleria); adaptation for epizoic dispersal by de-
velopment of a strong hook on top of the rachilla
(Uncinia), or by shedding numerous small fruits
into the surrounding mud (Fimbristylis).
3. Ants: some species of Carex, in particular those
inhabiting forest regions, with an oil-rich append-
age at the base of the utricle, Lepidosperma with
thick fleshy perianth parts.
4. Man: some species of Cyperus forming many long
slender stolons ending in a tuber easily detached
from the plant and capable of withstanding long
periods of unfavorable circumstances; other weedy
species having an elevated level of fruit production,
e.g., Cyperus iria, Schoenoplectus in rice fields.
5. Wind: in species of several genera the ripe fruit
is surrounded by long bristles, formed either by
elongated filaments (Androtrichum, Machaerina)
or by perianth hairs (Afrotrilepis, Eriophorum,
Scirpus), and the latter hairs may be conspicu-
ously plumose (Carpha, Costularia).
These specialized structures could indicate
adaptations to dispersal by one of these means,
but the number of actual observations is rather
small, and experimental control is needed for
confirmation.
Some species are known to be amphicarpous (or
basicarpous) (Haines 1971; Raynal 1976; Brown-
ing 1992; Bruhl1995); others are capable of build-
ing up a long persistent seed bank, e.g., Carex
pilulifera. Such phenomena should also be taken
into account when studying seed dispersal.
On the other hand, many species propagate veg-
etatively through extensive growth (and eventual
fragmentation) of rhizomes or stolons, more
rarely through the formation of tubers and even
bulbs, which may make them troublesome weeds
(Cyperus esculentus, C. rotundus). In many
helophytes, culms tend to curb down and start
forming new shoots in the floral region, a process
that is well known in the popular umbrella plant
(Cyperus involucratus ).
An excellent and more comprehensive review
of dispersal biology can be found in Kern {1974:
442-444).
PHYTOCHEMISTRY (by R. Hegnauer). A survey of
the vast phytochemical literature pertinent to the
Cyperaceae was given by Hegnauer (1963, 1986).
Metabolically, the Cyperaceae are similar to the
Poaceae in several respects. Both families have
starchy seeds, are silica accumulators, store
kestose- and isokestose-type oligosaccharides in
vegetative parts of all or some of its members and
have p-coumaric and ferulic acids esterified with
non-lignified cell walls (Harris and Hartley 1980).
Moreover, there is a rather convincing overall
similarity in flavonoid metabolism of the two
families, as evidenced by strong tendencies to
accumulate flavones (among others tricin and
luteolin-5-glycosides) and C-glycoflavones rather
than flavonols. Alkaloids and cyanogenic com-
pounds are rare in the Cyperaceae, in spite of as-
sertions to the contrary with regard to the latter
compounds by Dahlgren et al. (1985).
The morphology of epicuticular wax deposits
also suggests affinities between the Cyperaceae
and Poaceae and other members of Commelini-
florae, and of Bromeliiflorae, Zingiberiflorae, and
Areciflorae (all sensu Dahlgren et al. 1985). In all
these taxa Strelitzia type wax deposits occur
(Frolich and Barthlott 1988). It seems worthwhile
to mention that flavonoid and protoantho-
cyanidin metabolism also suggests affinities
between Cyperaceae, Juncaceae, Poaceae, and
Arecaceae (Harborne in Dahlgren and Clifford
 Cyperaceae
1982). With regard to the classification within the
family, phenolic and quinonoid constituents seem
to be of great promise. The same is probably valid
also for morphological aspects of silica and epicu-
ticular wax deposits.
CLASSIFICATION. The following subfamilies and
tribes can be recognized, which are characterized
by a combination of morphological characters
mainly referring to flower, inflorescence, and em-
bryo structure. With more data from molecular
studies becoming available, this classification will
no doubt need to be adjusted.
A most daring and deserving new classification
of the family has been suggested by Bruhl (1995).
The many differences with the system here pro-
posed have their origin in different interpretations
of morphological data, differences in contents
and size of the data sets, and a more strict applica-
tion of cladistic principles by Bruhl (1995). In
particular, I do not agree with many of Bruhl's
interpretations of morphological structures and
the great significance he attributes to anatomical
data (whose variability on the generic and specific
level is unknown). This must have strongly
influenced his results, in which sometimes un-
likely couples of genera are grouped together. A
discussion of these problems will be given
elsewhere.
In the following enumeration of suprageneric
taxa the approximate numbers of genera/species
are given in parenthesis.
I. Subfam. Mapanioideae (13/140) comprises a
number of genera which have a peculiar floral
structure: the lateral bisexual flowers are provided
with a lateral pair of usually larger, keeled, lami-
nar hypogynous scales.
1. Tribe Hypolytreae (9/130) is characterized by
large-leafed species, usually many spikelets per
inflorescence, and (very) poorly differentiated
embryos. These species are usually equatorial
forest dwellers.
2. Tribe Chrysitricheae (4/13) comprises four
small genera of :±:: open swamps, with a reduced
vegetative apparatus and inflorescence, and
probably a highly differentiated embryo. Their
phylogenetic relations are not clear. Hellmuthia
(upper flowers scaleless) and Chrysitrix (unusual
floral structure) are somewhat aberrant in this
subfamily. The disjunct Southern Hemisphere
distribution of this tribe suggests a Gondwana
origin.
151
II. In subfam. Cyperoideae (71/2380) at least 1
flower per spikelet is bisexual (with a few rare and
clearly derived exceptions), but the mapanioid lat-
eral pair of keeled hypogynous scales is lacking.
Apart from these symplesiomorphies, no other
character state can be used to circumscribe
this large and extremely diverse subfamily. Fur-
ther tribal subdivision is problematic because of
the high degree of morphological diversity of
nearly all tribes. The embryographical data are
extremely useful on this level, however, and are
here considered as highly indicative of natural
relationship.
3. Tribe Scirpeae (6/60) is characterized by
spikelets with spirally arranged glumes, which
usually are all fertile, and flowers with hypogy-
nous scales, at least in part of the species. Since
all of these characters are clear plesiomorphies,
this tribe most probably is a remainder
composed of genera without any pronounced
characters. The embryo is of the Carex, Schoen us,
or Fimbristylis type, three semophyletically close
types.
4. Tribe Fuireneae (5/90) is rather similar to the
preceding tribe as to its floral morphology, but
each of its genera possesses one or more highly
specialized structures. Their well-differentiated
embryos belong to the Schoenoplectus or
Bolboschoenus type and indicate a status as a
specialized offshoot.
5. Tribe Eleocharideae (3/200) is recognized
by its reduced vegetative apparatus, the fixed
unispiculate inflorescence, the unique embryo
type and the helio- and helophilous ecology. This
tribe seems related to both the previous and
(more) to the following tribe. Characters in
common with many Fuireneae include vegetative
morphology, bristlelike perianth, embryo with
broadened cotyledon. Similarities with the
Abildgaardieae are Kranz syndrome (rare), disti-
chous glumes (rare), differentiated and thickened
style base, moniliform stigmatic hairs.
6. Tribe Abildgaardieae (6/420) is remarkable
for its clearly differentiated style base, which is
often thickened and deciduous or persistent
on the achene. Distichous glumes appear for the
first time independently in several genera. The
Fimbristylis, Bulbostylis, and Abildgaardia type
embryos are semophyletically related. Also the
Kranz syndrome is present in nearly all species
of Abildgaardia, Bulbostylis, Crosslandia, Fimbri-
152 Cyperaceae
stylis, Nelmesia, and Nemum (Fimbristylis type).
Moniliform stigmatic hairs are reported in the
latter six genera.
7. In tribe Cypereae (19/900) the style base is not
distinct, not thickened, but sometimes persistent
as a narrow beak. Distichous glumes are common,
as well as a pronounced reduction of the spikelet
in the more specialized genera. The perianth has
disappeared completely. The Cyperus- or the very
similar Ficinia-type embryos are highly character-
istic for this group. The Kranz syndrome is also
very frequently observed here.
8. Tribe Dulichieae (3/10) is formed by three
small genera only, united by an exceptional char-
acter: the common presence of a fertile spikelet
prophyll (and fertile cladoprophyll, when
present), bearing a bisexual flower, plus a very
similarly formed embryo of the Carex type.
9. In tribe Schoeneae (29-700) are united nearly
all Cyperoideae with a (mostly) restricted number
of bisexual flowers per spikelet, often provided
with a ± well-developed perianth. The spikelets
are remarkable for the inclusion of the flower(s)
by the wings of the next glume. Carex and
Schoenus type embryos are prevalent, although
some more specialized forms (Helothrix type,
Carpha type) and a more reduced (secondary?)
form (Juncus type) are also known. The Kranz
syndrome is known from only a few species of
Rhynchospora. This group probably needs a fur-
ther division, but far more data are needed to de-
cide on this problem.
III. Subfam. Sclerioideae (15-340) is recognized
by spikelets with few glumes, 1-few of which each
subtend a strictly unisexual flower. A (sometimes
vestigial) perianth is often present. The subfamily
is perhaps heterogeneous: four tribes and two un-
classified genera are easily recognized.
10. In tribe Cryptangieae (4/50) the spikelets are
unisexual, with spirally arranged glumes, and a
perianth usually formed by 3 fimbriate scales op-
posite the flat sides of the achene. The Juncus type
embryo needs to be confirmed in more genera and
species.
11. Tribe Trilepideae (4/15) is characterized by a
panicle composed of many dense spikes of many
tiny spikelets with few distichous glumes. The pe-
rianth is as in the preceding tribe, and the embryo
is of the specialized Trilepis type.
12. Tribe Sclerieae (1/250) is formed by the
single, large genus Scleria. It has a basically pan-
iculate inflorescence, the spikelets are bisexual or
unisexual, the achene is surrounded at the base by
a hypogynium and a cupula, the homology of
which is rather obscure. The Fimbristylis type
embryo is often very well developed.
13. Tribe Bisboeckelereae (4/25) is recognized
by the (sometimes connate) empty (?) glumes
surrounding the apparently terminal (?) female
flower. Also the structure of the male spikelets is
unusual (glumes each with a single stamen). The
embryo belongs to the Carex, Schoenus, or
Fimbristylis type.
14. Under incertae sedis are included the two mo-
notypic genera Exochogyne and Koyamaea, whose
combination of characters does not fit any of the
tribes in Sclerioideae.
IV. Finally subfamily Caricoideae (5/2150) with
the single tribe Cariceae is easily recognized by the
unisexual flowers, the female flowers being com-
pletely enclosed by a fertile utriculiform spikelet
prophyll (often with 2 strong nerves), the absence
of perianth parts, and the embryo of the Carex or
very rarely of the Schoenus type.
AFFINITIES. In the past, different claims for
a close relationship between the Cyperaceae
and the Pandanales, Poales, and Juncales have
been made. A broad comparison of data from
many different sources points to a divergence
Restionaceae-Poaceae and Juncaceae-Cyperaceae
(Dahlgren and Rasmussen 1983; Dahlgren et al.
1985). The close relationship between Cyperaceae
and Juncaceae is also supported by more recent
morphological and molecular analyses (Linder
and Kellogg 1995; Simpson 1995). The presence of
diffuse centromeres and the meiosis of the
postreductional type may be considered as par-
ticularly strong arguments in favor of the close
relationship between Juncaceae, Cyperaceae, and
Thurniaceae. The possession of tetrad pollen in
the three families sometimes used to underpin
this relationship is not conclusive, because the
pollen tetrads in Juncaceae/Thurniaceae are
fundamentally different from the pseudomonads
of Cyperaceae.
DISTRIBUTION. The family has a worldwide dis-
tribution but is surprisingly absent from Antarc-
tica. It ranges from sea level to up to 5475m
( Carex nivalis Boott in the Karakorum), covering
 Cyperaceae 153

large areas of swamps and arctic vegetations. Fimbristylis are closely linked by similar rusts.
There are concentrations of genera in northern Carex is the host of at least three lineages of ad-
S America (Sclerioideae), southern Sudano- vanced species of Puccinia. These three lineages
Zambesian Africa (Cyperoideae), and SW Austra- are present as well in ]uncus and Luzula
lia (Schoeneae). Some of the larger genera are (Juncaceae), confirming the close relationship
particularly well developed (in number of species) between Cyperaceae and Juncaceae.
in certain regions: Eleocharis and Rhynchospora in Some species of Kobresia are known to have es-
tropical America, Fimbristylis in Malesia and adja- tablished a mutualistic symbiosis with ectomy-
cent N Australia, Bulbostylis, Cyperus, and Pycreus corrhizal fungi. The hyphae of these fungal
in tropical Africa, Lepidosperma, and Schoenus in symbionts permeate large volumes of soil and
Australia. obtain scarce elements, in particular phosphorus,
Endemism on the generic level is pronounced in which they pass on to the plant in exchange for
(tropical) America (21), tropical and S Africa (18) photosynthetates (Harley and Smith 1983).
and Australia (incl. New Zealand and New
Caledonia) (13), less so in tropical Asia with four FosSIL CYPERACEAE. (by Jan De Coninck, Gent).
endemic genera (plus six more extending into Fossils of Cyperaceae (mainly fruits or pollen,
Australia). Endemism on the specific level is much rarely other plant parts) are known with a reason-
more difficult to assess due to the lack of recent able amount of certitude from Paleocene times
revisions and reliable continental checklists. onward. Alleged late Cretaceous records are prob-
Many species are known from one single speci- ably not cyperaceous, or doubtful at least. There-
men or very few collections or a single locality. fore it seems reasonable to set the appearance of
this family in the Paleocene.
FUNGUS RELATIONSHIPS (by K. Van der The Cenozoic history of the Cyperaceae is rather
Gucht). According to the Fahrenholz rule, more well documented for N America and NW and
primitive taxa of parasites generally will occur central Europe. In Australia, New Zealand, and
on more primitive hosts, and vice versa. The dis- Antarctica, the record is less substantial. Only few
tribution of Ustilaginales in Cyperaceae seems to data on cyperaceous remains from African and NE
confirm this statement (Vanky 1995, 1996, and Asian Cenozoic deposits are available, and for S
previous papers). Thus, the more primitive Asia and S America records are even more scarce.
Cintractia is observed on many Cyperoideae,
but not on Caricoideae, while the specialized EcoNOMIC IMPORTANCE. The Cyperaceae are
Anthracoidea is only found on Caricoideae, and generally of limited economic value and often
within this subfamily only on the more specialized used locally. In a recent checklist (Inglis 1994),
taxa such as Carex subgen. Vignea, unispicate spe- well over 300 species are recorded for a variety of
cies of Carex, Uncinia, but not on Schoenoxiphium uses in many parts of the world. They are also of
or on Carex subgen. Indocarex. growing horticultural interest.
One single species of Anthracoidea, A. scirpi Several species are well known as noxious
(Kuhn) Kukkonen, is an exclusive parasite of weeds, among them the world's worst weed,
Trichophorum caespitosum and presents a good Cyperus rotundus (purple nutsedge), which, due
example of a major taxonomical jump to an eco- to its very resistant subterranean tubers, is ex-
logically associated species (Kukkonen 1964). tremely difficult to eradicate. Cyperus esculentus
The nearly complete lack of data for subfamilies (yellow nutsedge) is also an agressive weed
Mapanioideae and Sclerioideae is remarkable, but that recently has invaded NW Europe from the
probably reflects our poor knowledge rather than Americas. Several species of Cyperus, Eleocharis,
a real absence. Fimbristylis, and Schoenoplectus figure as impor-
Uredinales present a more complex picture. The tant weeds in SE Asia (Soerjani et al. 1987).
more primitive members of the more primitive Species of various genera are used for weaving,
genus Uromyces are known as parasites on thatching, or plaiting mats, such as Actinoscirpus
Cyperus and Scirpus. Bolboschoenus and Schoeno- grossus, Cladium mariscus, Desmoschoenus
plectus are infected by Uromyces, the more ad- spiralis, Eleocharis sphacelata, Lepironia arti-
vanced Schoenoplectus also by the more advanced culata, and Schoenoplectus lacustris.
Puccinia. Eriophorum (including Eriophoropsis) Cyperus involucratus is a well-known house
and Trichophorum, attacked by more specialized plant in temperate regions and, together with
species of Puccinia, emerge as 2 monophyletic Cyperus papyrus, a garden favorite in warmer
groups that arose from Scirpus. Eleocharis and countries. Species of Caustis are cultivated for
154 Cyperaceae

horticultural purposes, the beautifully curved - Not as above 12

dried stems being used in everlasting bouquets. 12. Inflorescence with many solitary spikelets; fruit ovoid to
ellipsoid, smooth and shining black. Australia, S New
Several species of Mapanioideae probably could Guinea) 2. Exocarya
provide bright innovations in tropical gardens - Inflorescence with digitate or capitate spikelets 13
with their orange to pinkish young shoots and 13. Inflorescence usually 3-4-anthelate; spikelets numerous,
leaves. small (OS:3mm), digitately arranged. N South America
Species of Schoenoplectus are used in sewage 6. Mapaniopsis
- Inflorescence 1-2-anthelate; spikelets many, larger,
treatment, for land-winning activities, and in scat- capitately arranged 5. Mapania
tered localities for building small boats or rafts, 14. Inflorescence paniculate to subcorymbose 15
the most famous being the totora boats of Lake - Inflorescence capitate or reduced to a single spikelet 18
Titicaca. 15. Inflorescence branches very short, panicle densely con-
tracted; spikelets with few lateral flowers ( + basal pair of
Cyperus papyrus, the source of ancient (and keeled scales) and 1 terminal flower (without such pair).
now tourist) papyrus rolls, is gaining interest as a SE Asia, N Australia, Polynesia 1. Scirpodendron
possible source for bioenergy. - Inflorescence more open; all flowers lateral ( + basal pair
of keeled scales) 16
16. Style 2-branched go to 8
KEY TO THE GENERA - Style 3-branched 17
17. Hypogynous scales 2, connate into a tube. W Africa
1. Flowers bisexual, with at least a basal, lateral (sometimes 4. Principina
::!:: connivent to connate) pair of ::!:: keeled laminar - Hypogynous scales 4-6 5. Mapania
(usually delicate) scales, or flowers with apical pistil(s) 18. Style 2-branched go to 8
surrounded by numerous (up to 100) stamens, each with - Style 3-branched 19
a subtending linear scale (Mapanioideae) 2 19. Hypogynous scales 2, only present in lower flowers of the
- Flowers bisexual or unisexual, without a lateral pair of spikelet. Coastal S Africa 12. Hellmuthia
keeled laminar scales 20 - Hypogynous scales 4-6 5. Mapania
2. Apical pistil(s) surrounded by numerous (up to 100) 20. All flowers unisexual 21
stamens, each with a subtending linear scale. S Africa - All flowers bisexual, or at least 1 flower bisexual, other
(SW Cape), SW Australia 13. Chrysitrix flowers usually male (Cyperoideae) 45
- Stamens surrounding the pistil <20, (lower) flowers with 21. Female flowers each axillated by and enclosed in a ::!::
a basal lateral (sometimes ::!:: connivent to connate) pair utriculiform prophyll (each subtended by a bract),
of ::!:: keeled scales 3 usually several-many together, and forming a female
3. Culms 4-5-angular, hollow, terete, septate; inflorescence spike or the basal (rarely the apical) part of a bisexual
capitate or reduced to 1 spikelet. New Guinea, Solomon spike (Caricoideae) 22
Is. 9. Capitularina - Not as above (Sclerioideae + abnormal species of
- Not as above 4 Cyperoideae) 26
4. Inflorescence reduced to a single, pseudolateral spikelet; 22. Rachilla of female spikelet protruding out of the utricle,
culms (sub )terete, ::!:: septate, all leaves reduced to a with an apical hook, formed by 1 empty recurved glume;
sheath 5 inflorescence a single spike with many basal female
- Not as above spikelets ("utricles") and a terminal male spikelet
5. Spikelets fusiform. Madagascar to Polynesia 102. Uncinia
11. Lepironia - Rachilla not protruding, or different 23
- Spikelets (depressed) globose. Australia, New Caledonia 23. Rachilla usually small and not protruding (rarely pro-
10. Chorizandra truding), utricles always completely closed (excepted for
6. Inflorescence with small primary bracts (often much a small apical orifice) 24
shorter than 6 em), or reduced to a single spikelet; culms - Rachilla protruding in at least a few spikelets; usually
lateral (rarely central) 7 some of the utricles partly open 25
- Inflorescence with larger, leaflike primary bracts 10 24. Inflorescence an ovoid to spherical spike, with white bracts
7. Style 2-branched 8 and glumes; style branches 3, conspicuously thickened-
- Style 3-branched 9 clavate; leaves reduced to a short sheath, only 1-few leaves
8. Hypogynous scales 2 (-3), often connate to various with a large, wide blade, without midnerve, margins
degrees 3. Hypolytrum conspicuously undulate. SE USA 104. Cymophyllus
- Hypogynous scales 4-6 5. Mapania - Not as above 103. Carex
9. Keel of basal lateral paired scales coarsely toothed; inflo- 25. Rachilla broadly linear, often with serrate margins; spike
rescence simply anthelate or reduced to a single spikelet. prophylls often utriculiform, subtending a female flower.
SE Asia, Polynesia 7. Paramapania S Africa, E African mountains, Madagascar
- Keel of basal lateral paired scales ciliate; inflorescence 101. Schoenoxiphium
capitate or reduced to a single spikelet 5. Mapania - Rachilla usually linear, not serrate; spike prophylls not
10. Inflorescence anthelate; style 2-branched 11 utriculiform, without flower; prophylls of female spike-
- Inflorescence paniculate, subcorymbose, capitate, or lets utriculiform but often more or less open. Holarctic,
reduced to 1 spikelet; style 2-3-branched 14 N Sumatran mountains 100. Kobresia
11. Basal margin of primary bracts with coarse yellowish 26. Inflorescence paniculate, with dense spikes of many
teeth; spikelets large, fusiform, glumes coriaceous. Tropi- few-glumed and 1-2-flowered spikelets, with fusiform,
cal S America 8. Diplasia long-beaked achenes, surrounded at the base by 3 long-
 Cyperaceae 155

fimbriate small scales opposite the sides of the achene - Female flower surrounded by ca.lO empty glumes; inflo-
(Trilepideae) 27 rescence paniculate with corymbose (rarely capitately
- Not as above 30 contracted) partial inflorescences. Tropical America
27. Flowers dioecious; leaves crowded on top of (branches 94. Becquerelia
of) a caudex, blade deciduous, leaving a truncate sheath. 40. Achene compressed ellipsoid or rounded trigonous, en-
W Africa to Cameroon 92. Microdracoides closed by a pubescent, nerveless glume; female flower
- Flowers monoecious; leaves different 28 surrounded by 3 glumes. Tropical America
28. Leaves distichous, with open sheath and deciduous blade; 97. Calyptrocarya
spikes bisexual. From Cameroon to E and S Africa, - Achene subtriquetrous, enclosed by a glabrous, many-
Madagascar 91. Coleochloa nerved glume; female spikelet surrounded by 2 glumes.
- Leaves spirally arranged, sheath closed, blade not Tropical America 96. Bisboeckelera
deciduous 29 41. Leaves basal, few, reduced to a tubular,::!:: hyaline sheath;
29. Leaves with ciliate ligule; spikes and spikelets 1-2-sexual. inflorescence reduced to 1 spikelet
W Africa to Gabon 90. Afrotrilepis 25. Eleocharis (E. chamaegyne)
- Leaves eligulate; spikes always unisexual. S America - Not as above 42
89. Trilepis 42. Glumes (spiro)distichous; leaves with well-developed
30. Spikelets with densely velvety hairy, cucullate glumes. S blade 43
Venezuela, Cerro de Ia Neblina 99. Koyamaea - Glumes spirally arranged; leaves reduced to a sheath 44
- Not as above 31 43. Basal spikelets many; rachilla of aerial spikelets with
31. Achene 2-merous, laterally compressed, without scales at short internodes, lower flowers male, upper flowers
the base; primary bracts navicular, spathelike. Central female. Australia 29. Crosslandia
and S America 98. Exochogyne - Basal spikelets absent; rachilla of aerial spikelets with
- Achene 2- or 3-merous, but not laterally compressed; elongated internodes, spikelets unisexual. Subtropical
primary bracts not navicular nor spathelike 32 and tropical America 41. Cyperus (C. can us,
32. Achene 3-merous, at the base with 3 small ciliate to C. dioicus, C. schomburgkianus)
fimbriate scales opposite the sides of the achene 44. Inflorescence open paniculate; hypogynous bristles 6. S
(Cryptangieae) 33 Africa 21. Pseudoschoenus
- Achene without hypogynous scales, but sometimes with a - Inflorescence capitate; hypogynous bristles absent. S
::!:: 3-lobed disk, but these lobes opposite the ribs, rarely Africa, Botswana 37. Scirpoides? ("Scirpus" dioicus)
with hypogynous bristles 36 45. Leaves basal, usually 2, reduced to a tubular, often ::!::
33. Caudex conspicuous; flowering culm lateral, conspicu- hyaline leaf sheath; vegetative and generative branchlets
ously compressed; hypogynous scales long-fimbriate 34 usually similar; inflorescence reduced to 1 terminal
- Caudex short or absent; flowering culm terminal, spikelet; lowermost scale shorter than the spikelet,
trigonous; hypogynous scales (shortly) ciliate 35 usually empty, rarely axillating a flower, other glumes
34. Inflorescence large, open paniculate; style base conical usually each axillating a flower; style base usually distinct,
87. Everardia thickened and persistent 25. Eleocharis
- Inflorescence small, compacted-capitate; style base - Not as above 46
clavate 88. Cephalocarpus 46. Bisexual flowers with hypogynous bristles or scales (the
35. Leaves broadly linear to lanceolate, pseudoverticillate in single adaxial rachilla remnant of some genera should
2s or 3s along a tall, woody culm; flowers dioecious not be confused with a bristle!) 47
86. Didymiandrum - Bisexual flowers without hypogynous bristles or scales
- Leaves linear, not pseudoverticillate; flowers usually 101
monoecious, spikelets unisexual 85. Lagenocarpus 47. Glumes (or glumelike scales) distichous, or number of
36. Female spikelets !-flowered or bisexual spikelets with 1 glumes reduced to 2 48
female flower; style base usually not distinct, not thick- - Glumes (or glumelike scales) spirally arranged 69
ened; most leaves with well-developed blade 37 48. Style 2-branched 49
- Female spikelets many-flowered; if 1-flowered, then style - Style 3 (-more)-branched 54
base distinct, thickened, persistent, and leaves reduced 49. Floating to submerged hydrophyte with many pseu-
to a tubular, ::!:: hyaline sheath (abnormal species of doverticillate vegetative branchlets and few, stouter
Cyperoideae) 41 generative culms; inflorescence of 1 spikelet; spikelet
37. Part of the spikelets bisexual with 1 female flower; if all with 2 elongated (8-12mm) glumes; hypogynous bristles
spikelets unisexual, then the female spikelets with (3-) 6-11, longer than the fruit 27. Websteria
reduced male flowers, or with a reduced rachilla top; - Not as above 50
male flowers usually 3-staminate; contraligule usually 50. Culms terete, hollow; inflorescence paniculate; prophyll
well developed 93. Scleria of spikelet axillating a bisexual flower. N America
- All spikelets unisexual, female spikelet !-flowered, with- 53. Dulichium
out any traces of the other sex, rachilla top completely - Not as above 51
reduced; male flowers usually !-staminate; contraligule 51. Style base deciduous 52
usually not developed (Bisboeckelereae) 38 - Style base persistent, often conspicuously thickened 53
38. Achenes glabrous (smooth or with various ornamenta- 52. Leaf blade up to 30 em long; glumes glabrous; fruit re-
tions), not enclosed by a utriculiform glume 39 markably incurved and dorsally concave. Australia
- Achenes rather tightly enclosed by either a shortly pubes- 66. Cyathochaeta
cent or a many-nerved utriculiform glume 40 - Leaf blade reduced; glumes fimbriate; fruit not incurved.
39. Female flower surrounded by only 2 empty glumes; in- New Zealand 62. Schoenus (S. tendo)
florescence capitate, or paniculate with capitate partial 53. Hypogynous bristles scabrid below, plumose upward.
inflorescences. Pan tropical 95. Diplacrum Tropical America 61. Pleurostachys
156 Cyperaceae

- Hypogynous bristles scabrid or not 60. Rhynchospora - Inflorescence prophyllate or not; stamens 3-6 (-8); hypo-
54. Cushion-forming plants; leaves (spiro) distichous, with gynous bristles 0-3-6 (or more), rather short, usually
open sheath; hypogynous scales or bristles persistent on linear, scabrid, basally not forming a disk; achenes usu-
the rachilla 67. Oreobolus ally with a large, thick, often scabrid beak 70. Tetraria
- Not as above 55 69. Style 2-branched 70
55. Inflorescence capitate; leaves ligulate 56 - Style 3 (or morel-branched 77
- Inflorescence (depauperate) paniculate; leaves often 70. Prophyll of spikelet axillating a bisexual flower 71
eligulate 59 - Prophyll of spikelet empty (or prophyll of spikelet
56. Upper internodes of rachilla elongated and prominently ~e~ 72
zig-zag; hypogynous bristles up to 6 62. Schoenus 71. Inflorescence anthelate. Sumatra 54. Sumatroscirpus
- Rachilla internodes short, or only uppermost internode - Inflorescence spikelike. Holarctis 55. Blysmus
somewhat elongated; hypogynous bristles 1-3 57 72. Style base ::!:: distinct, thickened and persistent (see 53)
57. Hypogynous bristles short, poorly developed; ligule - Style base not distinct, not thickened 73
ciliate. Australia 63. Gymnoschoenus 73. Spikelet 1-2 flowered; achene dorsally incurved.
- Hypogynous bristles > achene; ligule not ciliate 58 Australia 66. Cyathochaeta
58. Hypogynous bristles slender, plumose below, scabrid - Spikelets many-flowered; achene different 74
above; style base persistent, glabrous. Australia 74. Leaves usually bladeless or with a short blade; inflores-
65. Ptilothrix cence pseudolateral 24. Schoenoplectus
- Hypogynous bristles (at maturity) broadened and gla- - Leaves with well-developed blade 75
brous below, scabrid above; style base deciduous, slightly 75. Leaves eligulate 22. Bolboschoenus
scabrid. Australia 64. Mesomelaena - Leaves ligulate 76
59. Anthers conspicuously greenish yellow; leaves eligulate 76. Inflorescence pseudolateral, with spikelet or capitate
60 with few spikelets. America 18. Amphiscirpus
- Anthers not greenish yellow 61 - Inflorescence terminal, anthelate with numerous spike-
60. Annual plants with frequent amphicarpy; hypogynous lets. SE Asia 14. Scirpus (S. ternatanus)
bristles 6, 3 very short, and 3longer than achene. S Africa 77. Spikelets with several empty glumes, only 1-2 (rarely
69. Trianoptiles more) glumes, each subtending a flower 78
- Perennial plants without amphicarpy; hypogynous - Spikelets usually many-flowered, empty glume(s) rare
bristles 6, of subequallength 68. Carpha 82
61. Leaves distichous, laterally compressed to subterete, 78. Hypogynous bristles rarely developed, small; glumes and
never dorsiventrally flattened 82. Machaerina (elongated) filaments showing several types of fixing
- Leaves spirally arranged, if distichous, then dorsiven- mechanisms. SE Asia, Australia, Pacific islands
trally flattened 62 75. Gahnia
62. Partial inflorescences conspicuously corymbose; achene - Hypogynous bristles or scales 6, well developed 79
narrowly fusiform, stipitate and long-beaked. Venezuela 79. Hypogynous scales laminar 80
81. Rhynchocladium - Hypogynous scales rather linear, bristlelike 81
- Not as above 63 80. Hypogynous scales thick, spongy, deciduous with the
63. Leaves reduced to a sheath, but uppermost leaf ligulate, achene. SE Asia, Australia, New Zealand, New Caledonia
with well-developed, subterete, stiff blade; hypogynous 83. Lepidosperma
bristles 6, 2: achene, persistent on the rachilla; style un- - Hypogynous scales thin, membranous, persistent on the
usually long, 6-fid. S Africa 72. Neesenbeckia rachilla. Andean S America 17. Oreobolopsis
- Not as above 64 81. Hypogynous bristles deciduous with the achene; stamens
64. Upper internodes of rachilla elongated and prominently 6, filaments connate at the base; style 3-fid. S Africa
zig-zag 62. Schoenus 71. Cyathocoma
- Internodes short, or only uppermost internode elongated - Hypogynous bristles persistent on the rachilla; stamens 3;
65 style unusually long, 6-fid. S Africa 72. Neesenbeckia
65. Uppermost internode of rachilla elongated, thickened, 82. Inflorescence reduced to 1 spikelet 83
curved; hypogynous bristles poorly developed or absent. - Inflorescence paniculate, corymbose, anthelate, or
S Africa 73. Epischoenus capitate 91
- All internodes of rachilla short; hypogynous bristles 83. Leaves (almost) reduced to a sheath, or blade short 84
usually well developed 66 - Leaves with well-developed blade 86
66. Leaves conspicuously (spiro- )2-3-stichous, blade large 84. Inflorescence clearly pseudolateral, lowermost primary
and dorsiventrally fiat; inflorescence eprophyllate, bract erect and culmlike 24. Schoenoplectus
usually with numerous spikelets; glumes clearly - Inflorescence terminal 85
distichous; stamens 3; hypogynous bristles 6, usually > 85. Plants terrestrial, cespitose; all culms ::!:: similar
achene, ciliate to fimbriate 74. Costularia 19. Trichophorum
- Not as above 67 - Hydrophyte with submerged to floating branched stems;
67. Inflorescence eprophyllate, with many spikelets; glumes only a few robust branches bearing a spikelet. NE South
spirodistichous (to spirally) arranged, stamens 6; hypo- America 26. Egleria
gynous bristles 6, > achene, scabrid. S Africa 86. Leaves ligulate 87
71. Cyathocoma - Leaves eligulate 89
- Not as above 68 87. Hypogynous scales laminar, persistent on the rachilla.
68. Inflorescence prophyllate; stamens 3; hypogynous Andean S America 17. Oreobolopsis
bristles 6, short, sublaminar, often ciliolate, basally - Hypogynous bristles deciduous with the achene 88
connate into a disk; achene not beaked or with a small, 88. Hypogynous bristles 2:10, usually smooth and silky
short beak 84. Tricostularia 15. Eriophorum
 Cyperaceae 157

- Hypogynous bristles :56, usually scabrid and not silky - Achene laterally compressed 108
19. Trichophorum 108. Glumes deciduous, rachilla of spikelet usually persistent;
89. Small cushion-forming plant with basal, rosulate leaves. plants without curry smell 48. Pycreus
Andean S America 16. Phylloscirpus - Glumes persistent on a deciduous rachilla, spikelet de-
- Not as above 90 ciduous as a unit; pronounced curry smell
90. Primary bracts none, or glumelike; inflorescence reduced 49. Queenslandiella
to 1 spikelet, with blackish, glabrous glumes. N Zaire, 109. Spikelets with several empty glumes, only 1-2 (rarely
Uele 33. Nelmesia more) glumes each subtending a flower llO
- Primary bracts leaflike; inflorescence anthelate or capi- - Spikelets usually many-flowered, empty glume(s) rare,
tate, rarely reduced to 1 spikelet; glumes different or spikelets reduced to 1-2 glumes 120
22. Bolboschoenus 110. Inflorescence paniculate 111
91. Prophylls of spikes and spikelets axillating a bisexual - Inflorescence capitate 116
flower; inflorescence anthelate. Sumatra 111. Rachilla internodes elongated, conspicuously zig-zag
54. Sumatroscirpus 62. Schoenus
- Prophylls not axillating a flower 92 - Internodes short, or only upper internode elongated 112
92. Inflorescence paniculate 93 112. Inflorescence spadixlike, with spathelike bracts. Australia
- Inflorescence anthelate or capitate 94 77. Reedia
93. Culms terete; leaves and bracts reduced to a sheath; inflo- - Not as above 113
rescence with sinuous branches. S Africa 113. Uppermost rachilla internode elongated 114
21. Pseudoschoenus - Internodes short 115
- Culms 3-5-angular; leaves and bracts at least partly with 114. Inflorescence eprophyllate; glumes long persistent; lower
developed blade; inflorescence branches not sinuous flower male. S Africa 73. Epischoenus
20. Fuirena - Inflorescence prophyllate; glumes deciduous; upper
94. Inflorescence pseudoterminal, lowermost primary bract flower male. Hawaii, New Zealand 76. Morelotia
erect and culmlike; leaves usually reduced to a sheath, or 115. Leaves distichous, eligulate 82. Machaerina
blade short 24. Schoenoplectus - Leaves spirally arranged, often ligulate 70. Tetraria
- Inflorescence terminal, lowermost primary bract leaflike, 116. Culms densely hairy; blade much reduced; spikelets
spreading, more rarely erect 95 few-glumed, 1-flowered. Madagascar 58. Trichoschoenus
95. Hypogynous bristles ;:o::IO, much longer than the achene, - Not as above ll7
often silky 15. Eriophorum 117. Rhizome elongated; leaves eligulate; spikelets deciduous
- Hypogynous bristles :56, rarely silky 96 as a unit. S tropical Africa 44. Sphaerocyperus
96. Small cushion-forming plants with basal, rosulate leaves. - Rhizome short or plant tufted; leaves ligulate ll8
Andean S America 16. Phylloscirpus ll8. Leaves with a ciliate ligule, blade linear; inflorescence
- Not as above 97 spherical with numerous spikelets, spikelets 2-flowered.
97. All leaves basal and nearly reduced to a sheath; inflores- Australia 63. Gymnoschoenus
cence depauperate-anthelate. SE Asia - Ligule not ciliate, blade often reduced ll9
19. Trichophorum (T. subcapitatum) 119. Spikelets usually 2-more-flowered; rachilla internodes
- At least a few leaves with well-developed leaf blade, or if elongated and zig-zag; style base not distinct, not thick-
all leaves reduced to a sheath, then few-several leaves ened, deciduous or persistent, blade reduced or not
cauline 98 62. Schoen us
98. Leaves eligulate 22. Bolboschoenus - Spikelets !-flowered; rachilla internodes short; style base
- Leaves ligulate 99 distinct, thickened, deciduous; blade reduced
99. Inflorescence capitate 20. Fuirena 57. Actinoschoenus
- Inflorescence anthelate 100 120. Base of achene surrounded by a :±: cuplike, often 3-lobed
100. All leaves basal. SE Asia, NE Australia 23. Actinoscirpus disk; glumes with many parallel nerves. S Africa and E
- At least a few leaves cauline 14. Scirpus African mountains 34. Ficinia
101. Glumes (or glumelike scales) distichous, or number of - Not as above 121
glumes reduced to 1-2 102 121. Style base distinct, thickened, persistent or deciduous
- Glumes (or glumelike scales) spirally arranged 133 122
102. Style 2-branched 103 - Style base not distinct, not thickened, beaklike persistent
Style 3 (or more)-branched 109 124
103. Style base distinct, thickened, and persistent 122. Leaf sheath orifice with long hairs; style base often persis-
60. Rhynchospora tent, if deciduous, then achene usually transversely
- Not as above 104 rugose 30. Bulbostylis
104. Inflorescence of 1 to several sessile spikes, each with - Leaf sheath orifice without long hairs; style base
many bracts, subtending (highly) reduced spikelets with deciduous 123
1-few glumes 105 123. Achene >2mm, conspicuously clavate-stipitate; glumes
- Inflorescence anthelate or capitate, spikelets with many usually :±: coriaceous; inflorescence always with 1-few
distichous glumes 107 spikelets 31. Abildgaardia
105. Spikelet bract > > spikelet glume 52. Lipocarpha - Achene usually :51 mm, rarely stipitate; glumes often
- Spikelet bract<< spikelet glume 106 more delicate; inflorescence often with many spikelets
106. Glumes 2-more; achene laterally compressed 28. Fimbristylis
50. Kyllinga 124. Filaments conspicuously elongating after anthesis; per-
- Glumes 1; achene dorsiventrally compressed ennials on a seriate rhizome. Atlantic coast of S America
47. Ascolepis 40. Androtrichum
107. Achene dorsiventrally compressed 41. Cyperus - Not as above 125
158 Cyperaceae

125. Glumes 2 or more per spikelet 126 141. Inflorescence terminal, primary bracts spreading 142
- Spikelet reduced to a single glume (with or without the - Inflorescence pseudolateral, lowermost primary bract
spikelet prophyll), superposed to the spikelet bract 129 erect 144
126. Stoloniferous perennial; leaves many, cauline, thick; in- 142. Floating sedge with long stolons; glumes persistent,
florescence capitate; spikelets 3-4-glumed, upper rachilla apically thickened; achene corky thickened. Tropical
internode much enlarged, corky, clasping the achene. America and Africa 38. Oxycaryum
Pantropical, coastal 43. Remirea - Not as above 143
- Not as above 127 143. Inflorescence large, open anthelate. SE Asia
127. Dwarf annual; false spikelets with distichous bracts, each 14. Scirpus (S. ternatanus)
axillating a highly reduced 1-flowered spikelet with - Inflorescence small, capitate 52. Lipocarpha
prophyll and glume. Namibia, Zambia 51. Volkiella 144. Achene with a basal cuplike gynophore; plants perennial.
- Not as above 128 S Africa 34. Ficinia (F. latera/is)
128. Annual plants with curry smell; inflorescence anthelate - Not as above 145
with digitately arranged spikelets; spikelets deciduous 145. Glumes with many parallel nerves 36. Isolepis
as a unit; glumes with conspicuously winged keel. - Glumes 1-nerved, or with nerves converging towards the
Paleotropis 42. Courtoisina top 146
- Not as above 41. Cyperus 146. Glumes > 1.5 mm, usually much longer
129. Spikelet bract > > glume 130 24. Schoenoplectus
- Bract of spikelet < glume 131 - Glumelike scales (bracts of highly reduced spikelets)
130. Dwarf annual; bracts distichous. Namibia, Zambia small ( :s: 1mm) 52. Lipocarpha
51. Volkiella 147. Spikelets with several empty glumes, only 1-2 (rarely
- Annual or perennial plants; bracts spirally arranged more) glumes each axillating a flower 148
52. Lipocarpha - Spikelets usually many-flowered, empty glume (s) rare
131. Prophylls of spike and spikelets absent 47. Ascolepis 153
- Prophylls of spike and spikelets present 132 148. Inner side of glumes adpressed-pubescent near the top
132. Perennial; culm base bulbously thickened, basal leaf 149
sheaths semisucculent; glumes utriculiform, >4 mm - Not as above 150
long. S India 45. Ascopholis 149. Stamens 12-22; leaves basal or a few cauline. SW and W
- Annual plants; glumes <2mm long. Africa, Madagascar Australia 78. Evandra
46. Alinula - Stamens 3-6; leaves all cauline. Australia 79. Caustis
133. Style 2-branched 134 150. Inflorescence capitate; leaves reduced to a sheath. NE
- Style 3 (or more)-branched 147 Australia 56. Arthrostylis
134. Inflorescence paniculate 135 - Inflorescence paniculate; leaves with well-developed
- Inflorescence anthelate, capitate, or reduced to 1 spikelet blade 151
136 151. Inflorescence spadixlike, with partly enveloping
135. Style base not distinct, corky thickened, persistent; spike- spathelike primary bracts; stamens 6. SW and W
lets 2-3-flowered, lower flower male, upper flower(s) Australia 77. Reedia
bisexual; glumes long persistent 80. Cladium - Not as above 152
- Style base usually distinct and much thickened, persis- 152. Leaves eligulate, blade revolute; inflorescence
tent; spikelets 2-more-flowered, all flowers bisexual, or prophyllate; stamens 2(-3) 80. Cladium
upper flower(s) male; glumes usually deciduous - Leaves ligulate, blade involute; inflorescence
60. Rhynchospora eprophyllate; stamens (2- )3-6, filaments elongating after
136. Inflorescence anthelate; spikelets 1-flowered; glumes of anthesis, with different glume-fixing mechanisms
increasing length; stamens 2; style base thickened and 75. Gahnia
deciduous. E Australia 59. Trachystylis 153. Style base distinct, thickened, persistent, or deciduous
- Not as above 137 154
137. Glumes of increasing to subequallength; usually several - Style base not distinct, not thickened, often beaklike
glumes empty, 1-,2-few glumes subtending a flower; persistent 155
flowers bisexual, or upper flowers male; style base usually 154. Leaf sheath orifice with long hairs; style glabrous; style
distinct, thickened, persistent 60. Rhynchospora base often persistent, if deciduous, then achene usually
- Glumes subequal, usually each subtending a bisexual transversely rugose 30. Bulbostylis
flower 138 - Leaf sheath orifice without long hairs; style often
138. Style base distinct and thickened, persistent or deciduous fimbriolate; style base deciduous 28. Fimbristylis
139 155. Inflorescence pseudolateral, lowermost primary bract
- Style base not distinct, not thickened, sometimes beaklike erect and ± culmlike, long or short 156
persistent 140 - Inflorescence terminal, primary bracts ± spreading,
139. Style glabrous, style base persistent; leaf sheath orifice leaflike 158
with long hairs; basal spikelets often present. S Africa 156. Inflorescence often anthelate, branches flat-concave or
30. Bulbostylis (B. humilis) crescentiform in cross-section; if inflorescence capitate,
- Style usually fimbriolate, style base deciduous; leaf sheath then with many spikelets; glume nerves parallel; rhizome
orifice without long hairs 28. Fimbristylis thick, woody 37. Scirpoides
140. Glumes long persistent, dark reddish brown to black, ± - Not as above 157
scarious; achene blackish, conspicuously smooth and 157. Achene with a basal cuplike gynophore. S. and E Africa
shining. Africa 32. Nemum 34. Ficinia
- Glumes usually deciduous, if persistent, then achene - Not as above go to 145
different 141 158. Inflorescence paniculate 159
 Cyperaceae !59

- Inflorescence anthelate or capitate 161 flower. Rachilla apparently terminated by a flower.

!59. Inflorescence compacted, primary bracts partly adnate to Flowers bisexual. Terminal flower with a terminal
the axis; leaves stiff, yellowish. Coastal New Zealand
35. Desmoschoenus
pistil, surrounded by spirally arranged scales,
- Not as above 160 often each with a single stamen. Lateral flowers
160. Glumes with many parallel nerves, usually glabrous; dorsiventrally compressed, with a basal lateral
achene usually with a basal ± cuplike disk. S and E Africa pair of keeled and ::t: connate scales. Style 2-3-fid,
34. Ficinia style base not distinct, thickened, persistent. Fruit
- Glumes with few, apically converging nerves usually
pubescent to ciliate; achene without hypogynous disk, ovoid, with a large beak, ::t: drupaceous, ridged
often stipitate 20. Fuirena when dry. Two spp., Sri Lanka over SE Asia,
161. Inflorescence anthelate 162 Malesia, and NE Australia to Polynesia, in fresh-
- Inflorescence capitate or reduced to I spikelet( -like
structure) 165
162. Culms noded and leafy; inflorescence repeatedly
anthelate, with numerous spikelets. SE Asia
14. Scirpus (S. ternatanus)
- Culms scapose; inflorescence 1-2-anthelate with few to
many spikelets 163
163. Leaves equitant, blade subterete; rhizome elongated.
Africa 32. Nemum (N. equitans)
- Leaves spirally arranged, blade flat or reduced 164
164. Perennial; spikelets few, solitary. SE Asia
19. Trichophorum (T. subcapitatum)
- Annual plants; spikelets many, spikelike or capitately
compacted 41. Cyperus (C. hamulosus, C. michelianus)
165. Glumes with many parallel nerves; achene usually with a
basal ± cuplike disk. S Africa and E African mountains
34. Ficinia
- Not as above 166
166. Leaves ligulate, blade often reduced; inflorescence
reduced to 1 terminal spikelet 19. Trichophorum
- Leaves eligulate, blade usually well developed; inflores-
cence usually capitate 167
167. Glumes >7mm; achene >2mm, thick-walled. Coastal S
Africa 12. Hellmuthia
- Glumes much smaller; achene thin-walled 168
168. Small perennial plants; culm base covered by fibrous
remains of old leaf sheaths 39. Kyllingiella
- Annual plants; culm base different 169
169. Primary bracts long, leaflike; inflorescence of several
spikes, each with many crowded spikelets
41. Cyperus (C. hamulosus, C. michelianus)
- Primary bracts rather short; inflorescence of few false
spikelets (spikes!) with numerous false glumes (bracts of
reduced spikelets!) 52. Lipocarpha

I. Subfamily Mapanioideae C.B. Clarke (1897).

1. Tribe Hypolytreae Pres! ex Fenzl (1836).

1. Scirpodendron Zipp. ex Kurz Fig. 40

•
Scirpodendron Zipp. ex Kurz, J. Asiat. Soc. Bengal. 38, ii: 84 -
(1869); Kern, Fl. Males. I, 7: 456- 458 (1974), rev. .
.

Robust, tufted perennials. Culms (sub)scapose,

rather short, hidden between the basal leaves.
Leaves eligulate. Inflorescence densely paniculate
with numerous spikelets. Primary bracts leaflike,
not sheathing. Spikelets with several spirally
arranged persistent glumes, each subtending a
H
Fig. 40A-H. Cyperaceae. Scirpodendron ghaeri. A Habit. B
Portion of leaf. C Spikelet. D Lateral flower at anthesis. E
Same, diagram. F Infructescence. G Fruit. H Same, diagram-
matic cross-section. (Takhtajan 1982)
160 Cyperaceae

water tidal swamps and forests, also at the edge of Medium-sized perennial (base unknown). Culms
mangroves. probably 1-few-noded. Leaves eligulate (?). Inflo-
rescence open paniculate with many spikelets,
2. Exocarya Benth. branches semiverticillate. Primary bracts leaflike,
not (?) sheathing. Partial inflorescences corym-
Exocarya Benth., Hooker's Icon. Pl. 13: 5, t. 1206 (1877). bose, with 1-few spikelets at the top of their
branches. Spikelets with several spirally arranged
Creeping rhizomatous perennial. Culms many- persistent glumes, each subtending a flower.
noded, slender. Leaves eligulate, lower leaves of- Flowers bisexual. Scales 2 (?), lateral, laminar,
ten reduced to a sheath. Inflorescence anthelate completely united, slightly keeled. Stamens 3.
with many spikelets. Primary bracts leaflike. Style 3-fid, style base slightly thickened, persis-
Partial inflorescences anthelate, with solitary tent. Achene obovate, thick lenticular, 3-ribbed,
spikelets. Spikelets with many spirally arranged, beaked, rugose. Only one sp., P. grandis Uittien,
persistent glumes, lower glumes empty, small, Principe 1., W Africa. Known only from type
gradually passing into the larger fertile glumes. specimen, not seen by the author.
Flowers bisexual. Scales 4, basal pair lateral,
ciliately keeled, each with a single stamen, upper
2 scales posterio-anterior, not keeled, anterior 5. Mapania Aublet
scale with a stamen. Style deeply 2-fid, style base ±
Mapania Aublet, Hist. Pl. Guiane 1:47, t. 17 (1775); Simpson,
distinct, thickened, persistent. Achene ovoid to Rev. Mapania (1992).
ellipsoid, dorsiventrally compressed, style base Thoracostachyum Kurz (1869).
depressed-globose with a fine short beak, surface
smooth and shining black. Only one sp., E. Rhizomatous or stoloniferous perennials, roots
sclerioides (F. Muell.) Benth., NE Australia and SE thick, sometimes stiltlike on ascending rhizomes.
New Guinea (Fergusson Is.), in forest. Culms central or lateral, (sub)scapose. Lateral
culms few to many, with several basal leaves re-
3. Hypolytrum Rich. duced to a sheath. Leaves eligulate, sometimes
pseudopetiolate, rarely all leaves reduced to a
Hypolytrum Rich. in Pers., Syn. Pl. 1: 70 (1805); Nelmes, Kew
Bull. 10: 63-82 (1955) (Afr. spp.); Raynal, Adansonia II, 8:
sheath. Inflorescence rarely pseudolateral, capi-
424-430 (1968) (some Afr. spp.); Koyama, Darwiniana 16: tate, corymbiform, subanthelate or paniculate,
49-92 (1970) (Amer. spp.). when on lateral culms often reduced to 1-few
spikelets. Primary bracts small (on most lateral
Perennials, often on a thick woody rhizome. culms) or large and leaflike (on most central
Culms central or lateral (and then with a number culms). Spikelets with few to numerous spirally
of basal cataphylls), scapose or more rarely 1- arranged persistent glumes, each subtending a
several noded. Leaves eligulate, sometimes pseu- flower. Flowers bisexual. Scales 4-6, basal pair lat-
dopetiolate, blade sometimes lanceolate or eral, ± ciliate-keeled, each with a single stamen,
reduced (often in basal leaves). Inflorescence of the remaining 2-4 scales 0-2 with a stamen,
corymbose, paniculate or capitate, with numerous very rarely only 1 stamen per flower. Style 2-3-fid,
(-1) spikelets, often with 3( -more) collateral base not distinct, not or slightly thickened, persis-
branches per bract. Primary bracts small, scale- tent or not. Achene thick lenticular or rounded
like, or large and leaflike, patent to reflexed. Spike- trigonous, sometimes 3-ribbed, sometimes with a
lets with many spirally arranged persistent fleshy exocarp. About 70 spp., pantropical, mainly
glumes, each subtending a flower. Flowers bi- equatorial; in tropical rainforest and along open
sexual. Scales 2( -3 ), basal pair lateral, ± ciliate- swamps.
keeled, each with a single stamen, scales often
connate or ± fused. Style bifid, style base ±
persistent. Achene (ob)ovate to elliptic, dorsiven- 6. Mapaniopsis C.B. Clarke
trally biconvex, smooth, costate, wrinkled, tuber- Mapaniopsis C.B. Clarke, Kew Bull., Addit. Ser. 8: 55 (1908);
culate, or spongious. About 40 species, Koyama, Mem. N.Y. Bot. Gard. 17: 48-49 (1967).
pantropical, mainly equatorial, in forest.
Medium-sized to tall (tufted or rhizomatous?)
perennials. Culms few-many-noded. Leaves
4. Principina Uittien
eligulate. Inflorescence anthelate with numerous
Principina Uittien, Recueil Trav. Bot. Neerl. 32: 282 (1935). spikelets. Lower primary bracts large, leaflike.
 Cyperaceae
Partial inflorescences also anthelate, ultimate
branches with few-many digitately arranged
spikelets. Spikelets with several-many spirally
arranged persistent glumes, each subtending a
flower. Flowers bisexual. Scales 4-5, basal pair
lateral, each with a stamen, 1 median posterior
scale empty, 1-2 median or sublateral anterior
scales, each with a stamen. Style 2-fid, style base
not distinct, somewhat thickened, persistent,
smooth. Achene ovate, dorsiventrally (thick)
lenticular, beaked, surface smooth. Two spp., SE
Venezuela, French Guyana, and adjacent N Brazil,
in forest.
7. Paramapania Uittien
Paramapania Uittien, Recueil Trav. Bot. Neerl. 32: 186 (1935);
Kern, Fl. Males. I, 7: 484-489 (1974).
Rhizomatous perennials. Culms lateral, in the axil
of the lower leaves, or below the leaves, with some
basal bladeless sheaths, sometimes with 1-3
cauline sheaths also. Leaves basal, sometimes
pseudopetiolate, eligulate, blade rarely narrowly
oblong. Inflorescence anthelate or capitate with
few (-1) spikelets. Primary bracts very short.
Spikelets with many densely spirally arranged,
persistent glumes, upper glumes subtending a
flower. Flowers bisexual. Scales usually 5, basal
pair lateral, keeled, both subtending a stamen,
anterior scale absent, rarely present, with a sta-
men, posterior and 2 anterio-lateral scales empty.
Style 3( -4)-fid, style base not distinct, slightly
thickened, persistent. Achene ± obovate, sub-
terete to trigonous, beaked, surface smooth. Seven
spp., Malesia, P. radians (C.B. Clarke) Uittien
extending towards the W Pacific islands, in
rainforest.
8. Diplasia Rich.
Diplasia Rich. in Pers., Syn. Pl. 1: 70 (1805); Koyama, Mem.
N.Y. Bot. Gard. 17: 47-48 (1967).
Tall perennial with thick creeping rhizomes.
Culms few-noded, thick, robust. Leaves eligulate,
coriaceous. Inflorescence a large anthela of an-
thelate to ± corymbiform partial inflorescences
with many spikelets. Primary bracts large, leaflike,
not sheathing. Spikelets with many spirally ar-
ranged coriaceous persistent glumes, each sub-
tending a flower. Flowers bisexual. Scales 4, basal
pair lateral, spinulose-keeled, empty, 2 median
scales enclosing ca. 8 stamens. Style 2-fid, base not
distinct, scarcely thickened, persistent. Achene
161
ovate, dorsiventrally lenticular, smooth, shining
black, with a fine small beak. Only one sp., D.
karataefolia Rich., Costa Rica to N Peru and N
Brazil, in rainforest.
9. Capitularina Kern
Capitularina Kern, Fl. Males. I, 7: 458 (1974).
Shortly rhizomatous perennial. Culms scapose,
(4-)5-angular, septate. Leaves eligulate, blade pli-
cate, or leaves reduced to a sheath. Inflorescence
capitate with 1-many spikelets. Primary bracts
5-8, leaflike, suberect to spreading. Spikelets
with many densely spirally arranged deciduous
glumes, each subtending a flower. Flowers bi-
sexual. Scales many, membranous, basal pair lat-
eral, spathulate, cucullate, keeled, empty, the next
2 also empty, decussate, other scales ca. 10, each
subtending a stamen. Style 2-fid, style base thick-
ened, persistent. Achene obovate, dorsiventrally
(thick) lenticular, many-ribbed, on a hollow stipe,
beak thick and corky. Only one sp., the poly-
morphous C. involucrata (Valcken.) Kern, New
Guinea and the Solomon Is., in swampy forest.
2. Tribe Chrysitricheae Lestib. ex Fenzl (1836).
10. Chorizandra R. Br.
Chorizandra R. Br., Prodr.: 221 (1810).
Creeping rhizomatous perennials. Culms scapose,
mostly septate. Leaves reduced to a very oblique-
mouthed sheath. Inflorescence pseudolateral, re-
duced to a single spikelet, ± partly sunken into the
± widened base of the single, erect, stemlike bract.
Spikelet globose or ± compressed (from top to
base!), with many spirally arranged deciduous
glumes, each (except a few lower ones) subtend-
ing a flower. Rachilla extremely shortened, thick
crescentiform. Flowers bisexual. Scales many,
basal pair lateral, spathulate, ± keeled, each with
a single stamen, many middle scales smaller and
narrower, each with a stamen, several upper
empty. Style deeply 2-3-fid, style base not distinct,
slightly thickened, persistent. Achene thick abo-
void to lenticular, ± beaked, longitudinally ribbed
or reticulate. Seven spp., Australia and New
Caledonia, in swampy, semidry regions.
11. Lepironia Rich.
Lepironia Rich. in Pers., Syn. Pl. 1: 70 (1805); Kern, Fl. Males.
I, 7: 460-463 (1974).
162
Rhizomatous perennial with thick roots. Culms
scapose, septate. Leaves reduced to a sheath, open
in front, the margins overlapping. Inflorescence
pseudolateral, of 1 single spikelet. One empty
bract large, culmlike, erect. Spikelet with many
densely spirally arranged deciduous glumes, lower
empty, upper each subtending a flower. Rachilla
thick, spongy. Flowers bisexual. Scales many,
basal pair lateral, each subtending a single stamen,
the upper scales (up to 15) smaller, flat, many
subtending a stamen. Style 2-fid, style base not
distinct, slightly thickened, persistent. Achene
obovate, dorsiventrally compressed, plano-
convex, ± winged along the margins, beaked.
Only one sp., L. articulata (Retz.) Domin,
Madagascar over Sri Lanka, SE Asia, Malesia, NE
Australia towards Polynesia, in swampy places
mostly at low altitudes.
12. Hellmuthia Steudel
Hellmuthia Steudel, Syn. Pl. Glumac. 2: 90 (1855); Haines &
Lye, Bot. Not. 129: 61-67 (1976).
Medium-sized tufted perennial. Culms (sub)-
scapose. Leaves eligulate, lower leaves often
reduced to a sheath. Inflorescence capitate, with
(1- )2-10 spikelets. Lower primary bracts leaflike.
Spikelets with many densely spirally arranged
deciduous glumes, each subtending a flower
(sometimes a few lower glumes empty). Flowers
bisexual. Scales 2, transversal, keeled, each with
a stamen, 3rd stamen median anterior (scales
lacking in the upper flowers). Style deeply 3-fid,
style base not distinct, not thickened, deciduous.
Achene broadly obovate, stipitate, compressed
trigonous, surface minutely reticulate. Only one
sp., H. membranacea (Thunb.) R. Haines & Lye,
South Africa (coastal Cape), on coastal sands.
The taxonomic position of this genus is contro-
versial, and Bruhl (1995) tends to exclude it from
the Mapanioideae. In this respect, the morpho-
logical interpretation of the 2 floral scales is most
important.
13. Chrysitrix L.
Chrysitrix L., Mant. Pl. 2: 165, 304 (1771).
Tufted or creeping rhizomatous perennials with
thick roots. Culms scapose, ancipitous to terete.
Leaves distichous, eligulate, blade ancipitous or
terete, or reduced to a sheath. Inflorescence
pseudolateral, reduced to a single spikelet, partly
hidden by the base of a ± culmlike bract. Spikelet
with several spirally arranged empty, ± decidu-
Cyperaceae
ous, dark glumes, seated on a broadly cylindrical
rachilla, conically narrowed at the top, and there
bearing a single bisexual flower with many spirally
arranged lanceolate to linear scales, each with a
single stamen, top with a central pistil, style deeply
4-6-fid, style base not distinct, not thickened,
deciduous. Stamens with a short filament, a long
linear anther, and a very elongated linear crest;
rarely a second, lateral flower present in the axil of
the first glume; sometimes the apex of the conical
part of the rachilla divided into 3 smaller cones
each bearing a number of scales and a stamen,
plus a terminal pistil with a 2-fid style. Achene
stipitate, thick ovoid-ellipsoid to dorsiventrally
compressed, longitudinally ribbed. About four
spp., S Africa (SW Cape) and SW Australia in
damp places on slopes and flats.
The spikelet structure of these species is highly
unusual and is much in need of a more detailed
comparative study. The rare presence of a lateral
flower with the typically mapanioid arrangement
suggests the inclusion in the Mapanioideae.
II. Subfamily Cyperoideae Suessenguth (1939).
2. Tribe Scirpeae Kunth ex Dumort. (1827).
14. Scirpus L.
Scirpus L., Sp. Pl. 1: 47 (1753); Schuyler, Proc. Acad. Nat. Sci.
Philad. 119:295-323 (1967) (rev. N Amer.); Wilson, Telopea
2: 153-172 (1981) (rev. Austr.); Wilson, Taxon 38: 316-320
( 1989) (typification).
Trichophorum auct. plur., non Pers. (cf. Salmenkallio &
Kukkonen 1989).
Tufted or rhizomatous perennials. Culms few-
many-noded. Leaves ligulate. Inflorescence
anthelate with many to numerous spikelets. Pri-
mary bracts leaflike, spreading. Partial inflores-
cences anthelate to capitate. Spikelets with many
spirally arranged, deciduous glumes, each sub-
tending a flower. Flowers bisexual. Bristles (0-)3-
6, as long as to much longer than the fruit (and
then often extremely contorted), ± scabrid, de-
ciduous with the fruit. Stamens 1-3. Style (2-)
3-fid, style base not distinct, not thickened, persis-
tent. Achene (ob )ovate to elliptic, (compressed)
trigonous to dorsiventrally lenticular, beaked,
smooth. About 20 spp., Holarctis, SE and E Asia,
Malesia (2-0), SE Australia (1-1), and Andean
South America (1-1), in bogs, marshes, wet grass-
land, more rarely in wet woodland.
In Scirpus s.l. previously all scirpoid-looking
species without any clear diagnostic characters
have been dumped. They are now assigned to
more than 50 other genera. Several critical species
 Cyperaceae 163

have not yet been satisfactorily allocated. Also the

separation between Scirpus and Eriophorum is still
unsatisfactory and is here based on the number
of hypogynous scales (6 or less = Scirpus, 10 or
more = Eriophorum).

15. Eriophorum L.
Eriophorum L., Sp. Pl. 1:52 (1753); Palla, Bot. Zeit. 54: 141-158
(1896).
Erioscirpus Palla ( 1896).

Tufted perennials, sometimes rhizomatous,

stoloniferous, or tussock-forming. Culms scapose
or few-noded. Leaves ligulate, blade usually re-
duced in the cauline leaves. Inflorescence (com-
pound-)anthelate or reduced to a single spikelet,
fruiting spikelets sometimes drooping. Primary
bracts mostly small, rarely leaflike. Spikelets with
few to many spirally arranged deciduous glumes,
each subtending a flower. Flowers bisexual.
Bristles 10-many, extremely elongating after an-
thesis, silky, deciduous with the fruit. Stamens
1-3. Style 3-fid, style base not distinct, not
or slightly thickened, ::t persistent. Achene
obovate to oblong, trigonous, beak short to long,
surface smooth. About 20 spp., scattered over the
holarctic region, most species growing in wet
moorland and dominating some arctic vegetation
types.
Erioscirpus, with its remarkable species in
the Himalayas, might well deserve generic status,
as suggested by its rather deviating embryo
shape.

16. Phylloscirpus C.B. Clarke Fig. 41J

Phylloscirpus C.B. Clarke, Kew Bull., Addit. Ser. 8: 44 (1908);
Goetghebeur & Simpson, Kew Bull. 46: 174-175 (1991).

Small perennials on slender horizontal or vertical

rhizomes. Culms (sub)scapose, often very short-
ened. Leaves eligulate, blade canaliculate, up-
wardly subensiform. Inflorescence capitate with
up to 20 spikelets. Primary bracts short, not
sheathing, suberect. Spikelets with few-several
spirally arranged, (finally) deciduous glumes,
each subtending a flower. Flowers bisexual.
Bristles (3-)6, retrorsely scab rid, as long as or
longer than the fruit, deciduous with the fruit.
Stamens 3. Style 3-fid, style base not distinct, not
or slightly thickened, persistent. Achene obovate,
(compressed) trigonous, beaked, smooth, mi-
nutely reticulate-foveolate. About five spp.,
Andean S America, in marshy puna grassland and
bogs at high altitude.
Fig. 41A-J. Cyperaceae. A-C Websteria confervoides. A Habit.
B Spikelet. C Fruit, surrounded 3 filaments and many hypogy-
nous bristles. D- G Schoenoplectus lateriflorus. D Habit of cae-
spitose plant with pseudolateral heads. E Fruit. F Longitudinal
section of shoot basis showing a basal flower. G Achene. H, I
Isolepis fluitans. H Shoot. I Achene. J Phylloscirpus acaulis.
Rhizome with leaf rosette. (Takhtajan 1982)
17. Oreobolopsis T. Koyama & Guaglian.
Oreobolopsis T. Koyama & Guaglian., Darwiniana 28: 79
(1987).
Small, tufted perennial. Culms (sub)scapose.
Leaves ligulate, blade upwardly subensiform. In-
164
florescence sometimes + pseudolateral, reduced
to a single spikelet. Primary bracts 2, empty, lower
leaflike, suberect, upper reduced, not sheathing.
Spikelet with few, spirally arranged, deciduous
glumes, 1-2lower glumes empty, middle and up-
per glumes each subtending a flower. Flowers bi-
sexual. Scales 6, laminar, irregularly lobed at the
top, persistent on the rachilla. Stamens 3. Style
deeply 3-fid, style base not distinct, not thickened,
deciduous. Achene obovate, compressed trigo-
nous, slightly 3-ribbed, surface smooth. Only one
sp., 0. tepalifera T. Koyama & Guaglian., Bolivia
and Ecuador, in cushion-paramo or puna around
4000m.
The Californian Scirpus clementis Jones is some-
times included in (Scirpus section) Trichophorum
(Baeothryon auct.). However, the 6 hypogynous
scales are quite similar to those of Oreobolopsis
(laminar, pale brownish, with irregularly lobed
top) and are persistent on the rachilla. Therefore
this species might be included as a second species
in Oreobolopsis.
18. Amphiscirpus Oteng-Yeb.
Amphiscirpus Oteng-Yeb., Notes R. Bot. Gard. Edinburgh 33:
308 (1974); Goetghebeur & Simpson, Kew Bull. 46: 175-177
(1991).
Small perennial from a deep rhizome. Culms sea-
pose, subterete to slightly compressed, halfway
often with a V-formed ridge. Leaves usually stiffly
erect, ligulate, blade often with a similar ridge,
upwardly subensiform. Inflorescence :±:: pseu-
dolateral, (sub)capitate with 1-few spikelets. Pri-
mary bracts small, not sheathing, lowermost bract
larger and suberect. Spikelets with many spirally
arranged deciduous glumes, each subtending a
flower. Flowers bisexual. Bristles 3-6, smaller
than the fruit, retrorsely scabrid, deciduous
with the fruit. Stamens 3. Style 2-fid, style base
not distinct, not thickened, deciduous. Achene
obovate, dorsiventrally plano-convex to lenticu-
lar, smooth, and shining, minutely reticulate.
Only one sp., A. nevadensis (S. Watson) Oteng-
Yeb., W Canada, USA, Andean Chile, and Argen-
tina, on sandy, semisaline soil, often in dried-up
lake beds.
19. Trichophorum Pers.
Trichophorum Pers., Syn. Pl. 1: 69 (1805); Beetle, Am. J. Bot.
28: 469-476 (1941); Salmenkallio & Kukkonen, Taxon 38:
313-316 (1989) (typification).
Baeothryon auct. plur., non A. Dietr.
Eriophorella Holub (1984).
Cyperaceae
Tufted or creeping rhizomatous perennials.
Culms scapose. Leaves ligulate, blade much
reduced, rarely well developed, upwards subensi-
form or top conspicuously thickened. Inflores-
cence a single spikelet, rarely anthelate or capi-
tate with few spikelets. Primary bracts small,
lowermost bract suberect. Spikelets with several
spirally arranged deciduous glumes, each sub-
tending a flower. Flowers bisexual. Bristles 0-6,
elongating after anthesis, silky or not, deciduous
with the fruit. Stamens 3. Style 3-fid, style base not
distinct, slightly thickened, :±:: persistent. Achene
obovate, trigonous or dorsiventrally compressed,
sometimes :±:: 3-ribbed, slightly beaked, smooth.
About ten spp., holarctic, mountains of tropical
Asia, and Andean S America, often in alpine or
tundra vegetation or acid bogs, rarely in wood-
land.
The Andean Scirpus rigidus Boeckeler and
its relatives probably are to be included in this
genus, although their flowers lack the hypogynous
scales.
4. Tribe Fuireneae Reichenb. ex Fenzl (1836).
20. Fuirena Rottb.
Fuirena Rottb., Descr. Icon. Rar. Pl.: 70, t. 19 (1773); Kral, Sida
7: 309-354 (1978) (N & C Amer. spp.); Haines & Lye, Sedges
& Rushes E. Afr.: 41-52 (1983) (E Afr. spp).
Small to medium-sized annuals or rhizomatous
perennials. Culms usually many-noded (scapose
in F. enodis C.B. Clarke), 3-5-angular, occasion-
ally thickened at the base. Leaves tri- or pen-
tastichous, ligule tubular, membranous, blade
often reduced in the lower leaves. Inflorescence
paniculate to capitate with few to many spikelets,
rarely pseudolateral. Primary bracts often leaflike,
sheathing, lowermost bract sometimes erect.
Spikelets with many spiral, rarely (spiro)penta-
stichous, deciduous glumes, each subtending a
flower. Flowers bisexual. Bristles (0-1- )3, or 6,
and then in 2 series, the rib-opposed bristlelike,
the face-opposed often differentiated into vari-
ously shaped scales, rarely all bristles reduced or
absent, or only 1 scale developed, as long as or
shorter than the fruit, deciduous with the fruit.
Stamens 1-3. Style 3-fid, base not distinct, not or
slightly thickened, persistent. Achene obovate,
triquetrous to trigonous, often stipitate, beaked,
smooth, or with various ornamentations. About
30 spp., all tropical and warm temperate regions,
concentrated in America and Africa, mainly in
open, humid localities at low altitude.
 Cyperaceae
21. Pseudoschoenus (C.B. Clarke) Oteng-Yeb.
Pseudoschoenus (C.B. Clarke) Oteng-Yeb., Notes R. Bot. Gard.
Edinburgh 33: 308 (1974).
Medium-sized perennial on a thick, short, woody
rhizome. Culms scapose. Leaves reduced to a mu-
cronate sheath, eligulate. Inflorescence ± pseu-
dolateral, compound paniculate with many
spikelets, axis conspicuously sinuous. Primary
bracts short and rigid, sheathing, erect. Spikelets
with many spirally arranged, (finally) deciduous
glumes, each subtending a flower. Flowers bi-
sexual or polygamo-dioecious. Bristles 3-5( -6),
retrorsely scabrid, as long as or longer than the
fruit, deciduous with the fruit. Stamens 3 (reduced
in female flowers). Style 3-fid, style base ± dis-
tinct, somewhat thickened, persistent. Achene
narrowly obovate, plano-convex to trigonous,
slightly beaked, surface smooth. Only one sp., P.
inanis (Thunberg) Oteng-Yeboah, endemic in S
Africa, NE Cape, growing along stream sides at
higher altitudes.
22. Bolboschoenus (Asch.) Palla
Bolboschoenus (Asch.) Palla in Hallier & Brand, Koch's Syn.
Deut. Schweiz. Fl., ed. 3, 3: 2531 (1905); Wilson, Telopea 2:
156-157 (1981) (Austr. spp.); Browning et al., Brittonia 47:
433-445 (1995) (N. Amer. spp.).
Rhizomatous perennial, rhizome often forming
hard ovoid tubers. Culms many-noded, bulbously
thickened at the base. Leaves eligulate, blade often
reduced in the lower leaves. Inflorescence some-
times pseudolateral, (compound) anthelate or
capitate with 1-many spikelets. Primary bracts
leaflike, patent, lowermost one often suberect.
Spikelets with many spirally arranged, deciduous
glumes, each subtending a flower. Flowers bi-
sexual. Bristles 3-6, shorter to longer than the
fruit, deciduous with the fruit. Stamens 3. Style 2-
3-fid, style base not distinct, slightly or not thick-
ened, persistent. Achene obovate, dorsiventrally
lenticular or (compressed) trigonous, beaked,
smooth. Only one subcosmopolitan sp., B.
maritimus (L.) Palla, and probably ten or more
spp., in more restricted areas, often in saline or
brackish habitats.
Specific limits are less than clear and a world-
wide study is badly needed. Some authors do
not recognize this genus, and its species are
sometimes included in Schoenoplectus s.l.
(Haines and Lye 1983, Strong, Novon 3: 202-203.
1993).
165
23. Actinoscirpus (Ohwi) R. Haines & Lye
Actinoscirpus (Ohwi) R. Haines & Lye, Bot. Not. 124: 481
(1971); Kern, Fl. Males. I, 7: 497-499 (1974); Goetghebeur &
Simpson, Kew Bull. 46: 169-172 (1991).
Hymenochaeta P. Beauv. ex Lestib. (1819), nom. rej.
Medium-sized to tall perennial, with rather long,
slender, scaly stolons ending in a small tuber.
Culms scapose, thickened at the base, spongy and
inconspicuously septate. Leaves ligulate. Inflores-
cence anthelate with many spikelets. Primary
bracts leaflike, not sheathing, patent to reflexed.
Spikelets with many spirally arranged, deciduous
glumes, each subtending a flower. Flowers bi-
sexual. Bristles S-6, retrorsely scabrid, mostly
longer than the fruit, deciduous with the fruit. Sta-
mens 3. Style deeply 3-fid, style base not distinct,
not thickened, persistent. Achene obovate, com-
pressed trigonous, beaked, surface smooth.
Monotypic, A. grossus (L. f.) Goetghebeur & D.A.
Simpson, India to NE Australia, often abundant in
swampy areas at low altitudes.
24. Schoenoplectus (Reichb.) Palla Fig. 41D-G
Schoenoplectus (Reichb.) Palla, Verh. K.K. Zool.-Bot. Ges.
Wien 38, Sitzungsber.: 49 (1888), nom. cons.; Raynal,
Adansonia II, 16: 119-155 (1976) (rev. sect. Supini); Wilson,
Telopea 2: 157-161 (1981) (Austr. spp.); Haines & Lye,
Sedges & Rushes E. Afr.: 52-64 (1983) (E Afr. spp.).
Annuals or perennials, tufted or creeping. Culms
(sub )scapose, rarely septate. Leaves usually re-
duced to a sheath, more rarely developing a ligulate
blade. Female basal flowers present in axil of
1 or more leaves in most species of sect. Supini
[bisexual basal flowers only known from S.
aberrans (Chermezon) Raynal]. Inflorescence
pseudolateral, anthelate or more frequently capi-
tate with (1-)few-many spikelets. Lowermost pri-
mary bract often large, erect, stemlike. Spikelets
with many spiral, rarely pseudodistichous, decidu-
ous, rarely persistent glumes, often each subtend-
ing a flower. Flowers bisexual [partly male in S.
perrieri (Chermezon) Raynal]. Bristles 0-6, shorter
or longer than the fruit, smooth, scabrid or plu-
mose, deciduous with the fruit. Stamens 1-3. Style
2-3-fid, style base not distinct, not thickened, per-
sistent. Achene obovate, ± trigonous or dorsiven-
trally lenticular, beaked or not, smooth to rugo~e.
About 50 spp., widely distributed on all continents.
The perennial species in deep to shallow water, the
annual ones in seasonally wet depressions.
Various subdivisions were proposed, mainly
based on differences in habit (survey in Wilson
1981: 158-159), yet none of these is satisfactory.
166
5. Tribe Eleocharideae Goetghebeur (1985).
25. Eleocharis R. Br.
Eleocharis R. Br., Prodr.: 224 (1810); Svenson, Rhodora 31-41
(1929-1939); Blake, Proc. R. Soc. Queensland 50: 88-132
(1939) (Austr. spp.); Haines & Lye, Sedges & Rushes E Afr.:
64-76 (1983) (E Afr. spp.); Kukkonen, Ann. Bot. Fenn. 27:
109-117 (1990) (infragen. class.).
Baeothryon A. Dietr. (1833) (cf. Salmenkallio & Kukkonen
1989).
Chillania Roiv. (1933 ).
Chamaegyne Suesseng. (1943).
Helonema Suesseng. (1943).
Heleocharis, orthogr. var.
Small to medium-sized tufted annuals or rhi-
zomatous, stoloniferous, rarely bulbiferous peren-
nials, rarely amphicarpous. Culms scapose,
3-4-angular, ridged or terete, rarely septate,
ancipitous or bulbously thickened at the base.
Leaves 1-few per culm, eligulate, reduced to a ±
hyaline sheath or blade present as a small mucro.
Inflorescence a single spikelet. Spikelets with 2-
many spirally arranged, rarely (sub )distichous,
deciduous glumes, each subtending a flower. Low-
ermost glume ("associated bract") often empty or
with a vegetative bud (often developing into new
shoots when submerged), rarely flower-bearing.
Flowers bisexual. Bristles (0-)3-6( -10 ), shorter or
longer than the fruit, deciduous with the fruit. Sta-
mens 1-3. Style 2-3-fid, style base mostly distinct,
(sometimes only slightly) thickened, persistent.
Achene mostly obovate, lenticular or (com-
pressed) trigonous, beaked, smooth or variously
ornamented. About 200 spp., cosmopolitan, with
a marked concentration in (sub )tropical America;
nearly all (partly) submerged or on temporarily
wet or inundated muddy soil.
26. Egleria L. Eiten
Egleria L. Eiten, Phytologia 9: 481 (1964).
Submerged stoloniferous perennial(?), with float-
ing to partly emerged culms. Culms many-noded,
with false whorls of tiny vegetative branchlets, at
the base of a tubular leaf sheath (this "culm" prob-
ably representing a sympodial, elongated rhi-
zome). Leaves a tiny, hyaline sheath, mouth often
somewhat lobed. Inflorescence a single spikelet on
a thick peduncle basally surrounded by a tubular
leaf sheath. Spikelets with a basally sheathing and
flower-bearing "associated bract" and many spiral
deciduous (?) glumes, each subtending a flower.
Flowers bisexual. Bristles 5, scabrid, longer than
the fruit, deciduous with the fruit. Stamens 3, an-
Cyperaceae
thers with basal spongy appendages (Bruhl 1995:
173). Style 3-fid, style base slightly constricted,
persistent. Achene obovate, plano-convex, with a
narrow beak, smooth with a very fine cancellate
pattern. Only one sp., E. fluctuans L. Eiten, S
Venezuela and N Brazil.
27. Websteria S.H. Wright Fig. 41A-C
Websteria S.H. Wright, Bull. Torrey Bot. Club 14: 135 (1887);
Eiten, Ann. Mo. Bot. Gard. 63: 181-198 (1976).
Submerged to floating perennial. Culms many-
noded, much branched, with numerous vegetative
branchlets (with naked top) in successive false
whorls, intermingled with rare, more robust,
generative culms. Leaves a tubular, hyaline sheath.
Inflorescence a single spikelet on top of a genera-
tive culm. Spikelets with 2 distichous, deciduous,
much elongated glumes, the upper subtending a
flower. Flowers bisexual. Bristles (3- )6-11, some-
what longer than fruit, deciduous with the fruit.
Stamens 3. Style 2-fid, style base not distinct,
slightly thickened, persistent. Achene obovate,
dorsiventrally lenticular, minutely reticulate. Only
one sp., W. confervoides (Poiret) Hooper, scat-
tered occurrence in the tropics, locally extending
into subtropical regions.
6. Tribe Abildgaardieae Lye (1973).
28. Fimbristylis Vahl
Fimbristylis Vahl, Enum. Pl. 2: 285 (1805), nom. cons.; Kral,
Sida 4: 57-70, 101-136 (1971) (N & C Amer. spp); Kern, Fl.
Males. I, 7: 540-592 (1974); Haines & Lye, Sedges & Rushes
E. Afr.: 77-91 (1983); Goetghebeur & Coudijzer, Bull. Jard.
Bot. Belg. 54: 65-89 (1984) (C Afr. spp.); Latz, Nuytsia 7:
161-182 (1990) (Austr. spp.).
Tylocarya Nelmes (1949).
Annuals or tufted, more rarely creeping rhizo-
matous perennials. Culms (sub)scapose, terete
to 3-4-5-angular or ancipitous. Leaves rarely
(spiro )distichous or equitant, ligule absent or
present as a membranous or hairy rim, blade
sometimes ensiform or reduced to a sheath. Inflo-
rescence rarely ± pseudolateral, anthelate with
few to many spikelets, sometimes reduced to 1
single spikelet, more rarely capitate. Primary
bracts leaflike or short, lowermost bract rarely
erect. Spikelets with few to many spirally arranged,
rarely distichous, deciduous glumes, each sub-
tending a flower. Flowers bisexual. Bristles absent.
Stamens 1-3. Style 2-3-fid, style base distinct,
thickened, deciduous. Achene often obovate,
 Cyperaceae
trigonous, dorsiventrally lenticular, or rarely
subcylindrical, often with conspicuous ornamen-
tation. About 300 species, pan tropical to (warm-)
temperate, with a heavy concentration in SE Asia,
Malesia, and NE Australia; the annuals often on
temporarily wet sand, the perennials preferring
permanently wet conditions; several species on
disturbed grounds and often becoming weedy.
A natural classification has not been attained,
but a :±: convenient system of 18 sections has been
suggested by Kern (1974: 543), although Abild-
gaardia and Actinoschoenus here are raised to
generic rank.
29. Crosslandia W. Fitzg.
Crosslandia W. Fitzg., J. Proc. R. Soc. Western Australia 3: 122
(1918).
Tufted annual (or short-lived perennial?). Culms
scapose. Basal leaves with open sheath, eligulate,
subtending the basal spikelets, subbasal leaves
few, reduced to a mucronate sheath with a con-
spicuous scarious mouth. Basal spikelets abun-
dant, with few-many (spiro)distichous :±: leaflike
glumes, flowers female, with elongated, 3-fid
style. Inflorescence anthelate or capitate with few-
numerous spikelets. Primary bracts short, spread-
ing, lowermost bract suberect. Spikelets with
many densely (spiro )distichous, deciduous
glumes, each subtending a flower. Flowers uni-
sexual, lower glumes with male flowers and upper
glumes with female flowers. Bristles absent. Sta-
mens 2-3. Style 3-fid, base distinct, somewhat
thickened, deciduous. Achene obovate, rounded
trigonous, surface tuberculate. Only one sp., the
variable C. setifolia W. Fitzg., endemic in N
Australia, on dry sandy places, often in woodland.
30. Bulbostylis Kunth
Bulbostylis Kunth, Enum. Pl. 2: 205 (1837), nom. cons; Kral,
Sida 4: 57-70, 73-101 (1971) (N & C Amer. spp.); Haines &
Lye, Sedges & Rushes E. Afr.: 91-130 (1983); Goetghebeur &
Coudijzer, Bull. Jard. Bot. Belg. 55: 207-259 (1985) (C Afr.
spp.).
Annuals or tufted perennials, rarely with an elon-
gated rhizome, rarely forming a caudex. Culms
scapose, rarely many-noded. Leaves eligulate, but
with 2lateral tufts oflong white hairs at the sheath
mouth, rarely reduced to a sheath. Basal flowers
rarely present. Inflorescence anthelate or capitate
with few-many spikelets, or reduced to a single
spikelet, rarely pseudolateral. Primary bracts
167
short, rarely the lowermost bract leaflike and
erect. Spikelets often with many densely spirally
arranged (rarely distichous), deciduous glumes,
each subtending a flower. Flowers bisexual.
Bristles absent. Stamens 1-3. Style 3( -2)-fid, style
base distinct, thickened, persistent, rarely only
slightly thickened or deciduous. Achene obovate
to obpyriform, rounded trigonous, rarely dor-
siventrally lenticular, variously ornamented,
rarely smooth. About 100 spp., pantropical to
warm temperate, with concentrations in tropical
Africa and S America; most species in open, sunny
habitats on dry or temporarily wet soils.
31. Abildgaardia Vahl
Abildgaardia Vahl, Enum. Pl. 2: 296 (1805); Gordon-Gray, J.S.
Afr. Bot. 32: 141-152 (1966) (S Afr. spp.); Kral, Sida 4:57-73
(1971) (N & C Amer. spp.).
Annuals, or tufted to rhizomatous perennials.
Culms scapose. Leaves eligulate, sometimes re-
duced to a sheath. Inflorescence depauperate
anthelate or capitate with few spikelets, or reduced
to one single spikelet. Primary bracts short, not
sheathing. Spikelets with many densely (spiro-)
distichous, deciduous glumes, each subtending a
flower. Flowers bisexual. Bristles absent. Stamens
2-3. Style deeply 3-fid, style base distinct, thick-
ened, deciduous. Achene mostly obovate, stipi-
tate, rounded trigonous, rarely winged, surface
often tuberculate. About ten spp., tropics and
subtropics, concentrated in Australia, in dry, dis-
turbed grassland, woodland, or brackish marshes.
32. Nemum Desv. ex Ham.
Nemum Desv. ex Ham., Prodr. Pl. Ind. Occid.: 13 (1825);
Raynal, Adansonia II, 13: 146-151 (1973).
Annuals, or tufted to creeping rhizomatous peren-
nials. Culms scapose. Leaves often (spiro-)
distichous, ligulate (most perennials) or eligulate
(all annuals), with a bundle of long pale hairs at
both sides of the sheath mouth, blade rarely ensi-
form. Inflorescence sometimes :±: pseudolateral,
(compound) anthelate, but often capitate or re-
duced to 1-few spikelets. Primary bracts 1-few,
small and inconspicuous, not sheathing, often
stiffly erect. Spikelets with many densely spirally
arranged, persistent glumes, each subtending a
flower. Flowers bisexual. Bristles absent. Stamens
1-3. Style 2-(3-)fid, style base not distinct, not
thickened, deciduous. Achene obovate, dorsiven-
trally lenticular, rarely compressed trigonous,
smooth, often shining black, rarely grayish. About
168 Cyperaceae
ten spp., all in Sudano-Zambesian tropical Africa;
the annuals on temporarily wet soil, the perennials
on permanently marshy ground.
33. Nelmesia Van der Veken
Nelmesia Vander Veken, Bull. Jard. Bot Etat 25: 143 (1955).
Tufted annual(?). Culms scapose, 3-4-angular.
Leaves (sub )distichous, ligule an inconspicuous
rim or absent. Inflorescence a single spikelet. Pri-
mary bracts none (or similar to the glumes?).
Spikelet with many densely spirally arranged, fi-
nally deciduous glumes, 5-7 lower glumes empty,
others each subtending a flower. Flowers bisexual.
Hypogynous scales 1, adaxial, 2-fid at the top. Sta-
mens 3. Style 3-fid, style base ± distinct, slightly
thickened, dark, persistent. Achene (immature)
obovate, plano-convex, beaked, smooth. Only one
sp., N. melanostachya Van der Veken, N Zaire, in
temporarily wet, shallow depressions in the sa-
vanna region.
The presence of one adaxial hypogynous scale in
a species of this tribe is highly problematical and
would require a thorough morphological recon-
sideration of the nature of this "spikelet".
7. Tribe Cypereae Dumort. (1829).
34. Ficinia Schrader
Ficinia Schrader, Comm. Soc. Reg. Sci. Gottingen II, 7: 143
(1832), nom. cons.
Sickmannia Nees (1834).
Tufted perennials, frequently rhizomatous,
stoloniferous, or decumbent. Culms scapose,
more rarely few- to many-noded or branched.
Leaves often (conspicuously) ligulate, blade some-
times with scarious margins, or leaf reduced to a
sheath. Inflorescence rarely pseudolateral, mostly
capitate, more rarely compacted paniculate, spi-
cate, or spikelets scattered along a profusely
branched stem, with 1-many spikelets. Primary
bracts leaflike or short, lowermost bract some-
times erect. Spikelets with few to many spiral or
distichous, usually long persistent glumes, each
subtending a flower, or a few lower glumes empty.
Flowers bisexual. Bristles absent. Stamens 3. Style
deeply 3-fid (2-fid in F. lateralis Kunth), rarely
almost undivided, style base not distinct, decidu-
ous. Achene mostly obovate, rounded trigonous,
rarely biconvex, base mostly surrounded by a
tightly enveloping cupular to 3-lobed disk, usually
smooth. About 60 spp., S Africa (especially Cape),
a few extending along the E African mountains;
doubtful records from Madagascar, Australia, and
New Zealand, in open vegetatioq, often on sandy
or rocky ground.
The hypogynous disk is often considered highly
diagnostic for this genus. There are, however, a
few species with all typical ficinioid characters,
but without the disk (F. tenuifolia Kunth, F. rigida
Levyns), blurring the generic delimitation from
Isolepis. On the other hand, Scirpus nodosus Rottb.
(often referred to Scirpoides) shows several typical
ficinioid features (hypogynous disk, embryo of
Ficinia type), but its distribution is much wider
and includes the whole southern circumpolar re-
gion. This morphologically heterogeneous genus
is in need of a thorough revision.
35. Desmoschoenus Hook. f.
Desmoschoenus Hook. f., Fl. Nov.-Zel. 1: 271 (1853); Edgar, Fl.
New Zeal. 2: 170-171 (1970); Browning & Gordon-Gray,
N.Z.J. Bot. 34: 131-134 (1996).
Rhizomatous perennial. Culms scapose. Leaves
ligulate, coriaceous, scabrid, yellowish. Inflores-
cence condensed paniculate, partial inflorescences
with many spikelets in a dense cluster. Primary
bracts ± leaflike, partly adnate to the axis, widened
at the open sheathing base. Spikelets with many
densely spirally arranged deciduous glumes, 1-few
lower glumes empty, middle glumes each subtend-
ing a flower. Flowers bisexual. Bristles absent. Sta-
mens 3. Style deeply 3-fid, style base not distinct,
deciduous. Achene obovoid, concavo-convex,
smooth and shining, base surrounded by a small
cupular disk. Only one sp., D. spiralis (A. Rich.)
Hook. f., New Zealand, on coastal sand dunes.
36. Isolepis R. Br. Fig. 41H,I
Isolepis R. Br., Prodr.: 221 (1810); Raynal, Adansonia II, 17:
49-57 (1977); Wilson, Telopea 2: 161-170 (1981) (Austr.
spp.); Haines & Lye, Sedges & Rushes E. Afr.: 133-142
(1983).
Eleogiton Link (1827).
Annuals or mat-forming perennic:1ls, more rarely
creeping rhizomatous or stoloniferous. Culms
scapose or few- to many-noded. Leaves eligulate,
blade often much reduced. Inflorescence often
pseudolateral, capitate with few-many spikelets,
or reduced to a single spikelet, rarely anthelate.
Primary bracts leaflike or short, lowermost
bract often erect. Spikelets with few-many
spirally arranged, mostly deciduous glumes, each
subtending a flower, but lowermost glume some-
 Cyperaceae
times empty. Flowers bisexual. Bristles absent.
Stamens 1-3. Style (deeply) 2-3-fid, style base not
distinct, not or slightly thickened, persistent.
Achene mostly obovate, thick lenticular to
(rounded) trigonous, often 3-ribbed, beaked,
variously ornamented. About 60 spp., subcos-
mopolitan, with concentrations in southern
Africa and Australia, mostly in wetlands, or
semiaquatic.
Generic delimitation vs. Ficinia and Scirpoides is
highly problematic (see under these genera).
There are also a few (morphologically) typical
isolepidoid species, which have an embryo of the
Cyperus type (I. humillima (Benth.) K.L. Wilson, I.
inundata R. Br., and transitionally in I. hystrix
(Thunb.) Nees).
37. Scirpoides Scheuchzer ex Seguier
Scirpoides Scheuchzer ex Seguier, Pl. Veron. Suppl.: 73 ( 1754);
Wilson, Telopea 2: 162 (1981) (Austr. spp.); Browning &
Gordon-Gray, S. Afr. J. Bot. 60:315-320 (1994) (S Afr. spp.).
Holoschoenus Link (1827).
Karinia Reznicek & McVaugh (1993) (?).
Tufted or shortly rhizomatous perennials. Culms
scapose, thickened at the base. Leaves ligulate or
not, often reduced to a sheath. Inflorescence
pseudolateral, anthelate, or capitate, with many-
numerous spikelets. Primary bracts culmlike,
lowermost bract erect. Branches of partial inflo-
rescences conspicuously crescentiform in cross-
section. Spikelets with many spirally arranged,
long persistent (finally deciduous ?) glumes, each
subtending a flower. Flowers bisexual. Bristles ab-
sent. Stamens 2-3. Style deeply 3-fid, style base not
distinct, persistent. Achenes obovate, compressed
trigonous, beaked, finely puncticulate. About five
spp., S Africa and warmer parts of Eurasia; in
temporarily wet, often sandy habitats.
Problematic species include Scirpus nodosus
Rottb., which shows affinities with Ficinia, S.
pro lifer Rottb., which is more like a robust Isolepis
inundata R. Br., and S. dioecus Boeckeler, which is
dioecious. The monotypic and enigmatic Mexican
genus Karinia is only tentatively allocated here,
since some critical data are still missing.
38. Oxycaryum Nees
Oxycaryum Nees in Mart., Fl. Brasil. 2(1): 90 (1842); Lye, Bot.
Not. 124: 280-286 (1971).
Floating, stoloniferous perennial. Culms scapose.
Leaves (hairy) ligulate, lower leaf sheaths inflated.
Inflorescence anthelate to capitate, with few to
169
many spikelets. Primary bracts leaflike. Partial
inflorescences capitate with normal branching,
eprophyllate, bracts small and glumelike, not
sheathing. Spikelets with several spirally arranged,
persistent glumes, each subtending a flower, apex
of glumes conspicuously thickened. Flowers bi-
sexual. Bristles absent. Stamens 3. Style 2-fid, style
base not distinct, persistent. Achene narrowly
obovate, dorsiventrally plano-convex, beaked,
base, top and margins conspicuously corky. Only
one sp., 0. cubense (Poeppig & Kunth) Lye, scat-
tered in tropical America and Africa, hydrophytic.
39. Kyllingiella R. Haines & Lye
Kyllingiella R. Haines & Lye, Bot. Not. 131: 176 (1978); Haines
& Lye, Sedges & Rushes E. Afr.: 142-143 (1983).
Small, tufted perennials. Culms scapose, the bul-
bously thickened base covered by withered leaf
sheaths. Leaves eligulate. Inflorescence capitate,
with few-many spikelets. Primary bracts leaflike.
Spikelets with few-many spirally arranged, persis-
tent glumes, each subtending a flower. Flowers
bisexual. Bristles absent. Stamens 1(-2). Style 3-
fid, style base not distinct, slightly thickened,
persistent. Achene obovate, compressed subtri-
gonous, with a short beak, minutely puncticulate.
About three spp., tropical Africa and India, in
(temporarily) damp habitats.
40. Androtrichum (Brongn.) Brongn.
Androtrichum (Brongn.) Brongn. in Duperrey, Voy. Monde,
Bot. 2: 177, t. 32 (1834); Pfeiffer, Rev. Sudamer. Bot. 6: 179-
186 (1940).
Rhizomatous perennial with thick roots. Culms
scapose. Leaves reduced to a purple, apically in-
flated leaf sheath. Inflorescence ± pseudolateral,
condensed anthelate with several capitate partial
inflorescences. Primary bracts rather short, lower-
most bract erect, boat-shaped. Spikelets with
several distichous, persistent glumes, each sub-
tending a flower. Flowers bisexual. Bristles absent.
Stamens 3, filaments strongly elongating after
an thesis. Style deeply 3-fid, style base distinct,
slightly thickened, persistent. Achene elliptical,
trigonous, with a thick short beak, smooth. Only
one sp., A. trigynum (Sprengel) H. Pfeiffer, S Brazil
to N Argentina, in coastal dunes.
The inclusion of a second species [Scirpus
giganteus Kunth, as A. giganteum (Kunth) H.
Pfeiffer] as proposed by Pfeiffer (1940), has not yet
been convincingly demonstrated.
170 Cyperaceae
41. Cyperus L.
Cyperus L., Sp. Pl. 1: 44 (1753); Kiikenthal, Pflanzenreich IV.20
(101) (1935-1936); Kern, Fl. Males. I, 7: 592-646 (1974);
Haines & Lye, Sedges & Rushes E Afr.: 145-224, 252-267
(1983); Tucker, Syst. Bot. Monogr. 2, 85 pp. (1983) (C Amer.
spp.); Lye, Lidia 3: 37-72 (1992) (evaluation of Mariscus).
Anosporum Nees (1834).
juncellus (Griseb.) C.B. Clarke (1893).
Mariscus Vahl (1805).
Sorostachys Steudel (1850).
Torulinium Desv. (1825).
Annuals, or tufted, rhizomatous, stoloniferous or
bulbiferous perennials. Culms mostly scapose, tri-
angular to subterete, rarely winged, compressed
or septate. Leaves eligulate, sometimes reduced to
a sheath, sheaths sometimes semisucculent. Inflo-
rescence rarely pseudolateral, anthelate to capi-
tate with few-numerous spikelets, rarely spicate
or reduced to 1 single spikelet. Primary bracts of-
ten large and leaflike, lowermost bract rarely erect.
Partial inflorescences spikelike with spicately or
digitately arranged spikelets, or densely capitate.
Spikelets with few-many distichous (very rarely
spirally arranged) glumes, spikelet bract and
prophyll ± glumelike, each glume subtending a
flower, glumes deciduous or persistent. Rachilla
internodes elongated. Spikelets sometimes de-
ciduous as a unit, or breaking up into 1-flowered
units. Flowers bisexual, very rarely unisexual or
even dioecious. Bristles absent. Stamens 1-3.
Style (2-)3-fid, more rarely (sub )entire, style base
mostly not distinct, not thickened, persistent or
deciduous. Achene often obovate or elliptical,
(sub)trigonous, rarely dorsiventrally compressed,
rarely stipitate, beaked or not, smooth or vari-
ously ornamented, rarely corky thickened. About
550 spp., well represented in the tropics, extending
towards the temperate regions, ranging from very
dry to permanently submerged, from open sunny
places to dark forest, from acid bogs to brackish
marshes, several species on disturbed ground and
becoming weedy.
There is no general agreement either on the cir-
cumscription of the genus or on infrageneric clas-
sification. Pycreus and Kyllinga plus some highly
specialized smaller taxa are often excluded and
recognized at the generic level. Authors who in-
clude these taxa into Cyperus s.l. mostly maintain
them on the subgeneric level. Mariscus, in con-
trast, which is kept separate by several authors,
is rarely maintained at subgeneric level when in-
cluded in Cyperus, and is often divided into sev-
eral sections of widely scattered affinities. The
polyphyletic nature of Mariscus has convincingly
been demonstrated by Lye (1992) and is recog-
nized even by authors (Raynal 1973: 166) who
maintained it as a separate genus.
In a recent survey (Goetghebeur 1989) it was
argued that Cyperus, as circumscribed here, can be
divided into two subgenera: subgen. Cyperus with
the Kranz syndrome (chlorocyperoid anatomy:
subgenus Cyperus), and subgen. Anosporum with-
out Kranz cells (eucyperoid anatomy), the first
group often with spicately arranged spikelets,
the second with spikelets digitately arranged or
capitately condensed. Problems are caused, how-
ever, by the presence of several chlorocyperoid
sections with capitate inflorescences. Further
studies are needed for a better understanding of
this notoriously difficult genus.
42. Courtoisina Sojak
Courtoisina Sojak, Cas. Nar. Mus., Odd. Prir. 148: 193 (1980);
Haines & Lye, Sedges & Rushes E Afr.: 174-175 (1983).
Courtoisia Nees (1834), nom. illeg., non Marchand (1830).
Tufted annuals with a scent of curry. Culms sea-
pose. Leaves eligulate. Inflorescence anthelate,
with terminal clusters of many densely arranged
spikelets, rarely completely contracted. Primary
bracts leaflike, spreading. Spikelets with 2-several,
distichous, persistent, winged glumes, usually
each subtending a flower. Rachilla internodes
elongated. Spikelets deciduous as a unit. Flowers
bisexual. Bristles absent. Stamens 3. Style 3-fid,
style base not distinct, slightly thickened, persis-
tent. Achene oblong, trigonous, beaked, finely
puncticulate. Two spp., E and S Africa, one also in
Madagascar, India, and SE Asia; often on tempo-
rarily wet sandy soils.
43. Remirea Aublet
Remirea Aublet, Hist. Pl. Guiane 1: 44, t. 16 (1775); Hooper,
Kew Bull. 38: 479-480 (1983).
Small perennial with creeping rhizomes. Culms
many-noded, concealed by the leaf sheaths.
Leaves eligulate, blade rigid, subsucculent. Inflo-
rescence capitate with few-many spikelets. Pri-
mary bracts leaflike, suberect to spreading.
Spikelets with 3-4 distichous, persistent glumes,
1st and 2nd (fertile) glume large, 3rd small, sitting
on a finally much enlarged corky rachilla in-
ternode. Rachilla internodes elongated. Spikelet
deciduous as a unit. Flowers bisexual. Bristles
absent. Stamens 3. Style 3-fid, style base not dis-
tinct, not or slightly thickened, persistent. Achene
elliptical, rounded trigonous, beaked, finely
 Cyperaceae 171

puncticulate, clasped by the much enlarged, corky 46. Alinula Raynal

uppermost rachilla internode. Only one sp., R.
Alinula Raynal, Adansonia II, 17: 43 (1977); Goetghebeur &
maritima Aublet, pantropical along sandy sea Vorster, Bull. Jard. Bot. Belg. 58: 457-465 (1988).
shores and littoral dunes. Aliniella Raynal (1973), nom. illeg., non Skvortzow (1958).
The spikelet structure was longtime misunder- Marisculus Goetghebeur (1977).
stood, until Kern {1958: 795-798) provided a con-
vincing interpretation. Small tufted annuals. Culms scapose. Leaves
eligulate. Inflorescence sometimes ± pseu-
dolateral, (bis)anthelate to capitate with 1-many
44. Sphaerocyperus Lye
spikes. Primary bracts leaflike, not sheathing,
Sphaerocyperus Lye, Bot. Not. 125: 214 (1972); Browning & spreading or lowermost bract erect. Spikes with
Gordon-Gray, Nord. J. Bot. 13: 507-510 (1993). many densely spirally arranged spikelet bracts,
much smaller than their spikelet. Spikelets re-
Perennial with a creeping rhizome. Culms sea- duced to a small prophyll and a larger glume, in
pose, thickened at the base. Leaves eligulate, lower some species accompanied by a rachilla remnant,
leaves reduced to a sheath. Inflorescence capitate glumes open, rarely utriculiform. Flowers bi-
with !-several indistinct dense spikes of many sexual. Bristles absent. Stamens 1-2. Style deeply
spikelets. Primary bracts leaflike, spreading to re- 3-fid, style base not distinct, not or slightly thick-
flexed. Spikelets with several distichous, persistent ened, persistent. Achene oblong to obovate,
glumes of increasing length, largest penultimate rounded trigonous, (shortly) beaked, rarely sur-
glume subtending a flower. Rachilla internodes rounded at the base by a white collar of inflated
somewhat elongated. Spikelet deciduous as a unit. cells. Four spp., E and S Africa, one also in Mada-
Flowers bisexual. Bristles absent. Stamens 3. Style gascar, growing as ephemeral annuals on tempo-
3-fid, style base not distinct, not or slightly rarily wet sandy soils.
thickened, persistent. Achene narrowly obovate,
rounded trigonous, beaked, finely puncticulate.
Only one sp., S. erinaceus (Ridley) Lye, S tropical 47. Ascolepis Nees ex Steudel
Africa, often in open dry woodland. Ascolepis Nees ex Steudel, Syn. Pl. Glumac. 2: 105 (1855), nom.
cons.; Goetghebeur, Adansonia II, 19: 269-305 (1980);
Haines & Lye, Sedges & Rushes E Afr.: 302-311 (1983).
45. Ascopholis C. Fischer
Annuals or tufted perennials, rarely rhizomatous
Ascopholis C. Fischer, Kew Bull. 1931: 104 (1931), and in
or stoloniferous. Culms scapose, often bulbously
Hooker's Icon. Pl. 32: t. 3183 (1933); Nijalingappa &
thickened at the base. Leaves eligulate. Inflores-
Goetghebeur, Bioi. Jaarb. Dodonaea 57: 81-86 (1990).
cence capitate with 1-few spikes. Primary bracts
Small perennial with bulbously thickened stem leaflike, spreading to reflexed. Spikes often vividly
base. Culms seapose. Leaves eligulate, leaf sheaths colored, with many densely spirally arranged
swollen. Inflorescence capitate with several dense spikelet bracts, much smaller than their spikelet.
spikes of many spikelets, subtended by a small Spikelets reduced to 1 glume, eprophyllate, in
bract. Primary bracts leaflike, strongly reflexed some species accompanied by a rachilla remnant,
after an thesis. Spikelets reduced to a prophyll and glume polymorphic, rarely utriculiform, often
a single glume, this glume utriculiform and with elongated or swollen upper part. Flowers
subtending the single flower, also including a bisexual. Bristles absent. Stamens 1-3( -5). Style
bristlelike rachilla remnant (at the top with an 2-3(-5)-fid, style base not distinct, deciduous.
extremely reduced second glume). Spikelet de- Achene obovate to oblong, rounded trigonous to
ciduous as a unit. Flowers bisexual. Bristles ab- lenticular, rarely stipitate, surface finely punc-
sent. Stamens 3. Style deeply 3-fid, style base not ticulate. About 20 spp., pantropical, mainly in
distinct, persistent. Achene narrowly obovate, ± Africa, the perennials growing in marshes, the
compressed subtrigonous, shortly beaked, surface annuals preferring temporarily wet soils.
smooth. Only one sp., A. gamblei C. Fischer, S The rather common African A. protea Welw. is
India, in open grassland. highly polymorphic and inviting a biosystematic
study.
172 Cyperaceae

48. Pycreus P. Beauv. 50. Kyllinga Rottb.

Pycreus P. Beauv., Fl. Oware 2: 48, t. 86 (1816); Kiikenthal, Kyllinga Rottb., Descr. Icon. Rar. Pl.: 12, t. 4 (1773), nom.
Pflanzenreich IV.20 (101): 326-402 (1936); Haines & Lye, cons.; Kiikenthal, Pflanzenreich IV.20 (101): 566-614
Sedges & Rushes E. Afr.: 268-292 (1983). (1936); Getliffe, J.S. Afr. Bot. 49: 261-304 (1983) (S Afr.
spp.); Haines & Lye, Sedges & Rushes E. Afr.: 224-250
Annuals or tufted, rhizomatous, stoloniferous or (1983); Tucker, Rhodora 86:507-538 (1984) (C Amer. spp.).
tussock-forming perennials. Culms mostly sea-
pose. Leaves eligulate. Inflorescence sometimes Annuals or tufted to creeping perennials. Culms
pseudolateral, anthelate (rarely spicate) to capi- scapose to few-noded, sometimes 6-angular or
tate with few-many spikelets. Primary bracts winged. Leaves eligulate, lowermost leaves often
leaflike, not sheathing, sometimes the lowermost reduced to a sheath. Inflorescence rarely pseu-
bract larger and erect. Partial inflorescences often dolateral, ±capitate, with 1-many sessile, globose
± open spikelike, with few-many sessile spikelets to cylindrical spikes. Primary bracts mostly
in the axil of glumelike bracts. Spikelets with few- leaflike. Spikes with few-many, densely (rarely
many distichous, deciduous glumes, each sub- loosely) spirally arranged glumelike bracts, each
tending a flower. Rachilla internodes elongated. subtending a spikelet. Spikelets with 2-few disti-
Flowers bisexual. Bristles absent. Stamens 1-3. chous persistent glumes, 1-few subtending a
Style 2-fid, style base not distinct, mostly decidu- flower. Rachilla internodes elongated. Spikelets
ous. Achene usually obovate or rounded, laterally deciduous as a unit. Flowers bisexual. Bristles
lenticular to subcylindrical or subglobose, rarely absent. Stamens 1-3. Style 2-fid, style base not
subtrigonous, smooth, puncticulate, or trans- distinct, persistent. Achene often obovate, later-
versely rugose. About 100 spp., pantropical, a few ally lenticular, often minutely puncticulate. About
extending into the temperate regions, many in 60 spp., pantropical to warm temperate, concen-
Africa, growing in shallow water, rarely floating or trated in E Africa, often in rather dry to seasonally
submerged, or in permanently to seasonally wet wet grassland or woodland, some becoming
grassland, annual species often on thin soil overly- weedy.
ing rocks.
51. Volkiella Merxm. & Czech
49. Queenslandiella Domin Volkiella Menan. & Czech, Mitt. Bot. Staatssamml. Miinch. 8:
Queenslandiel/a Domin, Biblioth. Bot. 85: 415 (1915); Ballard, 317 (1953); Hooper, Kew Bull. 41: 945-948 (1986).
Hooker's Icon. Pl. V, 3: t. 3208 (1933).
Mariscopsis Chermezon (1919). Small tufted annual. Culms very small or up to
3 em long, scapose. Leaves few-absent, eligulate.
Tufted annual with scent of curry. Culms sea- Inflorescence capitate. Lower primary bracts
pose. Leaves eligulate. Inflorescence anthelate. leaflike. Partial inflorescences spikeletlike, but
Primary bracts leaflike, not sheathing. Partial in- composed of several-many distichous glumelike
florescences ± open spikelike, with few to many bracts, each subtending a highly reduced spikelet.
sessile spikelets subtended by a glumelike bract. Spikelets with a prophyll and a tiny, hyaline glume
Spikelets with few-several distichous, persistent subtending a flower. Flower bisexual. Bristles ab-
glumes, each subtending a flower. Rachilla inter- sent. Stamens 2. Style deeply 3-fid, style base not
nodes elongated. Spikelets deciduous as a unit. distinct, deciduous. Achene cordate-obovate,
Flowers bisexual. Bristles absent. Stamens 2. Style subtriquetrous, finely puncticulate. Only one sp.,
2-fid, style base not distinct, deciduous. Achene V. disticha Merxm. & Czech, N Namibia and Zam-
subcordate, laterally compressed, finely puncti- bia, an ephemerophyte of temporarily wet sandy
culate. Only one sp., Q. hyalina (Vahl) Ballard, E soils, often nearly completely buried in sand.
Africa over Madagascar, India, Sri Lanka, Malesia
to NE Australia, growing as an ephemerophyte of
temporarily wet sandy, often saline or mineral- 52. Lipocarpha R. Br.
rich soil. Lipocarpha R. Br. in Tuckey, Narr. Exped. Zaire 5: 459 (1818),
nom. cons.; Haines & Lye, Sedges & Rushes E. Afr.: 294-302
(1983); Hooper, Kew Bull. 41: 423-428 (1986); Goetghebeur
& Van den Borre, Agr. Univ. Wageningen Pap. 89-1,87 pp.
(1989).
Hemicarpha Nees (1834).
Rikliel/a Raynal (1973).
 Cyperaceae 173

Annuals or tufted or rhizomatous, rarely stolonif- Loosely tufted perennial. Culms few-noded. Lower
erous perennials. Culms scapose. Leaves eligulate. leaves ligulate. Inflorescence corymbose or
Inflorescence sometimes pseudolateral, capitate anthelate, with many corymbiform partial inflo-
with 1-many spikeletlike spikes. Lower primary rescences and numerous spikelets, cladoprophylls
bracts mostly leaflike, lowermost bract sometimes often fertile. Primary bracts leaflike, lower bracts
larger and erect. Spikes extremely spikeletlike, sheathing, upper bracts not. Spikelets with many
composed of many glumelike bracts, spirally ar- spirally arranged persistent glumes, each subtend-
ranged along a thickened to shortly conical axis, ing a flower (including the prophyll). Rachilla in-
each subtending a highly reduced spikelet. Spike- ternodes somewhat elongated. Spikelet deciduous
lets either prophyllate and with 1 small ± hyaline as a unit, or breaking up into 1- to few-flowered
glume, subtending a flower, or further reduced to parts. Flowers bisexual. Bristles 5-6, longer
the small ± hyaline prophyll, or even lacking this than the fruit, deciduous with the fruit. Stamens 3.
prophyll. Prophyll and glume, when present, often Style (2- )3-fid, style base not distinct, slightly
deciduous with the fruit. Flowers bisexual. Bristles thickened, persistent. Achene oblong, rounded
absent. Stamens 1-3. Style deeply 2-3-fid, style trigonous (or dorsiventrally lenticular), stipitate,
base not distinct, thickened or not, deciduous or beaked, finely puncticulate. Only one sp., S.
persistent. Achene ± obovoid to subcylindrical, junghuhnii (Miq.) Oteng-Yeb., N Sumatra, in
often constricted at the base, sometimes beaked, montane heathlands.
surface finely puncticulate. About 35 spp., pan-
tropical and extending into some temperate re-
gions, many in Africa, in open vegetation on 55. Blysmus Panzer ex Schultes
permanently to seasonally wet to humid soil, Blysmus Panzer ex Schultes, Mant. 2: 41 (1824), nom. cons.;
rarely in dry woodland. Oteng-Yeboah, Notes R. Bot. Gard. Edinburgh 35: 399-406
(1977).
Blysmopsis Oteng-Yeb. (1974).
8. Tribe Dulichieae Reichenb. ex Schultze-Motel
(1959).
Small perennials with a creeping rhizome. Culms
(sub )scapose. Leaves ligulate. Inflorescence spi-
53. Dulichium Rich.
cate with few-many pseudodistichous spikelets,
Dulichium Rich. in Pers., Syn. Pl. 1: 65 (1805); Kiikenthal, Bot. rarely with a lateral branch (with a fertile
Jahrb. Syst. 75: 485-488 (1952). cladoprophyll). Primary bracts mostly small, low-
ermost bract ± leaflike, suberect. Spikelets with
Perennial on a creeping rhizome. Culms many- few-many spirally arranged persistent glumes
noded, hollow. Lower leaves reduced to a sheath, (pseudodistichous in the terminal spikelet), each
upper leaves ligulate, contraligule rounded to glume (including the prophyll) subtending a
subtriangular. Inflorescence paniculate with many flower. Rachilla internodes somewhat elongated.
spikelets. Primary bracts leaf-like. Partial inflores- Spikelet deciduous as a unit, or breaking up into
cences spikelike, with pseudodistichous, distant 1-few-flowered parts. Flowers bisexual. Bristles
glumelike bracts, each subtending a spikelet. (1- )3-5, deciduous with the fruit. Stamens 3. Style
Spikelets with 4-8 distichous, persistent glumes, 2-fid, style base not distinct, thickened, persistent.
each (including the prophyll) subtending a flower. Achene oblong, dorsiventrally lenticular or
Rachilla internodes elongated, fruiting rachilla planoconvex, stipitate, beaked, smooth. About
thickened and finally breaking up into 1-flowered four spp., temperate Eurasia, B. rufus (Hudson)
parts. Flowers bisexual. Bristles (6- )8( -9), longer Link also in N America, in swamps, often on
than fruit, deciduous with the fruit. Stamens 3. brackish or compacted soils.
Style 2-fid, style base not distinct, somewhat thick-
ened, persistent. Achene oblong, dorsiventrally ±
lenticular, stipitate, and beaked, surface smooth. 9. Tribe Schoeneae Dumort. (1827).
Only one sp., D. arundinaceum (L.) Britton, tem-
perate to subtropical N America, in swamps and 56. Arthrostylis R. Br.
marshes; fossil records from W Europe.
Arthrostylis R. Br., Prodr.: 229 (1810); Kiikenthal, Repert.
Spec. Nov. Regni Veg. 53: 192-197 (1944).
54. Sumatroscirpus Oteng-Yeb.
Sumatroscirpus Oteng-Yeb., Notes Roy. Bot. Gard. Edinburgh Tufted perennial. Culms scapose, 3-5-angular.
33: 307 (1974). Leaves ligulate, reduced to a (mucronate) sheath.
174
Inflorescence capitate with few-several spikelets.
Primary bracts small. Spikelets with 5-7 spirally
arranged, deciduous glumes of increasing length,
1-flowered, penultimate larger glume subtending
a flower, enclosed by the wings of the next glum e.
Flowers bisexual. Bristles absent. Stamens 6. Style
deeply 3-fid, stigmatic branches thick-papillose,
style base distinct, slightly thickened, deciduous.
Achene obovate, rounded trigonous, substipitate,
finely reticulate. Only one sp., A. aphylla R. Br.,
NE Australia, on sandstone outcrops in open
woodland.
57. Actinoschoenus Benth.
Actinoschoenus Benth., Hooker's Icon. Pl. 14: 33, t. 1346
(1881); Raynal, Adansonia II, 7: 89-95 (1967).
Tufted, shortly rhizomatous or stoloniferous
perennials. Culms scapose. Leaves ligulate, blade
very short. Inflorescence capitate with 2-many
spikelets. Primary bracts small. Spikelets with
4-7 distichous, deciduous glumes of increasing
length, 1-(2-)flowered, flower subtended by the
penultimate larger glume enclosed by the wings
of the next glume. Rachilla internodes short,
somewhat elongated between the flowers. Flowers
bisexual. Bristles absent. Stamens 3. Style deeply
3-fid, style base distinct, thickened, deciduous.
Achene obovate, trigonous, ± 3-ribbed, smooth to
slightly tuberculate. Three spp. of scattered occur-
rence, Gabon, SE Zaire and Zambia, Madagascar,
Sri Lanka, SE Asia, Philippines, New Caledonia, in
open woodland on dry sandy soil.
A fourth sp., which agrees well in floral charac-
ters, Fimbristylis yunnanensis C.B. Clarke, could
be included here, but the decisive anatomical and
embryological data are still lacking.
58. Trichoschoenus Raynal
Trichoschoenus Raynal, Adansonia II, 8: 223 (1968).
Tufted perennial. Culms scapose, densely hairy.
Leaves reduced to a sheath, eligulate (?), blade
sometimes present as a short mucro. Inflorescence
capitate with many spikelets. Primary bracts
mostly as long as or shorter than spikelet. Spike-
lets with 3 distichous, deciduous glumes of in-
creasing length, 2nd glume subtending a flower,
enclosed by the wings of the next glume. Flowers
bisexual. Bristles absent. Stamens 3. Style deeply
3-fid, style base ± distinct, thickened, persistent.
Achene obovate, trigonous, with a thick short
beak, minutely reticulate. Only one sp., T. bosseri
Cyperaceae
Raynal, Madagascar, in open woodland on dry
sandy soil.
59. Trachystylis S. T. Blake
Trachystylis S.T. Blake, Proc. R. Soc. Queensland 48: 89 (1937);
Kiikenthal, Bot. Jahrb. Syst. 75: 493-497 (1952).
Medium-sized tufted perennial. Culms (sub-)
scapose. Leaves ligulate. Inflorescence anthelate
with many spikelets. Primary bracts rather
small. Spikelets with several spirally arranged,
deciduous glumes of increasing length, penulti-
mate larger glume subtending a flower enclosed
by the wings of the next glume (rarely a 2nd flower
present). Rachilla internodes short, somewhat
elongated between the flowers. Flowers bisexual.
Bristles absent. Stamens 2, lateral. Style 2-fid, style
base distinct, thickened, deciduous. Achene ob-
ovate, shortly stipitate, dorsiventrally (thick) len-
ticular, smooth, finely reticulate. Only one sp., T.
stradbrokensis (Domin) Kiik., E Australia, in open
woodland on dry sandy soil.
60. Rhynchospora Vahl Fig. 42A-C
Rhynchospora Vahl, Enum. Pl. 2: 229 (1805), nom. cons.;
Kiikenthal, Bot. Jahrb. Syst. 74: 375-509 (1949), 75: 90-195
(1950) & 75: 273-314 (1951); Haines & Lye, Sedges & Rushes
E. Afr.: 312-320 (1983); Thomas, Mem. N.Y. Bot. Gard. 37:
116 pp. (1984) (sect. Dichromena); Thomas, Brittonia 44:
14-44 (1992) (C Amer. spp.).
Dichromena Michaux (1803).
Psilocarya Torrey (1836).
Syntrinema H. Pfeiffer (1925).
Micropapyrus Suesseng. (1943).
Tufted or creeping rhizomatous perennials, more
rarely annuals, or stoloniferous to caudex-
formingperennials. Culms 1-many-noded. Leaves
eligulate. Inflorescence rarely ± pseudolateral,
paniculate, corymbose, anthelate, or capitate, with
1-numerous spikelets. Primary bracts leaflike or
short, sometimes partly white (part of section
Dichromena). Spikelets with few-several spirally
arranged to more rarely distichous, often decidu-
ous glumes of increasing lep.gth to subequal, larger
(1- )2-few glumes each subtending a flower, en-
closed by the wings of the next glume. Flowers all
bisexual, or upper flowers functionally male.
Bristles 0-6( -10?), deciduous with the fruit. Sta-
mens 1-3(-6-12), sometimes partly adnate. Style
deeply 2-fid or (nearly) undivided, style base
mostly distinct, (much) thickened, persistent.
Achene usually (ob )ovate and dorsiventrally
(thick) lenticular, beaked, yvith various ornamen-
 Cyperaceae
Fig. 42A-J. Cyperaceae. A-C Rhynchospora (sect. Dichro-
mena) latifolia. A Habit, head subtended by showy bracts.
B Spikelet. C Achene with enlarged style basis. D- F
Cephalocarpus dracaenula. D Habit. E Pistillate flower. F
Mature fruit, apical portion of the pericarp abscising. G-J
Oreobolus obtusangulus. G Habit. H Flower subtended by
glumes and bracts, only tips of tepals visible. I Persistent peri-
anth after shedding of the achene. J Achene. (Takhtajan 1982)
tations, more rarely smooth. About 250 spp.,
subcosmopolitan, with a marked concentration in
(sub)tropical America, most species preferring
humid to wet habitats in savanna vegetation.
175
61. Pleurostachys Brongn.
Pleurostachys Brongn. in Duperrey, Voy. Monde, Bot. 2: 172, t.
31 (1833); Kiikenthal, Bot. Jahrb. Syst. 75: 451-484 (1952).
Shortly rhizomatous perennials. Culms many-
noded and leafy, rarely scapose, often thickened at
the base. Leaves rarely all basal and then pseu-
dopetiolate, lower leaves often reduced to a ± in-
flated sheath, upper leaves eligulate, blade often
oblong. Inflorescence paniculate, with corymbose
to anthelate or contracted partial inflorescences,
with many-numerous spikelets. Primary bracts
leaflike or more rarely short, sheathing. Spikelets
small, with several to many spirally arranged or
more rarely distichous, long persistent glumes of
increasing length, lower glumes empty, middle
(3-)5-7 glumes each subtending a flower, en-
closed by the wings of the next glume, few upper
glumes empty. Lower 1-2 flowers bisexual, upper
flowers functionally male. Bristles 3-6, alope-
curoid (lower part not or slightly ciliolate, upper
part densely ciliate to fimbriate), shorter than to as
long as the fruit, deciduous with the fruit. Stamens
3. Style deeply 2-fid, style base scarcely distinct,
conspicuously thickened, persistent. Achene
small, (ob )ovate, dorsiventrally thick lenticular,
often rugose. About 30 spp., tropical and subtropi-
cal S America, mostly in forests.
62. Schoenus L.
Schoenus L., Sp. Pl. 1:42 (1753); Kiikenthal, Repert. Spec. Nov.
Regni Veg. 44: 2- 32, 65- 101, 161- 195 {1938); Kern, Fl.
Males. I, 7: 672-680 (1974).
Ptilanthelium Steudel (1855).
Tufted or rhizomatous perennials, rarely annuals
or short-living perennials. Culms mostly scapose,
rarely 3-4-angular, sometimes bulbously thick-
ened at the base, or creeping-decumbent and
branched from the nodes (semiaquatic spp. of
sect. Helothrix). Leaves ligulate, often reduced to a
sheath. Inflorescence sometimes ± pseudolateral,
paniculate to capitate, with 1-many spikelets. Pri-
mary bracts leaftike or short, sheathing or not,
lowermost bract sometimes erect. Spikelets with
few-many distichous, deciduous glumes of in-
creasing length, but upper glumes smaller, the
larger (1- )2-3( -9) glumes each subtending a
flower, enclosed by the wings of the next glume.
Rachilla internodes between and above the flowers
often elongated and curved (short in, e.g., subgen.
Pseudomesomelaena). Lower 1-2 flowers usually
bisexual, upper 1-2 flowers (functionally) male
(rarely inversed). Bristles 0-6, rarely laminar,
176 Cyperaceae

shorter or longer than the fruit, usually deciduous wards very much broadened at the base, top
with the fruit (persistent on the rachilla in sect. scabrid, abruptly acuminate, and twisted apically.
Helothrix). Stamens {1- )3-6. Style 3-fid [2-(3-) Stamens 3, connective produced into a long subu-
fid inS. tendo (Hook. f.) Hook. f.], style base not late appendage. Style trifid, style base distinct,
distinct, not or slightly thickened, deciduous, slightly thickened, deciduous, scabrid. Achene
rarely persistent. Achene often obovate and ovate, rounded trigonous, often seated on a thick-
rounded trigonous, rarely beaked, with various ened cuplike or 3-lobed gynophore. Five spp., SW
ornamentations. About 100· spp., concentrated Australia in heath formations.
in Australia and Malesia, S. nigricans L. sub-
cosmopolitan, often in humid grassland or
woodland. 65. Ptilothrix K.L. Wilson
A very particular species, S. clandestinus S.T. Ptilothrix K.L. Wilson, Telopea 5: 612 (1994); Kiikenthal,
Blake, is "forming small practically stemless mats Repert. Spec. Nov. Regni Veg. 47: 211-216 (1939) (sub
with only the leaf blades and styles projecting Ptilanthelium).
above the surface of the sand" (Blake, Proc. R. Soc. Ptilanthelium auct. non Steudel: Kiikenthal (1939).
Queensland 60: 50. 1949), recalling the situation of
the African Volkiella disticha Merxm. & Czech. Tufted perennial on a short woody, seriate
Schoenus asperocarpus F. Muell. is reported as rhizome. Culms scapose. Leaves spirodistichous,
poisonous to cattle (Huxtable et al., Austral. Veter. ligulate. Inflorescence capitate with many spike-
J. 70: 169-171. 1993). lets. Primary bracts small, short and stiff, sub erect,
sheathing. Spikelets with several distichous,
deciduous glumes of increasing length, upper 2
63. Gymnoschoenus Nees larger glumes each subtending a flower, enclosed
Gymnoschoenus Nees, Ann. Mag. Nat. Hist. I, 6: 47 (1841); by the wings of the next glume. Rachilla inter-
Kiikenthal, Repert. Spec. Nov. Regni Veg. 48: 56-60 (1940). nodes short, but upper internode elongated.
Lower flower bisexual, upper flower male or absent
Medium-sized-tall tufted perennials. Culms sea- (rarely bisexual?). Bristles 3, lower half fimbriate,
pose. Leaves with a ciliate ligule. Inflorescence upper half widened and scab rid, geniculate, longer
capitate with many spikelets. Primary bracts than the fruit, deciduous with the fruit. Stamens 3.
short. Spikelets with several distichous, persistent Style 3-fid, style base not distinct, elongate,
(?) glumes of increasing length, upper 2 larger persistent, glabrous. Achene oblong, rounded
glumes each subtending a flower, enclosed by the trigonous, long-beaked, smooth. Only one sp., P.
wings of the next glume. Lower flower functionally deusta (R. Br.) K.L. Wilson, E Australia, in open
male, upper flower bisexual. Bristles (0-)1-3, vegetation.
short and poorly developed. Stamens 3. Style 3-fid,
style base not distinct, thickened, persistent,
slightly scabrid to subglabrous. Achene obovate, 66. Cyathochaeta Nees
acutely trigonous, beaked, surface smooth. Two Cyathochaeta Nees in Lehmann, Pl. Preiss. 2: 86 (1846);
spp., Australia, in swamps or heathlike vegetation. Kiikenthal, Bot. Jahrb. Syst. 75: 488-493 (1952).

Tufted perennials. Culms few-noded. Leaves

64. Mesomelaena Nees
Mesomelaena Nees in Lehmann, Pl. Preiss. 2: 88 (1846);
Wilson, Telopea 2: 181-195 (1981).
Tufted perennials. Culms scapose or few-noded.
Leaves ligulate, sometimes reduced to a sheath.
Inflorescence often pseudolateral, capitate with
(1- )few-many spikelets. Primary bracts rather
short. Spikelets with several distichous, long per-
sistent, awned glumes of increasing length,
penultimate larger 2 glumes each subtending a
flower, enclosed by the wings of the next glume.
Lower flower functionally male, upper flower
bisexual. Bristles 3, before anthesis slender, after-
sometimes spirodistichous, ligulate. Inflorescence
paniculate with few-many spikelets. Primary
bracts ± leaflike, sheathing. Spikelets with 4 sub-
distichous to spirally arranged persistent glumes
of increasing length, upper 2 each subtending a
flower, enclosed by the wings of the next glume.
Lower flower functionally male, upper flower bi-
sexual. Bristles 4-5, fimbriate at the base, as long
as the fruit, deciduous with the fruit. Stamens 2,
lateral. Style 2-fid, style base not distinct, not
thickened, finally deciduous. Achene oblong, dor-
siventrally compressed and remarkably incurved
(dorsally very concave). Four spp., SW and E
Australia, in marshes.
 Cyperaceae
67. Oreobolus R. Br. Fig. 42G-J
Oreobolus R. Br., Prodr.: 235 (1810); Seberg, Bot. J. Linn. Soc.
96: 119-195 (1988), rev.
Voladeria Benoist (1933) (sub Juncaceae).
Schoenoides Seberg (1986).
Small, tufted to cushion-forming perennials.
Culms few-many-noded, leafy. Leaves ortho- or
spirodistichous, with an open sheath, eligulate,
base of blade often narrowed into a clear
pseudopetiole. Inflorescence reduced to 1-few
spikelets in a condensed depauperate panicle (±
concealed among the leaves before anthesis). Pri-
mary bracts leaflike, sheathing, upper (empty)
bracts gradually glumelike. Spikelets with 3-5 dis-
tichous, deciduous glumes, penultimate glume(s)
subtending 1(2-3) flower(s), enclosed by the
wings of the next glume. Flowers bisexual; upper
and lower flower, when present, functionally male.
Scales 6, about as long as fruit or shorter, narrowly
triangular to more bristlelike, persistent on the
rachilla. Stamens 3. Style deeply 3-fid, base not
distinct, slightly thickened, rarely persistent.
Achene obovate or pyriform, rarely fusiform,
rarely beaked, rounded trigonous, smooth. About
15 spp., Malesia to SE Australia, Tasmania, New
Zealand, Tahiti, Hawaii, Juan Fernandez Is.,
Falkland Is., W South America, Central America;
in wet alpine and subantarctic vegetation.
68. Carpha Banks & Sol. ex R. Br.
Carpha Banks & Sol. ex R. Br., Prodr.: 230 (1810); Kiiken-
thal, Repert. Spec. Nov. Regni Veg. 47: 101-119, 209-216
(1939); Reid & Arnold, Bothalia 15: 139-142 (1984) (S Afr.
spp.).
Asterochaete Nees (1834).
Oreograstis K. Schum. (1895).
Tufted, mat-forming (rarely stoloniferous) peren-
nials. Culms scapose or with a few nodes. Leaves
eligulate. Inflorescence open to condensed pan-
iculate with (sub )capitate partial inflorescences
with few to many spikelets. Lower primary bracts
± leaflike, sheathing. Spikelets with 3-6 disti-
chous, persistent glumes of increasing length,
lower glumes empty, larger 1-2( -3) glume(s) each
subtending a flower, enclosed by the wings of the
next glume. Flowers bisexual, sometimes upper or
lower flower functionally male. Bristles 6, ciliate to
(partly) fimbriate or plumose, shorter than to
much longer than the fruit, deciduous with the
fruit. Stamens 2-3, anthers conspicuously green-
ish yellow. Style 3-fid, base not distinct, (slightly)
thickened, persistent. Achene narrowly (ob)ovate
177
to oblong, trigonous, often with a long beak, sur-
face smooth or finely reticulate. About 15 spp., S
Africa and the Central African mountains, Mada-
gascar, Mascarenes, New Guinea, S Japan, SE Aus-
tralia, New Zealand, and Chile, in swamps from
low to high altitudes, often along stream sides or
rivulets.
69. Trianoptiles Fenzl
Trianoptiles Fenzl in Endl., Gen. Pl.: 113 (1836); Levyns, J.S.
Afr. Bot. 9: 21-26 (1943); Haines & Lye, Bot. Not. 130: 235-
240 (1977).
Small, loosely tufted annuals, with frequent
amphicarpy. Culms few-noded. Leaves eligulate.
Basal leaves often reduced to a sheath, subtending
a basal 1-flowered female spikelet. Basal spikelet
prophyllate, with a single utriculiform glume,
tightly wrapped around the female flower, top
conspicuously tubuliform, flower reduced to a 3-
carpellate pistil with an elongated style, achene
larger and thicker than the aerial fruits, ± irregu-
larly shaped. Inflorescence depauperate panicu-
late with 1-few spikelets. Primary bracts leaflike,
sheathing. Spikelets with 3-5 distichous, long per-
sistent glumes, of increasing length, larger (1-)2
glume(s) each subtending a flower, enclosed by
the wings of the next glume. Rachilla internodes
short, but upper rachilla internode conspicuously
elongated. Flowers bisexual. Scales 6, deciduous
with the fruit, outer very short and thick, inner
bristlelike. Stamens 3. Style 3-fid, style base not
distinct, thickened, persistent. Achene obovate,
trigonous, 3-ribbed, beaked, surface finely reti-
culate. Three spp., South Africa (SW Cape), in
wetland.
70. Tetraria P. Beauv.
Tetraria P. Beauv., Mem. Cl. Sci. Math. Inst. Natl. France
13(2): 54 (1816); Kiikenthal, Repert. Spec. Nov. Regni
Veg. 48: 195-250 (1940); Levyns, J.S. Afr. Bot. 13: 73-93
(1947).
Tufted perennials, or with a short woody seriate
rhizome. Culms scapose or few-noded, terete
to 3(-4)-angular. Leaves ligulate or not, some-
times reduced to a sheath. Inflorescence open
or contracted paniculate. Primary bracts short
or long and leaflike, sheathing or not. Spikelets
with several to many distichous, long persistent
(finally deciduous ?) glumes of increasing length,
larger (1-)2-3(-4) glumes each subtending a
flower, enclosed by the wings of the next glume.
178
Lower 1-2 (-3) flowers usually functionally male,
upper 1-2 flowers bisexual (rarely male). Bristles
0-3-6 (or more), rather short, or absent. Stamens
3-6(-8). Style 3(-4-9)-fid, style base at first
distinct, slightly to much thickened, persistent,
rarely tuberculate. Achene (ob )ovate, rounded
3-( 4-)angular, beaked, surface often smooth.
About 50 spp., most in S Africa (ca. 45), a few
in the mountains of SE and Central Africa (2),
SW Australia (6), and New Zealand (1). T.
borneensis Kern in lowland Borneo doubtfully
kept here. Growing in rather dry, sandy, or rocky
places on mountain slopes, more rarely in marshy
places.
71. Cyathocoma Nees
Cyathocoma Nees, Linnaea 9: 300 (1834); Browning &
Gordon-Gray, S. Afr. J. Bot. 62: 250-257 (1996).
Macrochaetium Steudel (1855).
Medium-sized tufted perennials. Culms scapose.
Leaves eligulate. Inflorescence compacted panicu-
late with many spikelets. Primary bracts leaflike,
sheathing, suberect. Spikelets with 3-5 spirally ar-
ranged, mostly persistent glumes of increasing
length, the larger 2-3(4) each subtending a flower,
enclosed by the wings of the next glume. One
flower bisexual, the other flower(s) functionally
male, position variable. Bristles 6, basally connate,
deciduous with the fruit. Stamens 6. Style 3-fid,
style base distinct, much thickened, persistent.
Achene (ob )ovate, rounded trigona us, or sub-
globular, 3-ribbed, stipitate, beaked, smooth.
Probably three spp., S Africa (S Cape and Natal),
on mountain slopes.
This critical genus is superficially similar to
Tetraria, but differs by the spikelet with fewer and
spirally arranged glumes, a bisexual lower flower,
and basally connate bristles.
72. Neesenbeckia Levyns
Neesenbeckia Levyns, J.S. Afr. Bot. 13: 74 (1947).
Medium-sized to tall tufted perennial. Culms
(sub)scapose or 1-noded. Basal leaves reduced to
a sheath, uppermost leaf subbasal or cauline,
ligulate, blade terete, stiff. Inflorescence ± pseu-
dolateral, contracted paniculate, with many spike-
lets. Primary bracts rather short and stiff, erect,
sheathing. Spikelets with several subdistichous,
persistent glumes of increasing length, 2 middle
glumes largest, each subtending a flower, enclosed
by the wings of the next glum e. Lower flower func-
Cyperaceae
tionally male, upper flower bisexual. Bristles 6, as
long as to longer than the fruit, persistent on the
rachilla. Stamens 3. Style elongated, 6-fid, style
base distinct, thickened, remarkably large and
persistent, smooth. Achene ovoid, subterete, con-
stricted at the base, beaked, irregularly rugose.
Only one sp., N. punctoria (Vahl) Levyns, South
Africa (SW Cape), at stream sides.
This critical species is rather doubtfully kept
separate from the closely related Tetraria. In
Levyns (op.cit.: 47) some of the differences are
enumerated: leaves with reduced blade, but the
uppermost leaf blade large and stemlike, glumes
not clearly distichous, bristles very long, style un-
usually large and terminating in 6 branches. Also
the extremely enlarged style base grown over the
upper half of the fruit, and the bristles persistent
on the rachilla should be noticed.
73. Epischoenus C.B. Clarke
Epischoenus C.B. Clarke in Thiselton-Dyer, Fl. Cap. 7: 273
(1898); Levyns, J.S. Afr. Bot. 25: 69-82 (1959); Browning &
Gordon-Gray, S. Afr. J. Bot. 61: 147-152 (1995).
Medium-sized tufted perennials. Culms (sub)
scapose, terete to trigonous, rarely 4-angular or
compressed. Leaves rarely distich a us (?), eligulate,
often reduced to a sheath, blade rarely ancipitous.
Inflorescence (reduced) paniculate, with few-
many spikelets. Lower primary bracts mostly
rather short and stiff. Spikelets with few-many
distichous long persistent (finally deciduous ?)
glumes of increasing length, upper 2-4 each sub-
tending a flower, enclosed by the wings of the next
glume. Rachilla internodes short, but last intern-
ode elongated, thickened, and curved. Lower 1-3
flowers functionally male, upper flower bisexual.
Bristles absent (sometimes 3 poorly developed).
Stamens 3. Style 3-fid, style base distinct or not, ±
thickened, persistent. Achene ellipsoid to abo-
void, often 3-ribbed, ± beaked, sometimes seated
on a 3-lobed cuplike structure, surface smooth to
tuberculate. About eight spp., South Africa (SW
and S Cape), in damp to marshy places, often low-
to mid-montane.
74. Costularia C.B. Clarke
Costularia C.B. Clarke in Thiselton-Dyer, Fl. Cap. 7: 274
(1898); Kiikenthal, Repert. Spec. Nov. Regni Veg. 46: 13-32,
65-76 (1939); Raynal, Adansonia II, 14: 337-377 (1974) (rev.
New Caledonia); Browning & Gordon-Gray, S. Afr. J. Bot.
61: 66-71 (1995).
Lophoschoenus Stapf (1914).
 Cyperaceae
Tufted or more rarely shortly rhizomatous peren-
nials, sometimes sympodial caudex-forming
dwarfshrubs. Culms scapose or few-noded. Leaves
usually basally crowded or on top of the caudex,
sometimes (spiro- )distichous, eligulate, blade
sometime deciduous. Inflorescence paniculate
with many-numerous spikelets. Primary bracts ±
leaflike, sheathing. Spikelets with several disti-
chous, deciduous glumes of increasing length, the
larger (1-)2(-3) each subtending a flower, en-
closed by the wings of the next flower. The lower
flower (functionally) male or bisexual, the upper
flower bisexual or functiona!ly female. Bristles 6,
fimbriate to ciliate, deciduous with the fruit. Sta-
mens 3. Style 3-fid, style base often distinct (at
anthesis), thickened, persistent. Achene ovate or
oblong, rounded trigonous, often 3-ribbed, ±
stipitate, beaked, smooth or rugulose. About 20
spp., New Caledonia (12) over Malesia (1),
Seychelles, Madagascar, to southern Africa (2);
in scrubby vegetation on rocky ground, rarely in
forest fringes.
The inclusion of Lophoschoenus in Costularia
is debatable, and the vegetative differences
(tristichous leaves without clear midrib) are con-
spicuous. However, the genus Costularia s.l. is in
need of a worldwide revision, and I have here re-
frained from recognizing Lophoschoenus on the
generic level.
75. Gahnia Forster & Forster f.
Gahnia Forster & Forster f., Char. Gen. Pl.: 26 (1775);
Ki.ikenthal, Repert. Spec. Nov. Regni Veg. 52:52-111 (1943);
Kern, Fl. Males. I, 7: 703-710 (1974).
Tufted or shortly rhizomatous perennials, some-
times with a long, erect, or ascending caudex.
Culms scapose or few-noded. Leaves ligulate,
blade flat or inrolled, often inverted (Wilson,
Telopea 5: 601-602. 1994). Inflorescence terminal,
paniculate, partial inflorescences often ± con-
densed, with many-numerous spikelets. Primary
bracts leaflike, sheathing. Spikelets mostly dark
brown or black, with 5-8 spirally arranged,
persistent glumes, 1-2 upper glume(s) each
subtending a flower, enclosed by the wings of
the next glume. Lower flower functionally male,
upper bisexual. Bristles mostly absent. Stamens
(2- )3-6, filaments persistent at the base of the
fruit, with several glume-fixing mechanisms. Style
3(4)-fid, base rather distinct, slightly thickened,
sometimes persistent. Achene ovate to elliptic,
subterete to subtrigonous, sometimes beaked,
often yellowish to reddish colored, with hard,
179
thick fruitwall, endocarp sometimes with grooved
inner side. About 30 spp., SE Asia over Malesia,
Australia, New Zealand, to the Pacific islands, in
swampy to wet places in lowland and at high
altitudes.
76. Morelotia Gaudich.
Morelotia Gaudich. in Freycinet, Voy. Uranie, Bot.: 416, t. 28
(1829); Blake, Contr. Queensland Herb. 8: 38-39 (1969).
Tufted or shortly rhizomatous perennials. Culms
scapose or few-noded. Leaves eligulate, blade flat
to revolute. Inflorescence paniculate with many
spikelets. Lower primary bracts leaflike, sheath-
ing. Spikelets with several (sub)distichous, coria-
ceous, deciduous glumes of increasing length,
upper 2 larger glumes each subtending a flower,
enclosed by the wings of the next glume. Rachilla
internodes short, but upper internode ± elon-
gated and curved. Lower flower bisexual, upper
functionally male. Bristles absent. Stamens 3, fila-
ments elongating after an thesis and clasped by the
partly inrolled glumes. Style 3-fid, style base not or
scarcely distinct, conspicuously corky thickened,
persistent. Achene elliptic-oblong, rounded tri-
gonous, 3-ribbed, with a persistent corky style
base. Two spp., Hawaii, New Zealand, on dry open
hillsides.
77. Reedia F. Muell.
Reedia F. Muell., Fragm. 1: 239, t. 10 (1859); Ki.ikenthal,
Repert. Spec. Nov. Regni Veg. 53: 209-212 (1944).
Tufted perennial on woody rhizome. Culms sea-
pose. Leaves eligulate (?), sheaths often ± inflated.
Inflorescence condensed paniculate with numer-
ous spikelets, resembling a spadix, enclosed
by several large spathelike bracts, these lower
primary bracts empty, bract at the base of
the inflorescence probably subtending a partial
inflorescence, higher bracts smaller and partly
hidden by the densely arranged spikelets. Spike-
lets with many densely spiral to subdistichously
arranged, deciduous (?) glumes of increasing
length, upper 2-3larger glumes each subtending a
flower. Lower flower functionally male, upper bi-
sexual. Bristles absent. Stamens 6, top of filaments
connivent, in the bisexual flower enclosing the
style branches. Style 3-fid, style base not distinct,
thickened, persistent. Achene obovate, rounded
trigonous, beaked, surface smooth, dark and shin-
ing. Only one sp., R. spathacea F. Muell., SW and
W Australia, in swamps.
180 Cyperaceae
78. Evandra R. Br.
Evandra R. Br., Prodr.: 239 (1810); Kiikenthal, Repert. Spec.
Nov. Regni Veg. 53: 187-219 {1994).
Rhizomatous perennials. Culms scapose or few-
noded. Leaves eligulate (?). Inflorescence some-
times pseudolateral, paniculate or capitate with
few-many spikelets. Lower primary bracts
leaflike, sheathing or not, lowermost bract some-
times erect. Spikelets with many spirally arranged
persistent glumes of increasing length, upper
{1- )2 glumes each subtending a flower, glumes
acuminate, top recurved, inner side near the top
densely adpressed-pubescent. Lower flower func-
tionally male, upper flower bisexual. Bristles ab-
sent. Stamens 12-22. Style 8-fid, style base ±
distinct, thickened, persistent. Achene narrowly
obovoid, with a thick rounded beak constricted at
its base, smooth. Two spp., SW and W Australia,
on wet spots in heathland.
79. Caustis R. Br.
Caustis R. Br., Prodr.: 239 (1810); Kiikenthal, Repert. Spec.
Nov. Regni Veg. 53: 85-100 {1944).
Perennials on a short-long creeping rhizome.
Culms many-noded, often strongly sinuous.
Leaves eligulate, blade small, rarely long, inner
side densely adpressed-pubescent. Inflorescence
paniculate with many spikelets. Primary bracts
with reduced blade, sheathing. Partial inflores-
cences sometimes with numerous sterile lateral
branchlets, dense and only slightly curved at first,
often elongating and strongly recurved or coiled at
maturity. Spikelets with few-several spirally ar-
ranged, long persistent glumes of increasing
length, upper 1-2larger glumes each subtending a
flower, glumes acuminate, top recurved, inner
side near the top densely appressed-pubescent.
Lower flower functionally male or absent, upper
flower bisexual. Bristles absent. Stamens 3-6. Style
deeply 3-fid, style base constricted at the base,
thickened, persistent. Achene ovoid to ellipsoid,
rounded and 3-ribbed, mostly beaked, often finely
reticulate. About six spp., Australia, in open forest
or scrub, on dry sandy soil, also at the edge of
streams.
80. Cladium P. Browne
Cladium P. Browne, Civ. Nat. Hist. Jamaica: 114 (1756);
Kiikenthal, Repert. Spec. Nov. Regni Veg. 51: 1-8, 183-192
(1942).
Rhizomatous and stoloniferous perennials. Culms
few-noded. Leaves eligulate, blade flat or revolute.
Inflorescence paniculate with numerous spikelets.
Primary bracts large, leaflike, sheathing. Partial
inflorescences anthelate, corymbose or capitately
contracted. Spikelets with few-many ± spirally
arranged persistent glumes of increasing length,
2-3 upper glumes each subtending a flower. Lower
flower mostly functionally male, upper flower(s)
bisexual. Bristles absent. Stamens 2( -3 ). Style
(2- )3-fid, style base not distinct, thickened, persis-
tent. Achene ovoid, with a thick corky beak, seated
on a broad disklike structure, smooth and shining
to rugulose. About four spp., C. mariscus (L.) Pohl
subcosmopolitan; in swamps and marshes, often
in brackish or calcareous habitats.
81. Rhynchocladium T. Koyama
Rhynchocladium T. Koyama, Mem. N.Y. Bot. Gard. 23: 86
(1972).
Perennial on a suberect rhizome. Culms 1-2-
noded. Leaves eligulate, blade flat or revolute.
Inflorescence paniculate with many spikelets.
Primary bracts long, leaflike, sheathing. Partial in-
florescences ± corymbiform, secondary bracts
short, not sheathing. Spikelets with several disti-
chous, deciduous glumes of increasing length
towards middle glumes, lower glumes sometimes
persistent. Middle and upper 4-6 larger glumes
each subtending a flower. Spikelet (or its upper
part) deciduous as a unit. Lower flower bisexual,
upper flowers male. Bristles 4( -6), somewhat
longer than the fruit, deciduous with the fruit (?).
Stamens 2{-3). Style shortly 3-fid, style base not
distinct, somewhat thickened, persistent. Achene
oblong, trigonous, stipitate, 3-ribbed, long-
beaked, smooth. Only one sp., R. steyermarkii (T.
Koyama) T. Koyama, S Venezuela, in woodland
and savannah.
82. Machaerina Vahl
Machaerina Vahl, Enum. Pl. 2:238 {1805); Kiikenthal, Repert.
Spec. Nov. Regni Veg. 51: 1-17, 139-193 {1942); Raynal,
Adansonia II, 12: 103-112 {1972) (Madagascan spp.).
Baumea Gaudich. (1829).
Creeping rhizomatous or stoloniferous perenni-
als. Culms scapose or few-noded, ancipitous, com-
pressed or (sub)terete, sometimes septate. Leaves
distichous, eligulate, blade ensiform to (sub-)
terete, sometimes septate, rarely reduced to a
 Cyperaceae
sheath. Inflorescence sometimes ± pseudolateral,
paniculate with many spikelets, main axis often
sinuous, partial inflorescences sometimes capi-
tately contracted. Primary bracts often leaflike,
sheathing. Spikelets with 2-10 distichous, long
persistent glumes of increasing length, with (1-)
2-4{5) flowers subtended by the larger glumes.
Lower 1-2 flowers bisexual, upper flower(s) func-
tionally male. Bristles absent, poorly developed, or
up to 6, delicate, deciduous with the fruit. Stamens
3. Style 3-fid, style base not distinct, thickened,
persistent. Achene ovate to oblong, terete to tri-
quetrous, ±3-ribbed to winged, stipitate or sessile,
beaked, smooth to rugose. About 50 spp., Mada-
gascar and the Mascarenes over SE Asia, Malesia,
SE Australia, New Zealand, New Caledonia, Pacific
islands to tropical South America and the West
Indies, in wetlands, sometimes as floating mats, or
in woodlands, often at higher altitudes.
83. Lepidosperma Labill.
Lepidosperma Labill., Nov. Holl. Pl. 1: 14, t. 11 (1805);
Kiikenthal, Repert. Spec. Nov. Regni Veg. 50: 19-50, 112-
128 (1941).
Tufted, rhizomatous, or stoloniferous perennials.
Culms scapose, rarely few-noded, terete, com-
pressed 3-4-angular, or ancipitous. Leaves
(spiro)distichous, ligulate, blade terete, often all
reduced to a sheath. Inflorescence rarely ±
pseudolateral, paniculate or spicately condensed
with few-many spikelets. Primary bracts rather
short, more rarely leaflike, lowermost bract often
suberect. Spikelets with few-several spirally
arranged, persistent glumes of increasing length,
upper 1-2{-4) larger glumes each subtending a
flower. Lower flowers, when present, functionally
male, upper flower bisexual. Scales 6, thin and
bristlelike at anthesis, afterwards basally spongy,
thickening. Stamens 3. Style 3{4)-fid, style base
distinct (at anthesis), thickened, persistent.
Achene (ob)ovate to oblong, rounded to 3-ribbed,
crowned by a mostly thick and obtuse apex, rarely
with a long beak, smooth or finely reticulate.
About 55 spp., SE Asia and Malesia (1) towards
Australia (ca. 50), New Zealand (3), and New
Caledonia {4), along rivers and in woodland,
rarely in mountain heath vegetation.
The flower of L. forsythii A.A. Hamilton has only
3 scales, which remain thin, but remarkably has
the 3 filaments inflated at the base and remaining
attached to the fruit (Wilson, Telopea 5: 608.
1994), mimicking the real thing.
181
84. Tricostularia Nees
Tricostularia Nees in Lehmann, Nov. Stirp. Pug. 8: 50 (1844);
Kiikenthal, Repert. Spec. Nov. Regni Veg. 53: 212-217
(1944).
Tufted, rhizomatous perennials. Culms scapose or
few-noded. Leaves eligulate, sometimes reduced
to a sheath. Inflorescence paniculate with few to
many spikelets. Primary bracts rather stiff and
short, sheathing. Spikelets with several distichous,
deciduous glumes of increasing length, upper 1-3
larger glumes each subtending a flower. Lower
flower(s) functionally male, rarely absent, upper
flower bisexual. Bristles {3-)6, sublaminar, basally
adnate into a small cuplike structure, (much)
shorter than the fruit, deciduous with the fruit.
Stamens 3. Style 3-fid, style base± distinct, not or
slightly thickened, deciduous, rarely persistent.
Achene obovoid, ±stipitate, ±3-ribbed, rarely
beaked, surface hairy. Six spp., Sri Lanka, SE Asia,
and Malesia (1), to Australia {5) and New
Caledonia (2), in open heath or scrubland, on
humid sandy soils.
III. Subfamily Sclerioideae C.B. Clarke (1908).
10. Tribe Cryptangieae Bentham (1881).
85. Lagenocarpus Nees
Lagenocarpus Nees, Linnaea 9: 304 (1834); Koyama &
Maguire, Mem. N.Y. Bot. Gard. 12:35-51 (1965) (part. rev.).
Cryptangium Schrader ex Nees (1842).
Rhizomatous or stoloniferous perennials. Culms
scapose or several-many-noded. Leaves eligulate,
contraligule ± triangular, lower leaves sometimes
reduced to a sheath. Inflorescence paniculate with
many-numerous spikelets. Primary bracts leaf-
like, sheathing. Partial inflorescences anthelate or
± contracted. Lower spikelets (or lower partial
inflorescences) male, upper female, or flowers dio-
ecious. Male spikelets with many densely spirally
arranged, persistent glumes, middle and upper
glumes subtending a male flower. Male flowers
with 1-6 stamens. Female spikelets with few-
several spirally arranged, persistent glumes,
penultimate glume subtending a female flower.
Female flower surrounded at the base, opposite
the sides, by 3 small ciliate scales. Style 3-fid, style
base distinct or not, thickened or not, mostly per-
sistent. Achene ovate to obovate, rounded
trigonous, often 3-ribbed, ±beaked, smooth or
foveolate, rarely scabrid. About 30 spp., tropical
182
Central and S America, West Indies, in dry to wet
savannah, often on sandy soil or rocky slopes.
86. Didymiandrum Gilly
Didymiandrum Gillay, Bull. Torrey Bot. Club 68: 330 (1941);
Koyama & Maguire, Mem. N.Y. Bot. Gard. 12: 51-52 (1965).
Tal rhizomatous perennial. Culms many-noded,
woody. Lower leaves reduced to a sheath, middle
and upper leaves pseudoverticillate in 2 s or 3 s,
sheathing, eligulate. Inflorescence paniculate with
many spikelets. Primary bracts leaflike, sheathing.
Flowers dioecious. Male spikelets with many
densely spirally arranged, persistent glumes, 1-
few middle or apical glumes each subtending a
male flower. Male flowers with 3 stamens. Female
spikelets with many densely spirally arranged,
persistent glumes, penultimate glume subtending
a female flower. Female flower surrounded at the
base, opposite the sides, by 3 small, fimbriolate
scales, deciduous with the fruit. Style 3-fid, style
base :±: distinct, thickened, persistent. Achene
obpyriform to obovate, trigonous, 3-ribbed, beak
pyramidal, smooth. Only one sp., D. stellatum
(Boeckeler) Gilly, N S America, in mountain forest
and woodland.
87. Everardia Ridley ex Thurn
Everardia Ridley ex Thurn, Timehri 5: 210 (1886); Koyama &
Maguire, Mem. N.Y. Bot. Gard. 12: 20-35 (1965).
Pseudoeverardia Gilly (1951).
Dwarf shrubs with woody caudex covered by
fibrous leaf sheath remnants and adventitious
roots. Culms lateral, scapose, compressed. Leaves
crowded on top of the caudex, eligulate, con-
traligule :±: triangular. Inflorescence paniculate
with many-numerous spikelets. Primary bracts :±:
leaflike, sheathing. Lower branches (or lower par-
tial inflorescences) usually with male spikelets,
upper with female spikelets, rarely dioecious.
Male spikelet with spiral, persistent glumes, lower
empty, middle and upper each subtending a male
flower with 2-9 stamens. Female spikelet with few
spiral, persistent glumes, penultimate glume sub-
tending a female flower with a pistil, surrounded
at the base, opposite the sides, by 3 small :±: trian-
gular, long-fimbriate scales. Style 3-fid, base not
distinct, not or slightly thickened, persistent.
Achene oblong to obovate, rounded trigonous, 3-
ribbed, often long-beaked, beak smooth or
scabrid, surface smooth. About 12 spp., N South
America, terrestrial or epiphytic, on bare sand-
stone rocks, escarpments, in crevices.
Cyperaceae
88. Cephalocarpus Nees Fig. 42D-F
Cephalocarpus Nees in Mart., Fl. Brasil. 2(1): 162, t.l8 (1842);
Koyama & Maguire, Mem. N.Y. Bot. Gar d. 12: 17-19 ( 1965).
Tufted or isolated caudex-forming dwarfshrubs,
caudex covered by decaying leaf sheaths and
adventitious roots. Culms lateral, scapose, com-
pressed. Leaves crowded on top of the caudex,
eligulate. Inflorescences contracted paniculate or
capitate with few-several spikelets. Primary bracts
small, sheathing, suberect. Inflorescences uni-
sexual or bisexual, flowers often dioecious. Male
spikelets with spiral, persistent glumes, lower and
upper glumes smaller and empty, middle 5-9 larg-
est each subtending a mostly 2-staminate flower.
Female spikelet with few spiral, persistent glumes,
penultimate glume subtending a flower (rarely a
2nd upper and nonfunctional flower present).
Female flower with a pistil, surrounded at the base
by 3 small, fimbriate scales. Style 3-fid, style base
distinct, partly thickened and persistent. Achene
(ob)ovate, terete to subtrigonous, :±:3-ribbed, per-
sistent style base finally clavate, smooth. About
three spp., N South America, terrestrial or epi-
phytic, in open vegetation or on rocks in lowland
forests.
11. Tribe Trilepideae Goetghebeur (1985).
89. Trilepis Nees
Trilepis Nees, Linnaea 9: 305 (1834); Koyama & Maguire,
Mem. N.Y. Bot. Gard. 12: 14-17 (1965).
Fintelmannia Kunth (1837).
Tufted perennials or dwarf shrubs with a
(branched) caudex, covered by :±: fibrous leaf
sheath remnants and adventitious roots. Culms
lateral, scapose, compressed. Leaves crowded on
top of the caudex, eligulate, contraligula triangu-
lar. Inflorescence paniculate with few-many
spikeletlike spikes. Primary bracts :±: leaflike,
sheathing. Spikes with numerous lateral spikelets,
subtended by spirally arranged glumelike bracts.
Basal spikes male, apical spikes female. Spikelets
with few distichous, persistent glumes, 1-2 each
subtending a flower, lateral spikelets with an
irregularly shaped prophyll. Male flower with 3
stamens. Female flower with a pistil, surrounded
at the base and opposite the sides by 3 small, long
fimbriate hypogynous scales, deciduous with the
fruit. Style 3-fid, style base not distinct, tapering,
persistent. Achene fusiform, trigonous, long-
beaked, smooth. About three spp., Guayana and
SE Brazil, on bare, granitic rocks.
 Cyperaceae 183

90. Afrotrilepis (Gilly) Raynal tapering, persistent. Achene fusiform, trigonous,

long-beaked, surface smooth. About seven spp.,
Afrotrilepis (Gilly) Raynal, Adansonia II, 3: 258 (1963); Haines
& Lye, Bot. Not. 126: 330-339 (1973).
W Cameroon, W Angola, E and S Africa, Madagas-
car; on hot, temporarily dry, bare rocks.
Mat-forming perennials with branched rhizomes,
forming a short caudex. Culms scapose. Leaves ::!::: 92. Microdracoides Hua Fig. 43
crowded on top of a caudex, leaf sheaths closed
but with a conspicuous longitudinal fold, ligule Microdracoides Hua, Bull. Mus. Natl. Hist. Nat. 12: 421 (1906);
Chermezon, Bull. Soc. Bot. Fr. 80: 90-97 (1933).
fimbriate, contraligule triangular. Inflorescence
paniculate with few-many spikel~tlike s~ikes. P.ri-
Medium-sized dwarfshrub with caudex covered
mary bracts ::!::: leaflike, sheathmg. Spikes .with
by leaf sheaths and adventitious roots. Culms lat-
numerous lateral spikelets, subtended by spually
eral, scapose. Leaves crowded on top of the
arranged glumelike bracts. Spikes usually
caudex, eligulate (?), blade deciduous. Inflores-
bisexual, rarely male. Spikelets unisexual or
cence paniculate with many spi~eletlik~ spik~s.
bisexual, with few distichous, persistent glumes,
Primary bracts ::!::: leaflike, sheathmg. Sp1kes with
the larger 1-2 each subtending a flower, lateral
numerous lateral spikelets, subtended by small,
spikelets with an irregularly shaped . prop~yll.
densely spirally arranged glumelike bracts. Spikes
Basal spikelets with 1-2 male flowers, apical spike-
unisexual, flowers dioecious. Spikelets unisexual,
lets mostly with 1 male and 1 female flower, rarely
with few distichous, persistent glumes, 1-2 larger
completely male or female. Male flower with 1-3
each subtending a flower, lateral spikelets with an
stamens. Female flower with a pistil, surrounded
irregularly shaped prophyll. Male flower 3-
at the base and opposite the sides by 3 hypogynous
staminate. Female flower with a pistil, surrounded
scales, deciduous with the fruit. Style 3-fid, style
at the base and opposite the sides by 3 hypogynous
base not distinct, tapering, persistent. Achene
scales deciduous with the fruit. Style 3-fid, style
fusiform, trigonous, long-beaked, smooth. Two
base not distinct, tapering, persistent. Achene
spp., tropical W Africa, from Senegal to Gabon; on
fusiform, trigonous, long-beaked, surface smooth.
bare granitic rocks.
Only one sp., M. squamosus Hua, scattered in
Guinee, Sierra Leone, Nigeria, Cameroun, on bare
91. Coleochloa Gilly rocks.

Coleochloa Gilly, Brittonia 5: 12 (1943); Nelmes, Kew Bull.

1953: 373- 381 (1953); Haines & Lye, Bot. Not. 126: 330-339
(1973) (rev. E Afr.) .
Eriospora Hochst. ex A. Rich. (1850), nom. illeg., non Berkeley
& Broome (1850).

Tufted or cushion-forming perennials with exten-

sively branched rhizomes, sometimes forming a
small caudex. Culms (sub )seapose, compressed.
Leaves distichous, leaf sheath open, ligule
fimbriate, blade flat or inrolled, deciduous. Inflo-
rescence paniculate with few-many spikeletlike
spikes. Primary bracts ::!::: leaflike. Spikes .with
numerous lateral spikelets subtended by sp1rally
arranged glumelike bracts. Spikes bisexual. Spike-
lets unisexual or bisexual, with few distichous,
persistent glumes, the larger 1-2 each subtending
a flower, lateral spikelets with an irregularly
shaped prophyll. Basal spikelets with 1-2 male
flowers, apical spikelets mostly with 1 male and 1
female flower, more rarely completely female.
Male flower with 2-3 stamens. Female flower with
Fig. 43A-E. Cyperaceae. Microdracoides squam.osus. A .stems
a pistil, surrounded at the base and oppos.ite the with terminal leaf rosettes. B Tip of branch with pamcle. C
sides by 3 small hypogynous scales, dec~d~ous Pistillate spikelet. D Fruit subtended by hypogynous scales.
with the fruit. Style 3-fid, style base not distmct, E Hypogynous scale. (Takhtajan 1982)
184 Cyperaceae
12. Tribe Sclerieae Kunth ex Fenzl (1836).
93. Scleria Bergius
Scleria Bergius, K. Vetensk. Acad. Hand!. 26: 142, t.4-5 (1765);
Core, Brittonia 2: 1-105 (1936) (rev. Amer.); Robinson, Kew
Bull. 18: 487-551 (1966) (rev. SE Afr.).
Acriulus Ridley (1884).
Small annuals or medium-sized to tall tufted,
rhizomatous, or even scrambling perennials.
Culms mostly many-noded. Leaves sometimes
pseudoverticillate, sheaths sometimes conspicu-
ously winged, (hairy) ligule rarely present, con-
traligule often present, sometimes as a large
outgrowth, blade rarely lanceolate or pseudo-
petiolate, sometimes pseudopremorse. Inflores-
cence polymorphic, basically paniculate, but often
with contracted partial inflorescences. Spikelets
with few-many distichous glumes, lower glumes
empty, larger glume subtending a female flower,
or a female (lower) and 1-few upper glumes each
a male flower, or male flowers only, glume length
increasing towards the female flower. Male and
female spikelets intermingled, or female spikelets
only basal, rarely plants dioecious. Male flower
with 3 stamens. Female flower with 3-fid style,
style base mostly not distinct, thickened or not,
persistent or deciduous, ovary surrounded at the
base by a variously shaped (often 3-lobed, margin
entire, ciliate or serrate) and sometimes reduced
hypogynium, lobes - when present - opposite the
ribs of the fruit, deciduous with the fruit. Achene
globose to ovoid, rounded to trigonous, smooth or
variously sculptured, glabrous or hairy, wall often
bony hard, white or discolored, sometimes
beaked. About 250 spp., pantropical, locally ex-
tending to (warm-)temperate regions; most spp.
in seasonally to permanently damp to waterlogged
places, often on sandy soils, many annuals on sea-
sonally damp shallow soils overlying rocks, more
rarely growing in open to dense woodland.
13. Tribe Bisboeckelereae Pax ex L.T. Eiten
(1976).
94. Becquerelia Brongn.
Becquerelia Brongn. in Duperrey, Voy. Monde, Bot. 2: 161, t.
27 (1833); Koyama, Mem. N.Y. Bot. Gard. 17: 25-32 (1967).
Tufted or shortly rhizomatous, rarely stolonifer-
ous perennials. Culm scapose or not. Leaves
sometimes all basal and then often pseudo-
petiolate, eligulate, leaf blade sometimes lan-
ceolate. Inflorescence paniculate with many-
numerous spikelets. Primary bracts leaflike,
sheathing. Partial inflorescences usually corym-
bose, rarely capitately contracted, with several-
many lateral male or female spikelets, branches
terminated by a female spikelet. Male spikelet
prophyllate, with ca. 5 distichous, persistent
glumes, the lower each subtending a monandrous
male flower. Female spikelet with ca. 10 empty,
persistent glumes surrounding a female flower in
(pseudo?)terminal position. Female flower with
a deeply 3-fid style, style base not distinct, not
thickened, deciduous, base surrounded by a
subtrigonous disc. Achene (depressed) globose,
smooth, rugulose, or tuberculate. Five spp., tropi-
cal Central and S America, in forest or at the edge
of swamps in savannah.
95. Diplacrum R. Br.
Diplacrum R. Br., Prodr.: 240 (1810); Koyama, Mem. N.Y. Bot.
Gard. 17: 32-35 (1967) (Amer. spp.).
Pteroscleria Nees (1842).
Annuals or tufted, rarely stoloniferous perennials.
Culms scapose, often short. Leaves eligulate, blade
sometimes lanceolate. Inflorescence paniculate
or capitate with many spikelets. Primary bracts
leaflike, sheathing. Partial inflorescences anthelate
or capitately contracted, lateral spikelets usually
male, terminal spikelets usually female. Male
spikelets with few distichous, persistent glumes,
each subtending a monandrous male flower. Fe-
male spikelet with 2 distichous, persistent glumes,
sometimes deciduous with the fruit, surrounding
a (pseudo?)terminal female flower with a pistil
seated on a basal trilobed disc, lobes opposite
the 3 main ribs, style 3-fid, style base not distinct,
not thickened, deciduous. Achene subglobose to
ovoid, inconspicuously 3-ribbed, beak short,
smooth, ribbed, or reticulate. Seven spp., pan-
tropical, in forest, often along streams or near
waterfalls, or on damp places in savanna, often on
white sand.
96. Bisboeckelera Kuntze
Bisboeckelera Kuntze, Rev. Gen. Pl. 2: 747 (1891); Koyama,
Mem. N.Y. Bot. Gard. 17: 35-39 (1967).
Tufted, rarely shortly rhizomatous perennials.
Culms central or lateral, scapose. Leaves eligulate,
blade often oblong and pseudopetiolate, or leaves
reduced to a sheath. Inflorescence capitate or
anthelate with few-many spikelets. Primary bracts
leaflike or short, (shortly) sheathing. Partial inflo-
 Cyperaceae
rescences capitately contracted, male spikelets 2-
3, lateral, terminal spikelets usually female. Spikes
bisexual, with 2 bracts subtending 1-2 male spike-
lets and terminated by a female spikelet. Male
spikelet with several distichous, persistent glumes,
1-2 subtending a monandrous male flower.
Female spikelet only a (pseudo?)terminal female
flower, tightly surrounded by a many-nerved
utriculiform glume. Female flower with a pistil,
seated on a basal trilobed disc, lobes opposite the
3 main ribs, style 3-fid, style base not distinct,
deciduous to ± persistent. Achene obovate to
cordate, trigonous to triquetrous, beaked, sur-
face smooth. Four spp., N South America, in
forest.
97. Calyptrocarya Nees
Calyptrocarya Nees, Linnaea 9: 304 (1834); Koyama, Mem.
N.Y. Bot. Gard. 17:40-46 (1967); Davidse & Kral, Ann. Mo.
Bot. Garden 75: 853-861 (1988).
Rhizomatous or stoloniferous perennials. Culms
central or lateral, (sub)scapose, trigonous. Leaves
eligulate, sometimes with a triangular con-
traligule, blade sometimes lanceolate. Inflores-
cence paniculate or capitate with few-many
spikelets. Primary bracts leaflike, sheathing.
Partial inflorescences ± corymbose to capitately
contracted, with many lateral bisexual or func-
tionally male or female spikelets, terminal spike-
lets male or female. Male spikelet with a few
distichous, persistent glumes, each subtending a
monandrous male flower. Functionally female
spike with 3 tristichous bracts, each subtending a
neuter spikelet with a few empty distichous, per-
sistent glumes, these spikelets surrounding a fe-
male spikelet, reduced to a (pseudo?)terminal
female flower, very tightly clasped by a nerveless
utriculiform glume. Female flower with a pistil,
seated on a basal inconspicuous disc, style
deeply 2-3-fid, style base not distinct, not thick-
ened, deciduous. Achene obovate, thick lenticular
to rounded trigonous, beak short, surface smooth.
Eight spp., tropical Central and S America, in
forest.
Incertae sedis
98. Exochogyne C.B. Clarke
Exochogyne C.B. Clarke, Verh. Bot. Vereins Prov.
Brandenburg 47: 101 (1905); Koyama & Maguire, Mem. N.Y.
Bot. Gard. 12: 52-54 (1965).
185
Perennial with short creeping rhizome. Culms sea-
pose. Leaves eligulate, contraligule rounded
triangular. Inflorescence paniculate with many
spikelets, main axis conspicuously sinuous. Pri-
mary bracts boat -shaped, shortly sheathing, tightly
clasping the partial inflorescences (only part of the
spikelets visible). Lower (=long pedicelled) spike-
lets female, upper (=shortly pedicelled) spikelets
male. Male spikelet with several distichous, persis-
tent glumes, middle and upper glumes subtending
male flowers with 2 (up to 6?) stamens. Female
spikelet with several distichous, persistent glumes,
penultimate glume (rarely the lower glume also)
subtending a female flower with a pistil lacking
hypogynous scales. Style 2-fid, style base distinct,
thickened, persistent. Achene (ob )ovate, laterally
compressed, 2-ribbed, shortly stipitate, beaked,
surface smooth. Only one sp., E. amazonica C.B.
Clarke, scattered inN South America and SE Brazil,
in savanna or scrub.
Usually, this genus is considered a member of
the Cryptangieae. However, some disturbing dif-
ferences such as the slender habit, the distichous
glumes, the laterally compressed bicarpellate
ovary, and the lack of a perianth are out of order
in that tribe (Goetghebeur, Gen. Cyp.: 937-938,
1986). Lacking further evidence, I provisionally
consider this and the following genus as incertae
sedis.
99. Koyamaea W.W. Thomas & Davidse
Koyamaea W.W. Thomas & Davidse, Syst. Bot. 14: 189 (1989).
Loosely tufted to shortly rhizomatous perennial.
Culm lateral, few-many-noded, velvety hairy.
Leaves with a ciliate ligule, contraligule shortly
triangular, hairy, lower leaves reduced to a sheath.
Inflorescence capitate with few-several spikelets.
Lower primary bracts leaflike, sheathing. Spikelets
densely pilose, with many spirally arranged, per-
sistent (?) glumes, several empty basal glumes,
1 female flower, many male flowers, and a few
empty upper glumes, top cucullate. Male flower
3-staminate, some of the lower male flowers with
1-2 bristles. Female flower with 2-5 hypogynous
bristles, shorter than the fruit, probably deciduous
with the fruit. Style deeply 3-fid, style base not
distinct, thickened, persistent. Achene elliptic,
rounded triangular, 3-ribbed, beaked, smooth.
Only one sp., K. neblinensis W.W. Thomas &
Davidse, endemic on the Cerro de la Neblina (SE
Venezuela and adjacent Brazil), in dry scrub or in
more mesic savannah.
186
IV. Subfamily Caricoideae Pax (1886).
14. Tribe Cariceae Kunth ex Dumort. (1827).
100. Kobresia Willd.
Kobresia Willd., Sp. Pl. 4:205 (1805); Kiikenthal, Pflanzenreich
IV. 20: 33-50 (1909); Rajbhandari & Ohba in Ohba & Malia,
Himalayan Pl. 2: 117-167 (1991) (rev. Nepal).
Elyna Schrader (1806).
Tufted or shortly rhizomatous perennials. Culms
mostly scapose. Leaves ligulate, blade often
reduced. Inflorescence paniculate or spicate, with
few-many spikelets, cladoprophylls not utriculi-
form, empty. Primary bracts leaflike or short.
Ultimate partial inflorescences spikelike, bisexual
or unisexual, with few-many spirally arranged
deciduous or persistent bracts (or persistent
glumes), either subtending (proximally) a
bisexual spikelet or a female spikelet, or (distally)
a male flower. Bisexual spikelet with a basal ±
utriculiform prophyll, subtending a female flower
and a rachilla bearing at the top few-several
glumes, each subtending a male flower. Female
spikelet reduced to a (more to less open)
utriculiform flower-bearing prophyll, enclosing
the ± well-developed, often laminar rachilla,
ovary and style, only style branches and often the
rachilla protruding. Male flower 3-staminate,
without bristles. Female flower with a 3-fid style
(2-fid in K. macrantha Boeckeler), style base not
distinct, not or slightly thickened, persistent.
Achene (ob )ovate to oblong, trigonous, rarely
biconvex, rarely stipitate, ±beaked, surface
smooth. About 50 spp., holarctic region with a
concentration in the Himalayas, K. kobresioidea
(Kiik.) Kern on the mountains of N Sumatra. In
open, alpine vegetation, rarely in shady or moist
places.
101. Schoenoxiphium Nees
Schoenoxiphium Nees, Linnaea 7: 531 (1832); Levyns, ].S. Afr.
Bot. 11: 79-89 (1945); Kukkonen, Bothalia 14: 819-823
(1983).
Tufted or (shortly) rhizomatous perennials.
Culms scapose. Leaves ligulate. Inflorescence
open to condensed paniculate or spicate, with
few-many spikelets, cladoprophylls sometimes ±
utriculiform, swollen at the base, and subtending a
female flower. Primary bracts leaflike or short,
sheathing or not. Ultimate partial inflorescences
spikelike, bisexual or unisexual, with few-many
spirally arranged deciduous or persistent bracts
Cyperaceae
(or glumes), either subtending (proximally) a
bisexual spikelet or a female spikelet, or (distally)
a male flower. Bisexual spikelet with a basal ±
utriculiform prophyll, subtending a female flower,
and a rachilla bearing at the top few-several
glumes, each subtending a male flower. Female
spikelet reduced to a ± open utriculiform flower-
bearing prophyll, enclosing the well-developed,
often laminar rachilla, ovary and style, only style
branches and often the rachilla protruding. Male
flower 3-staminate, without bristles. Female
flower with a 3-fid style, style base not distinct, not
or slightly thickened, persistent. Achene oblong or
(ob )ovate, trigonous, often beaked, smooth.
About 12 spp., S Africa, on the E African moun-
tains (2), and in Madagascar (2), in humid grass-
land and dense to open forest.
102. Uncinia Pers.
Uncinia Pers., Syn. Pl. 2: 534 (1807); Kiikenthal, Pflanzenreich
IV. 20: 50-67 (1909); Hamlin, Rec. Domin. Mus. 3: 85-88
(1958) (infragen. class.); Hamlin, Domin. Mus. Bull. 19: 106
p. (1959) (rev. New Zealand); Kukkonen, Kew Bull. 21: 93-
97 (1967) (structure of spikelet).
Tufted, rhizomatous, or rarely stoloniferous pe-
rennials. Culms scapose, rarely noded. Leaves
ligulate. Inflorescence a bisexual spike with in the
lower part female spikelets, and apically male
flowers, forming a terminal male spikelet. Primary
bracts mostly small and glumelike, or a few lower
bracts ± leaflike, deciduous or persistent. Male
spikelet with few-many densely spirally arranged
glumes, each subtending a male flower with 2-3
stamens with broad or narrow filaments. Female
spikelets 1-flowered, rachilla elongated, with
a terminal recurved hook, formed by a thick,
±coriaceous glume, prophyll utriculiform, sub-
tending the female flower. Female flower without
bristles, with a 3-fid style, style base not distinct,
often ± thickened and persistent, ovary seated on
a small disk with 3 basal knobs, opposite the ribs.
Achene oblong to ovate, (rounded) trigonous,
beaked, surface smooth. About 60 spp., Malesia
over Australia, New Zealand, S Pacific islands
(plus Hawaii) to Central and S America, in humid
grassland and temperate forest, often at higher
altitudes.
Uncinia kingii Boott was long considered an
oddity in Uncinia. Many abnormal character
states came to light, and, as a result, the species
was quite correctly transferred to Carex as C.
kingii (Boott) Reznicek.
 Cyperaceae 187

103. Carex L. Fig. 44

Carex L., Spl. Pl. 2: 972 (1753); Ktikenthal, Pflanzenreich IV.
20: 67-767 (1909); Smith & Faulkner, Bot. Rev. 42: 53-81
(1976); Robertson, Taxon 28: 535-548 (1979); Reznicek,
Can. J. Bot. 68: 1409-1432 (1990).
Vignea P. Beauv. ex Lestib. (1819).
Vesicarex Steyerm. (1951).

Tufted, rhizomatous, stoloniferous, or tussock-

forming perennials. Culms central or lateral,
mostly scapose, rarely noded. Leaves ligulate.
Inflorescence rarely pseudolateral, paniculate,
often partly or completely contracted, rarely
corymbose or anthelate with few-numerous
spikes, less frequently reduced to a single spike,
cladoprophylls sometimes swollen at the base,
utriculiform, and subtending a female flower.
Inflorescence mostly bisexual, rarely unisexual or
flowers dioecious. Primary bracts leaflike or not,
sheathing or not. Partial inflorescences spikelike,
unisexual or bisexual, with few-many spirally
arranged bracts (or glumes), subtending either a
female spikelet or a male flower. Spikes male,
female, or bisexual, and then mostly male flowers
apically, rarely basally, or intermingled. Male
flower (2- )3-staminate, without bristles. Female
spikelet reduced to the (mostly underdeveloped to
virtually absent) rachilla and a utriculiform,
flower-bearing prophyll, completely enclosing the
rachilla, ovary, and style, only the style branches
protruding and very rarely also the well-devel-
oped rachilla (very rarely bearing 1-few reduced
glumes). Female flower with a 2-3-fid style, style
base not distinct, rarely thickened, persistent.
Achene often obovate, trigonous, or dorsiven-
trally compressed, sometimes remarkably dis-
formed. Perhaps ca. 2000 spp., cosmopolitan, with
concentrations in some cold holarctic regions.
The species of this giant genus occupy a diversity
of habitats, but most species grow under wet, cold
conditions in bogs, alpine and arctic vegetations.
Forest and thermophilic species are much rarer,
there are also a few species inhabiting disturbed Fig. 44A-J. Cyperaceae. A Carex physodes. Fruiting plant,
grounds, sometimes becoming weedy. note inflated utricles. B-E C. dioica. B Habit. C Pistillate spike.
The monographer Kiikenthal (op.cit.) divided D Staminate spike. E Utricle at fruiting stage. F-1 C. vulpina. F
Inflorescence. G Staminate flower. H Pistillate flower, utricle
the genus into four subgenera. His subgenus
opened to show pistil. I Utricle at fruit maturity. J Achene.
Primocarex (based on the single character state of (Takhtajan 1982)
unispicate inflorescences) is now widely consid-
ered as a polyphyletic assemblage of species with
phylogenetic affinities to highly diverse groups in cence bracts mostly leaflike, often sheathing,
the other three subgenera or even out of Carex. In cladoprophylls present, spike prophylls never
a recent review by Reznicek (op.cit.), the three utriculiform, pistils 3-, rarely 2-carpellate; subgen.
remaining subgenera Carex, Vignea, and Indo- Vignea: inflorescence bracts rarely leaflike, never
carex are recognized as such, and (here succinctly) sheathing, cladoprophylls absent, spike prophylls
characterized as follows. Subgen. Carex: inflores- absent, if present never utriculiform, pistil 2-,
188 Cyperaceae
rarely 3-carpellate; subgen. Indocarex: inflores-
cence bracts mostly leaflike and sheathing,
cladoprophylls present, spike prophylls often
utriculiform and sometimes subtending a female
flower, pistils 3-carpellate.
104. Cymophyllus Mackenzie
Cymophyllus Mackenzie ex Britton in Britton & Brown, Ill. Fl.
N. U.S., ed. 2, 1: 441 (1913); Kartesz & Gandhi, Rhodora 93:
136-140 (1991).
Tufted perennial on an oblique rhizome. Culms
lateral, (sub)scapose. Lower 4-5 leaves reduced
to an open sheath with overlapping margins,
eligulate, uppermost (?) leaf sheathless (?),
eligulate, blade (or = expanded sheath?) large and
wide, with undulate margins. Inflorescence a
single spike with many spirally arranged small
whitish bracts, lower bracts each subtending a
female spikelet, upper bracts each subtending
a 3-staminate male flower. Female spikelet with
a rachilla, ca. half as long as the fruit, and a large,
whitish, inflated utriculiform prophyll, sub-
tending a female flower. Female flower with a
3-fid style, stigmatic branches conspicuously
thickened-clavate, style base not distinct, not
thickened, deciduous, ovary seated on a disk
with 3 basal knobs, opposite the ribs. Achene :±
obovate, trigonous, smooth. Only one sp., C.
fraserianus (Ker Gawler) Kartesz & Gandhi,
Appalachian Mts., SE USA, in forest.
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190 Cyperaceae/Dasypogonaceae

anatomy and carbon isotope ratios. Ann. Bot. II, 64: 425-
438.
Dasypogonaceae
Van der Veken, P. 1965. Contribution a l'embryographie
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(eds.) Flowering plants in Australia. Adelaide: Rigby, pp.
364-368. Dasypogonaceae Dumort., Anal. Fam. Pl.: 54-55 (1892).
Kingiaceae End!. (1838).
Calectasiaceae End!. (1838).
Baxteriaceae Takhtajan (1994).

Low to tall perennials, shrubby or arborescent,

with condensed to tall stem, the stem sometimes
(Kingia) with a cover of aerial roots penetrating
the persistent leaf bases; leaves spirally inserted,
dorsiventral, more or less V- or U-shaped in
transection. Inflorescences solitary or fasciculated
globular heads on bracteate peduncles, or flowers
solitary and subtended by scarious bracts; flowers
hermaphrodite; perianth indurated, persistent;
tepals variously free or united, the 2 whorls
similar, cream white, reddish brown or purple to
blue. Stamens 6; usually attached to base of tepals;
anthers tetrasporangiate, basifixed or rarely dor-
sifixed, dehiscing introrsely or rarely extrorsely by
slits or 2 pores. Ovary syncarpous, 3-locular, each
locule with 1 ovule arising from axile placenta, or
(Calectasia) 1-locular with 3 basally attached
ovules; style single, with capitate or 3-lobed
stigma; ovules anatropous, bitegmic and cras-
sinucellar;fruit indehiscent, enclosed in persistent
perianth, or rarely capsular and explosively dehis-
cent (Baxteria); seeds subspherical with pale yel-
low testa, enclosed inside ovary and perianth
parts; endosperm copious, without starch; embryo
broad, resticted to the lower part of the seed.
An endemic Australian family of four genera
and eight species, restricted to S and SW Australia.

VEGETATIVE MORPHOLOGY. The plants are either

rhizomatous with single or branched aerial stems
or have a single erect, often massive stem with a
terminal leaf rosette. In Kingia adventitious roots
arise a few em below the stem apex and grow
downards penetrating the persistent leaf bases, as
in the Velloziaceae or Bromeliaceae. The leaves
are linear or strap-shaped and range from a few
mm to 1m in length. In cross-section the leaf
blades are flat, circular, triangular or rhomboid in
outline. The lamina may contract slightly towards
the base but never develop a petiole; a leaf sheath
is always well developed.
 Dasypogonaceae 191

VEGETATIVE ANATOMY. The roots have vessels

with simple (Dasypogon, Calectasia) or scalari-
form (Kingia, Baxteria) perforation plates. In the
stem and leaves, vessels are lacking in all genera.
The massive rhizomes and aerial stems of Kingia
and some Dasypogon are not due to (anomalous)
secondary growth, as in the habitually similar
Xanthorrhoea, but result from extensive primary
thickening processes (Fahn 1954b, 1961; Wagner
1977; Staff and Waterhouse 1981; Rudall and
Chase 1996).
The leaves have stomata on both surfaces or sto-
mata may be restricted to the abaxial surfaces
only. In Kingia and Baxteria, the stomata are of
the common paracytic/tetracytic type, while in
Dasypogon and Calectasia several cells surround
the stomata. Baxteria resembles Kingia in having
both phloem fibres and a larger region of thinner-
walled fibrous cells present within the outer
bundle sheaths, but differs in having sclerenchy-
matous girders extending from the outer bundle- D
sheath cells to the epidermis. All genera often have
2 peripheral phloem strands at the vascular
Fig. 45A-D. Dasypogonaceae. Kingia australis. A Habit. B
bundles. Dasypogon differs from the other genera Globular head on bracteate peduncle. C Flower. D Stamen.
in having chlorenchyma present only on the (Takhtajan 1982)
abaxial side of the leaf, while the vascular bundles
are embedded in parenchymatous tissue at the
adaxial side. Dasypogon and Calectasia resemble like, membranous or fleshy, but not petaloid. In
each other in lacking girders, but the leaf of Kingia the abaxial surface is densely pilose. In
Calectasia is reduced and has only about 3 vascu- Calectasia the tepals are indurated at maturity.
lar bundles. Calcium oxalate raphides are known The number of veins per tepal varies from 1 to
only from the flowers but no other organs of about 9. The tepals vary in length from 5-60mm.
Dasypogon or Calectasia, while in Baxteria and The 6 stamens are usually adnate to the bases of
Kingia they are completely lacking. Spherical, the tepals. The anthers are 4-sporangiate and 2-
druselike silica boides are present in the leaf epi- locular, basifixed or dorsifixed (Dasypogon) and
dermis, usually 1 per cell. In Kingia and Baxteria, introrse with longitudinal or apical porate
these bodies were mistaken as clustered calcium (Calectasia, Fig. 46I) dehiscence. The anthers may
oxalate crystals by Fahn (1954a). Fine silica sand be up to 25 mm long. Calectasia cyanea has
or amorphous silica crystals are also present in the diporate anthers. The ovary is superior, syncar-
epidermal cells of Dasypogon and Calectasia, in pous and 3-locular with 1-2 ovules per locule,
contrast to Xanthorrhoea and Lomandra, where except in Calectasia, which has a 1-locular ovary
silica is replaced by styloidlike calcium oxalate with 3 basal ovules. Septal nectaries are present as
crystals. shallow basal grooves in at least Dasypogon and
Kingia, but are absent in Calectasia.
INFLORESCENCE AND FLORAL STRUCTURE. The
globular heads of Kingia (Fig. 45) and Dasypogon EMBRYOLOGY. The ovules of Dasypogon, Calec-
(Fig. 46) probably comprise cymule units. In tasia and Kingia have a small embryo sac and a
Calectasia and Baxteria the flowers are solitary large starch-filled chalazosperm. This chalaza-
and terminal, although the multiple subtending sperm is subdermal and lacks central axially-
bracts for each flower suggest reduction from a oriented cells. Both integuments form the
more complex inflorescence type. The 3 + 3 tepals micropyle (Rudall 1994); these features setting
are persistent and similar in both whorls. The them apart from Asparagales.
tepals are either more or less free (Kingia), or
united basally (Dasypogon), or into of floral tube POLLEN MORPHOLOGY. The pollen grains in
(Calectasia, Fig. 46H). The tepals are parchment- Dasypogon and Calectasia are sulcate and micro-
192 Dasypogonaceae
Fig. 46A-K. Dasypogonaceae. A-E Dasypogon bromeliifolius.
A Habit. B Flower. C Segment of outer perianth whorl. D
Segment of inner perianth whorl. E Stamen. F-K Calectasia
cyanea. F Habit. G Leaf. H Flower. I Poricidal stamen, ventral
view with tepa!. J Stamen, dorsal view. K Pistil. (Takhtajan
1982)
reticulate; those of Kingia "extended" -sulcate and
psilate, i.e. with the sulcus extending to the proxi-
mal 1/2 of the grain; and those of Baxteria
"unipantocolpate" and microreticulate, i.e. the
exine is divided by the single, repeatedly branched
colpus into geometrically defined areas, a nearly
unique type among angiosperms (Chanda and
Ghosh 1976).
KARYOLOGY. Base chromosome numbers are
x = 7 in Kingia and Dasypogon, 14 in Baxteria and
9 in Calectasia (data from Anway 1969; Keighery
1984). In Calectasia, there is a cytological aberra-
tion of the 9th chromosome, which results in most
of the genetic material passing as a supergene,
causing permanent, unchangeable heterozygosity
(Anway 1969).
PoLLINATION. Kingia and Dasypogon produce
copious nectar and are visited by honey-eater
birds (Meliphagidae) and numerous flies, bees
(both native species and the introduced European
honeybee Apis mellifera) and ants, especially large
Iridomyrmex species, although the ants are prob-
ably nectar thieves (J.G. Conran, unpubl. observ.).
In Calectasia, stylar elongation is synchronised so
that the stigma in level with the anther pores at
anthesis, and the flowers are believed to be self-
pollinated (Anway 1969). Nevertheless, the an-
thers of two of the species change colour from
yellow to orange-red or brownish once the flower
ceases to be receptive, suggesting a visual cue for
biotic pollinators. The reddish brown flowers of
Baxteria smell of rotting meat and are presumably
pollinated by carrion flies.
FRUIT, SEED AND DISPERSAL. The fruits of the
Dasypogonaceae are indehiscent and enclosed
within the persistent and often undurated peri-
anth remnants, except of Baxteria where the fruit
is an explosively dehiscent capsule. The capsule in
Baxteria is septifragal, the valves each splitting in
half, the inner wall forming a tensioned plate
which flings the seed outwards at dehiscence
(Keighery 1983). Most fruits mature only a single
seed, producing a nut, although several seeds may
develop per ovary. The seeds in Dasypogon are
starchless, with a mealy, very thin-walled paren-
chyma. The inner layers of the seed coat collapse
during development. The embryo is lens-shaped
and broad, similar to the members of the
Commeliniflorae (J.G. Conran, unpubl. observ.).
All genera are fire-resistant or resprouters, and
flowering is enhanced dramatically in the season
following burning. Seed dispersal is passive in
most taxa, but ballistochrous in Baxteria, where
dehiscence of capsule is explosive (Keightery
1983).
PHYTOCHEMISTRY. Cell wall-bound ferulic acid
is present in all four genera (Rudall and Cad-
dick 1994). The record of chelidonic acid for
Dasypogon bromeliifolius (Ramstad 1953) seems
to be the only occurrence of this compound
 Dasypogonaceae
outside the Lilianae and merits further inves-
tigation.
DISTRIBUTION AND HABITATS. The Dasypo-
gonaceae are restricted to Australia. Baxteria,
Dasypogon and Kingia are endemic to the coastal
region of SW Western Australia. Calectasia ex-
tends from W Australia to Victoria. The members
of the family grow in a diversity of habitats includ-
ing Eucalyptus and Yarra woodland, heathlands
and exposed rocks.
SYSTEMATICS AND RELATIONSHIPS. The evi-
dence presented above suggests that all four
genera seem to be somewhat isolated morphologi-
cally. Although the family is normally placed in
the Asparagales (Dahlgren et al. 1985), various
characters, including the lack of petaloid perianth,
ovule structure, anatomical traits, cell wall-bound
ferulate and especially the molecular data are not
compatible with this placement, and they certainly
do not belong within the Xanthorrhoeaceae, in
which they were included by Cronquist (1981).
The rcbL tree of Chase et al. (1995) placed the
three dasypogonoid genera that were studied on a
well-supported branch within the commelinoids.
Although the number of commelinoid genera in-
cluded in the analysis is still relatively small, the
proximity of the Dasypogonaceae to the palms
may not be without significance, if one thinks of
the palmlike mode of stem-thickening in Kingia,
in which the sunken apical meristem leads to an
enormously wide thickening of the apical region
of the stem. Although Takhtajan (1994) has re-
cently erected a separate family (Baxteriaceae) for
Baxteria, current evidence supports the mono-
phyly of the Daspogonaceae as an early evolu-
tionary offshoot from the line leading to the
commelinoids and grasses, possibly near the
Rapateaceae, with which they share ovule, silica
body and some morphological features (Rudall
and Chase 1996).
KEY TO THE GENERA
1. Flowers solitary; perianth segments indurated 2 Arborescent woody perennial with leaves crowded
- Flowers in globular heads; perianth segments not indurated
3
2. Flowers borne at ground level; tepals free, 70-80mm long;
fruit and explosive capsule 4. Baxteria
- Flowers borne on shoot apices; tepals fused, up to 30 mm
long; fruit indehiscent, enclosed within the persistent
perianth 2. Calectasia
3. Inflorescence terminal, single per shoot; broacts on
peduncle widely spaced; outer tepals fused 1. Dasypogon
- Inflorescence axillary, numerous per shoot; bracts on
peduncle numerous, imbricate; outer tepals free 3. Kingia
193
1. Dasypogon R. Br. Fig. 46A-E
Dasypogon R. Br., Prod.: 263 (1810); George, Fl. Australia 46:
142-146 (1986).
Rosulate or palmlike perennials; stems short to
c. 3m tall, surrounded by persistent leaf sheaths;
leaves dispersed along the stems or apically
crowded, linear, expanded basally into a broad
sheath, margin toothed; inflorescence a globose
head on terminal peduncle bearing scattered
bracts; floral bracts navicular; flowers sessile;
outer tepals persistent, united in a tube, with stiff
dorsal hairs; inner tepals almost free, narrower
than outer; stamens separate from or adhering to
tepal bases; anthers dorsifixed, versatile; ovary
sessile, 3-angled, 3-locular, locules 1-ovulate;
style subulate, stigma small; fruit indehiscent
within persistent perianth, 1-2-seeded. Three sp.,
endemic to SW Western Australia.
2. Calectasia R. Br. Fig. 46F-K
Calectasia R. Br. Prod.: 263 (1810); George, Fl. Australia 46:
170-171 (1986).
Rhizomatic perennial with erect stem and wiry
roots. Leaves sessile, linear, the base indurated,
persistent and sheathing; flowers sessile, sur-
rounded by scarious bracts; tepals subequal,
united in tube in lower half, the lobes spreading
horizontally and blue or purple; stamens inserted
at base of perianth tube, fused basally to the tepals;
anthers basifixed, bright yellow or orange, dehisc-
ing by longitudinal slits or two terminal pores (C.
cyanea); ovary shortly stipitate, 1-locular, with 3
basal ovules; style filiform; fruit dry, indehiscent,
enclosed in the slightly hardened tube of the peri-
anth; seed oblong. Three spp., two in SW Western
Australia and one endemic to W Victoria.
3. Kingia R. Br. Fig. 45
Kingia R. Br., Ann. Sci. Nat. 8: 211 (1826);' George, Fl. Austra-
lia 45: 146-148 (1986).
apically; caudex covered with aerial roots pen-
etrating the persistent leaf bases; leaves linear, up
to 1m long, expanded at the base into a persistent
loose sheath, margins usually serrulate; inflores-
cences many, axillary just below stem apex,
headlike, their peduncles covered by imbricate
sheathing bracts; flowers sessile, tepals all similar,
persistent, free; stamens nearly free, 3 adhering
basally to inner tepals; anthers basifixed, introrse;
194 Dasypogonaceae

ovary sessile, 3-locular, locules with 1 erect ovule;
style subulate, stigma minutely 3-toothed; fruit
1-seeded within persistent perianth. Only one sp.,
K. australis R. Br., a conspicuous plant in SW
Western Australia.
4. Baxteria R. Br.
Baxteria R. Br. ex Hook., Hooker's J. Bot. Kew Gard. Misc. 2:
492, 494 (1843), nom. cons.; George, Fl. Australia 45: 141-
142 (1986).
Rudall, P., Chase, M. 1996. Systematics of the Xanthor-
rhoeaceae sensu lato: evidence for polyphyly. Telopea 6:
185-203.
Staff, LA., Waterhouse, J.T. 1981. The biology of arborescent
monocotyledons with special reference to Australian
species. In: Pate, J.G., McComb, A.J. (eds.) The biology of
Australian plants. Nedlands: University of W.A. Press,
pp. 216-257.
Takhtajan, A.L. 1982. See general references.
Takhtajan, A.L. 1994. Novae semejstva odnodolan'ch. Bot. Zh.
79: 65-66.
Wagner, W.M. 1977. Vessel types of monocotyledons: a sur-
vey. Bot. Not. 130: 383-402.

Perennial with short rhizome and thick roots;

stem condensed, bearing tufts oflinear leaves with
sheathing bases; flowers large, solitary, terminal,
sessile, surrounded by several bracts; tepals
subequal, free, indurated, not opening widely; sta-
mens 6, attached to the bases of tepals; anthers
basifixed, introrse; ovary 3-locular, each locule
with one erect ovule; style subulate, with small
stigma; capsule ovoid; dehiscence septifragal, the
valves also splitting into 2 halves, each half with
the inner wall forming a plate held under tension
and springing upwards to eject the seed; seeds glo-
bose. Only one sp., B. australis R. Br., restricted to
SW Western Australia.

Selected Bibliography

Anway, J.C. 1969. The evolution and taxonomy of Calectasia

cyanea R. Br. (Xanthorrhoeaceae) in terms of its present-
day variation and cytogenetics. Aust. J. Bot. 17: 147-159.
Chanda, S., Ghosh, K. 1976. Pollen morphology and its evolu-
tionary significance in Xanthorrhoeaceae. In: Ferguson,
I.K., Muller, J. (eds.) The evolutionary significance of the
exine. London: Academic Press, pp. 527-559.
Chase, M.W. eta!. 1995. See general references.
Cronquist, A. 1981. See general references.
Dahlgren, R.M.T. eta!. 1985. See general references.
Fahn, A. 1954a. The anatomical structure of the Xanthor-
rhoeaceae Dumort. Bot. J. Linn. Soc. 55: 158-184.
Fahn, A. 1954b. Metaxylem elements in some families of the
Monocotyledonae. New Phytol. 53: 530-540.
Fahn, A. 1961. the anatomical structure of the Xanthor-
rhoeaceae Dumort. and its taxonomic position. Recent Adv.
Bot. 1: 155-160.
Keighery, G.J. 1983. Ballistochory (explosive seed dispersal) in
Baxteria R. Br. (Xanthorrhoeaceae). W. Aust. Nat. 15: 163-
166.
Keighery, G.J. 1984. Chromosome numbers of Australian
Liliaceae. Feddes Repert. 95: 523-532.
Ramstad, E. 1953. Ober das Vorkommen und die Verbreitung
von Chelidonsaure in einigen Pfianzenfamilien. Pharm.
Acta Helv. 28: 45-55.
Rudall, P. 1994. The ovule and embryo sac in Xanthor-
rhoeaceae sensu lato. Flora 189: 335-351.
Rudall, P., Caddick, L.R. 1994. Investigation of the presence of
phenolic compounds in monocotyledonous cell walls using
UV fluorescence microscopy. Ann. Bot. 74: 483-491.
 Ecdeiocoleaceae 195

Ecdeiocoleaceae ridge. Below the sclerenchyma cap are the

palisadelike, radially elongated chlorenchymatous
H.P. LINDER, B.G. BRIGGS, peg cells in 2-3 layers. Each bicollateral vascular
and L.A.S. JoHNSON 1 bundle is surrounded by a sheath of scleren-
chyma. The peripheral vascular bundles are
smaller than the medullary vascular bundles
(Cutler 1969, Briggs and Johnson 1998).

INFLORESCENCE AND FLOWER STRUCTURE. The

Ecdeiocoleaceae D.F. Cutler & Airy Shaw, Kew Bull. 19: 495 flowers are borne in the axils of bracts, and are
(1965). aggregated into dense spikelike heads, with the
male and female flowers occupying alternating
Evergreen, rushlike, caespitose, monoecious sections of the head (Fig. 47). The flowers are
plants, with a villous or pubescent creeping rhi- dorsiventrally flattened, with 6 tepals, 4 or 6
zome. Culms photosynthetic, simple or with 1-3 tetrasporangiate stamens and hi- or tri-locular
inflorescence-bearing branches, terete, surface ovaries. The styluli are covered on 1 side with long
striate. Leaves reduced to tightly convoluted
sheaths in the adult plants; sheath split to the base,
usually with a mucro which may be developed into
a small awn. Inflorescence of 1- few spikelets, ter-
minating culm or inflorescence branches, glumes
all or mostly fertile, either male or female at base
and flower sex alternating once or twice along
spikelet axis. Flowers subtended by a glossy, dark
brown or black, rigid, ovate bract longer than the
flower, unisexual. Perianth of 6 tepals in 2 whorls,
dorsiventrally compressed, the outer 2 lateral seg-
ments conduplicate and keeled, the inner 4 more
or less flat, glabrous apart from the pubescent
keel. Male flowers with a minute pistillode and 4
or 6 stamens; filaments free, slender; anthers
bilocular, dorsifixed, 2-lobed, dehiscing by lon-
gitudinal slits. Female flowers with staminodes;
ovary superior, free, 2-celled with 2 or 3 bushy
styluli; each locule with a solitary atropous pendu-
lous ovule. Fruit a 3-locular capsule or a nut.
Two monotypic genera from southwestern
Australia, restricted to nutrient-poor Kwongan
soils.

VEGETATIVE MORPHOLOGY AND ANATOMY. The

plants are caespitose, rushlike, about 1m tall, with
well-developed rhizomes (Fig. 47). The culms are
the photosynthetic organs, the leaves have been
reduced to 1-4 sheaths on each culm. The culms
are deeply grooved, with the stomata near the
bases of the grooves, which are further occluded
by interlocking papillae. The ridges between the
grooves are capped by thick sclerenchyma which
sometimes extends down the sides of the groove in
the upper culm and in Georgeantha is connected
to the parenchyma around the outer vascular
Fig. 47A-F. Ecdeiocoleaceae. Georgeantha hexandra. A
bundles by a sclerenchyma rib central in each Habit. B Spikelet. C Staminate flower. D Same, with some
tepals and stamens removed. E Pistillate flower. F Dehiscing
'Deceased capsule, freeing a seed. (Orig.)
196 Ecdeiocoleaceae
and branching stigmatic branches that are them-
selves papillose; the stigmas are Dry.
EMBRYOLOGY. The development of the male ga-
metophyte is not known. The bitegmic, atropous
ovules are tenuinucellate. The megagametophyte
is tetrasporic, most similar to the Drusa type,
without antipodal cells. The micropylar nucellar
cells are in 1 or 2 layers, and are isodiametric in
shape. The nucellus is almost circular. There is no
evidence of starch grains in the mature megaga-
metophyte (Rudall1990).
POLLEN MORPHOLOGY. Chanda (1966) and Ladd
(1977) reported the pollen to be graminoid, with
an annulate aperture with a plug (Fig. 48). The
annulus is formed by a thickening of the foot layer
(Linder and Ferguson 1985). The interapertural
wall is smooth, and lacks scrobiculi (Ladd 1977).
The grains are 37 X 41 mm (Chanda 1966).
KARYOLOGY. Only approximate counts are avail-
able (B. Briggs, unpubl.), of 2n = 64-66 for
Georgeantha and 2n = ca. 48 for Ecdeiocolea.
FRUIT AND SEED. The ovary is a dehiscent capsule
or a nut, thus the fruit or seed is the dispersal unit.
REPRODUCTIVE BIOLOGY. The family is wind-
pollinated. Plants survive fire by resprouting from
the rhizome.
Fig. 48. Ecdeiocoleaceae. Georgean tha hexandra, pollen
showing small aperture with annulus and operculum, SEM
X2200. (Photo C. Porter)
ECOLOGY AND DISTRIBUTION. This family is re-
stricted to the very nutrient-poor sandy Kwongan
soils of western Australia.
AFFINITIES. Until1965 the family was included in
Restionaceae. Cutler and Shaw (1965) separated
the family from Restionaceae on the basis of the
culm anatomy: this has been further corroborated
by the very different embryology and the lack of
scrobiculi in the pollen. However, the family re-
sembles Restionaceae and Poaceae in the atro-
pous, pendulous, tenuinucellate ovule, single in
the loculus. Further, the family, like Restionaceae,
Joinvilleaceae and Poaceae has the 28-kb inver-
sion (Doyle et al. 1992). Although a close relation-
ship to Restionaceae is evident, it is not clear what
the exact relationship is between Restionaceae,
Ecdeiocoleaceae and Anarthriaceae (Linder and
Rudall1993; Kellogg and Linder 1995).
KEY TO THE GENERA
1. Culms with 2 or 3 branches, sheaths caducous; stamens 6,
fruit a capsule 1. Georgeantha
- Culms unbranched, sheaths persistent, stamens 4, fruit a
nut 2. Ecdeiocolea
1. Georgeantha B. G. Briggs &
L.A. S. Johnson Fig. 47
Georgeantha B. G. Briggs & L. A. S. Johnson, Telopea 7: 307
(1998).
Culms with 2 or 3 inflorescence branches, the low-
est internode and base of the second internode
densely woolly with long white hairs, with 3 or 4
persistent sheaths, the lower sheaths caducous.
Spikelets with 8-12 flowers. Male flowers with soft
tepals and 6 stamens. Female flowers with 3-locu-
lar ovary (although often 1 or 2loculi aborted) and
3 styluli. Seeds patterned with convex, strongly
lobed cells. Culms with grooves extending halfway
to base of chlorenchyma, stomates and accompa-
nying sclereids at base of grooves, ridges with a
central sclerenchyma rib 1-3 cells wide extending
from the parenchyma around the outer bundles to
the sclerenchyma cap. One sp., G. hexandra B. G.
Briggs & L.A. S. Johnson, Jurien-Eneabba region
north of Perth in the west of Western Australia, in
seasonally moist Kwongan sites in a region oflow
rainfall.
2. Ecdeiocolea F. Muell.
Ecdeiocolea F. Muell., Fragm. Phytogr. Austral. 8: 236 (1874)
 Ecdeiocoleaceae/Eriocaulaceae 197

Caespitose; rhizome hairs inconspicuous, covered Eriocaulaceae

by glossy cataphylls. Culms unbranched, glabrous,
the lowest 1(2) internodes pruinose, with 1(2) per- T.STUTZEL
sistent, appressed sheaths high on the culm.
Spikelets solitary with 20-50 flowers (the lowest
few not fully developing). Male flowers with rigid
tepals and 4 stamens. Female flowers with 2-
locular ovary and 2 styluli. Fruit a nut. Seeds with
an irregular pattern oflongitudinallines of convex
cells. Culm with grooves extending to base of chlo- Eriocaulaceae P. Beauv. ex Desv. in Ann. Sci. Nat. (Paris) 13: 47
renchyma, stomates and accompanying sclereids (1828), nom. cons.
on sides of grooves, ridges without a central
sclerenchyma rib. Only one sp., E. monostachya F. Medium-sized perennials, hapaxanthic perennials
Muell., widespread in Kwongan habitats in drier or annual herbs, usually rosulate, often tufted,
parts of the west of extratropical Western Austra- rarely caulescent, with horizontal or erect rhi-
lia, in well-drained sites in a region oflow rainfall. zomes. Indumentum of multicellular uniseriate
and sometimes glandular hairs, terminal cell
simple or bibranched (malpighiaceaous hairs).
Selected Bibliography Leaves usually spirally arranged, rarely distichous,
in some species distichous in the vegetative parts
Briggs, B.G., Johnson, L.A.S. 1998. Georgeantha hexandra, a and spirally in the inflorescence or vice versa.
new genus and species of Ecdeiocoleaceae (Poales) from
Western Australia. Telopea 7: 307-312. Inflorescences forming single headlike indetermi-
Chanda, S. 1966. On the pollen morphology of the Cen- nate spikes (here called heads) on thin peduncles,
trolepidaceae, Restionaceae and Flagellariaceae, with or composed of many (up to 1000) such heads.
special reference to taxonomy. Grana 6: 355-415. Peduncles with a closed sheath at the base. Heads
Cutler, D.P. 1969. Juncales. In: Metcalfe C.R. (ed.) Anatomy of composed of 10 to more than 1000 unisexual
the monocotyledons, Vol. 4. Oxford: Clarendon Press.
Cutler, D.P., Shaw, H.K.A. 1965. Anarthriaceae and Ecdeio- flowers. Heads subtended by an involucrum
coleaceae: two new monocotyledonous families, separated (involucra! bracts) different in size and shape
from the Restionaceae. Kew Bull. 19: 160-168. from bracts subtending flowers; the two types con-
Doyle, J.J., Davis, J.I., Soreng, R.J., Garvin, D., Anderson, M.J. nected by intermediates. Inner involucra! bracts
1992. Chloroplast DNA inversions and the origin of the
grass family (Gramineae). Proc. Nat!. Acad. Sci. U.S.A. 89:
petaloid and capable of hygroscopic movements
7722-7726. in Paepalanthus subgen. Xeractis and Syngon-
Kellogg, E.A., Linder, H.P. 1995. Phylogeny of Poales. In: anthus sect. Eulepis. Flowers generally unisexual
Rudall, P.J., Cribb, P.J., Cutler D.P., Humphries, C.J. (eds.) (hermaphrodite only in Rondonantus flabelli-
Monocotyledons: systematics and evolution. pp. London: formis, Syngonanthus amazonicus and few related
Royal Botanic Gardens, Kew, pp. 511-542.
Ladd, P.G. 1977. Pollen morphology of some members of the
species), mostly monoecious, rarely and then not
Restionaceae and related families, with notes on the fossil constantly dioecious, heterochlamydeous, actino-
record. Grana 16: 1-14. morphic or slightly zygomorphic, trimerous or
Linder, H.P., Rudall, P.J. 1993. The megagametophyte in dimerous, nearly always isomerous, rarely
Anarthria (Anarthriaceae, Poales) and its implications for heteromerous (2- and 3-merous). Sepal whorl and
the phylogeny of the Poales. Am. J. Bot. 80: 1455-1464.
Linder, H.P., Ferguson, I.K. 1985. On the pollen morphology bracts white, brownish, yellow, or dark green to
and phylogeny of the Restionales and Poales. Grana 24: black. Petal whorl generally colorless or white,
65-76. often hyaline, sometimes yellow or brown when
Meney, K.A., Pate, J.S. (eds) 1998. Australian rushes-biology, dried, rarely (Syngonanthus anthemidiflorus)
identification and conservation of Restionaceae and allied petals and gynoecium yellow. Stamens usually 3 or
families. Perth: University of Western Australia Press.
Rudall, P.J. 1990. Development of the ovule and megagame- 2, oppositipetalous, or 6 or 4 in 2 whorls (only
tophyte in Ecdeiocolea monostachya .. Austr. Syst. Bot. 3: Eriocaulon, Mesanthemum), Syngonanthus ama-
365-274. zonicus with a single stamen. Anthers introrse,
basifixed or dorsifixed, 2-thecate 4-sporangiate, in
some species 2-thecate 2-sporangiate by facial
reduction (Lachnocaulon, Blastocaulon, some
spp. of Paepalanthus). Ovary 3- or 2-locular, each
locule with a single atropous pendulous ovule.
Fruit a dorsicidal loculicidal capsule. Seed coat
longitudinally striated or "hairy" by residuals of
198 Eriocaulaceae

anticlinal walls of the inner layer of the outer

integument. Embryo small, bell-shaped, poorly
differentiated, at the micropylar end laterally
attached to the starchy endosperm.
A family of 10 genera and about 700-1400
species, from humid lowland areas to the alpine
region of the tropics and subtropics, best devel-
oped in S America; two spp. of Eriocaulon, tem-
perate regions of the Northern Hemisphere.

VEGETATIVE MORPHOLOGY. Perennial species

often have thick rhizomes; some species form a
stout trunk up to 80 em tall covered by adventi-
tious roots. In most species, the leaves are rosulate
(Figs. 49, 50). Nonrosulate, leafy stems occur
in some aquatic species with long floating axes, in
a few species with reptant stems, and in species
with erect, up to 3m high stems (Paepalanthus
sect. Actinocephalus). Erect elongated leafy stems
can also be formed by intense intercalary growth
during and after the differentiation of the inflores-
cence (P. polyanthus). In some species, the leaf
bases form water-storing cisterns similar to in
many Bromeliaceae, which seem to serve as fire
protection. Normally, the leaves are narrow
and grasslike, but in larger species and species
with water cisterns they can be up to 15 em
wide (E. macrobolax, P. bromelioides). The mor-
phology of the leaf apex is often of diagnostic
value.

VEGETATIVE ANATOMY (Poulsen 1888; Mal-

manche 1919). The roots have a polyarchic
vascular bundle with parenchymatic elements in
the center. Root hairs rise from special short
trichoblasts or are lacking, as in many species of
muddy habitats. Vessels with simple or scalari-
form perforations may occur, tracheids are always
present. The endodermis often has U- or 0-
shaped, brown, thickened walls. In Eriocaulon and
Mesanthemum the innermost cell layers of the Fig. 49A-R. Eriocaulaceae. A-L Eriocaulon woodii. A
cortex also have thickened walls and thus the Stoloniferous plant. B Involucra! bract. C Sepal. D Staminate
endodermis appears multilayered. In Eriocaulon, flower. E Same, calyx removed. F Expanded calyx of staminate
the outer part of the root cortex forms a highly flower. G Pistillate flower. H Sepal. I Petal with "epipetalous"
specialized aerenchyma formed of alternating gland. J Pistil. K Seed. L E. ussuriense, habit. M, N E. setaceum.
M Habit. N Leaf. 0, P E. longipetalum. Q Habit. R Polymerous
transversal layers of stellate parenchyma cells and flower. Q, R E. sieboldianum. Q Staminate flower, the sepals
radial rows of longitudinally extended and fused into an adaxial spathe, the petals lifted on an
flattened cells without any contact to the laterally anthophore. R Pistillate flower. (Takhtajan 1982)
neighboring cells. Thus the root appears seg-
mented vermiform. In Mesanthemum, Syngon-
anthus, Rondonanthus, and some species of In the leafy stem there is little or no pith. The
Leiothrix, the aerenchyma is less specialized; biconcentric, leptocentric, or collateral bundles
Paepalanthus, Tonina, Lachnocaulon, and most may have tracheids and vessels of the spiral type
species of Leiothrix have roots without aeren- and net type. Both scalariform and simple per-
chyma. forations occur. In aquatic species, the xylem is
 Eriocaulaceae
Fig. SOA-R. Eriocaulaceae. A-D Leiothrix nubigena. A Habit.
B Staminate flower. C Pistillate flower. D Pistil with carina!
appendices and commissural stylodia. E-1 Ton ina fluviatilis.
E Habit. F Staminate flower. G Staminate flower, sepals
removed, petal tube distally expanded. H Pistillate flower, the
petals reduced to small hairy lobes; note bibranched stylodia.
I Seed. J, K Syngonanthus anthemidijlorus. J Habit. K Pistil
with appendices. L-Q Philodice hoffmannseggii. L Habit. M
Flower head. N Staminate flower with basally fused petals. 0
Pistillate flower, the petals fused in the middle and basally
free. P Expanded petal tube of pistillate flower. Q Pistil
with carinate appendices and commissural stylodia. R Mes-
anthemum rutenbergianum, petal tube, basally petals free.
(Takhtajan 1982)
199
reduced and vessels may be lacking (Tonina). A
cortex is generally well developed, especially in the
basal part of the stem, where it may form a thick
ventilation tissue. For one species of Syngon-
anthus abnormal secondary thickening growth
similar to that in Dracaena has been reported
(Castro 1986) but it is more likely that it is a kind
of late primary thickening growth.
The peduncles have a very characteristic
anatomy. They are obtuse or acutely angled and
generally more or less twisted. Aerenchyma
strands can be found beneath or between the
angles. The longitudinal aerenchymatous parts
are separated by a more or less sclerified mecha-
nical tissue ofT- or H-shaped strands. The central
cylinder consists of 2 circles of concentric or
biconcentric vascular bundles, the outer and the
inner circle separated by a sclerenchymatic
sheath. The outer bundles are opposite the aeren-
chyma, the inner opposite the mechanical ele-
ments. The bundles contain spiral tracheids with
scalariform perforations. In the peduncles gener-
ally a parenchymatous pith is developed.
The leaves are bifacial, mostly hypostomatic,
rarely amphistomatic, and often hairy. The sto-
mata are paracytic with 2 lateral subsidiary cells.
In some aquatic species stomata are lacking. The
leaves usually have a single-layered cutinized epi-
dermis; in some species an adaxial, more rarely
also abaxial1-3-layered hypodermis occours. The
epiderms on the abaxial and adaxial sides are
linked by several buttresses of mechanical tissue,
each containing 1 (rarely more than 1) vascular
bundle. The buttresses run from the leaf base to or
close to the apex and separate the leaf into several
parallel air channels with sparse or no palisade
parenchyma. In some species, bundles without
buttresses alternate regularly with buttressed
bundles. The chlorenchyma consists of either a
diffuse aerenchyma or of transversal diaphragmas
of star parenchyma. The latter is common in
Eriocaulon and causes the "fenestration" of the
leaves. The leaf bundles may be anastomosing, but
anastomosing bundles are not buttressed and
therefore hardly to be seen. The bundles are
always surrounded by a bundle sheath of large
cells without chloroplasts. The anatomy of
the closed peduncle sheath resembles that of the
leaves, although anastomosing bundles are more
common.
Hairs are often present on nearly all parts of the
plant and are of taxonomic importance. Multi-
cellular uniseriate acute hairs with smooth surface
represent the only type found on stems bearing
rosulate leaves, whereas elongated parts of stems,
200 Eriocaulaceae

peduncle, sheath, and the leaves may have also calyx, the sepals are in lateral position. There are
glandular hairs and bibranched so-called mal- 2 whorls of stamens (Eriocaulon and Mes-
pighiaceous hairs of different size and sculpture. anthemum) or only 1 whorl of oppositipetalous
On vegetative parts, all kind of hairs have an iso- stamens (all other genera), and a trilocular or
diametric foot cell and a more or less globose bilocular gynoecium. The flowers are unisexual by
basal cell, the latter sometimes black, dark brown, late abortion of either stamens or carpels. The
or red and often persisting on older parts of the estivation is a pert. The sepals are generally free or
plant when the rest of the indumentum has been fused only at the base; rarely, a tube is formed. In
lost. Bracts, sepals, and petals never bear mal- some species of Eriocaulon (mostly in male, rarely
pighiaceous hairs but often capitate hairs with in female flowers) the sepals are fused into an
complicated outer and inner sculpture; they are adaxial spathe. The sepals seem to lack veins, but
common especially on the apical part of the sepals often have 1 median vascular bundle which does
in Paepalanthus. The foot cells of perianth hairs not reach the apex. The sepals are usually hairy on
are normal epidermal cells. their margins and on the abaxial surface. The
adaxial surface is usually glabrous. Sometimes the
INFLORESCENCE STRUCTURE (Stiitzel 1984). The hairs are crowded at the apex (many species of
basic unit of inflorescences in all species of the Paepalanthus); in some species the sepals are gla-
family is a head-shaped indeterminate spike (Figs. brous. The sepals are usually brownish, green or
49, 50). The heads are borne on peduncles and are nearly black, rarely hyaline to white. The hairs are
generally supplied with a closed sheath at their always white.
base formed by a single leaf. The sheath may be In Eriocaulon, the petals are free in both female
shorter or longer than the other leaves and either and male flowers and are abaxially glabrous
ends in a small reduced leaf blade or is cut and ventrally and on the margins hairy. In male
transversally. A closed sheath is lacking in flowers an anthophore can be found between the
Philodice and in a few species of Paepalanthus calyx and the petal whorl. In Eriocaulon and
subgen. Platycaulon. The heads have an involu- Mesanthemum, the filaments of the inner stamens
crum of few-numerous "outer bracts" or involu- are fused with the petals from their base up to or
cra! bracts. The flowers are generally borne in axils near to the gland. A similar but shorter fusion is
of "inner bracts", but in Syngonanthus the inner present in Rondonanthus flabelliformis. In male
flowers have no subtending leaves. flowers of the other genera, the petals are generally
An inflorescence system may consist of 1 or fused to a tube, which is usually glabrous, but
few-many heads (up to more than 1000). A typical hairy on the margins in Mesanthemum; in
unit of such inflorescences is a dibotryum with a Paepalanthus subgen. Xeractis the tube is hairy on
terminal raceme in which the lateral peduncles are margins and inside. In the female flowers of
not subtended by bracts (pherophylls) but have a Mesanthemum, the petals are congenitally fused
closed sheath. The lowermost heads in such a in the middle part, but free apically (primarily) as
dibotryum may be replaced by a complete dibo- well as basally (secondarily, schizogenous). They
tryum. The condensation of 1-several dibotryas are hairy on the margins and ventrally. The petals
results in an umbel-like inflorescence as in of the female flowers in Syngonanthus, and
Syngonanthus caulescens or Paepalanthus subgen. Philodice and the hermaphrodite flowers in
Actinocephalus. In Paepalanthus subgen. Actino- Rondonanthus flabelliformis show the same
cephalus the inflorescence is composed of several peculiar kind of fusion; it does not occur in male
to numerous umbels like this and thus may con- flowers. In Tonina, Blastocaulon, Paepalanthus,
sist of more than 1000 heads. Leiothrix, Syngonanthus, and Philodice the petals
In Paepalanthus subgen. Platycaulon the of the male flowers are fused to a glabrous, gener-
peduncles are flattened and bear several heads. ally 3-lobed tube. The female flowers have free
This inflorescence is an axillarydibotryum without petals in Blastocaulon, Tonina (reduced to small
elongated internodes within the inflorescence and hairy lobes), Paepalanthus and Leiothrix. In
the scape (peduncle) is beneath the dibotryum. Rondonanthus the petals are either free in flowers
of both sexes or in female flowers show the same
FLOWER STRUCTURE. The hypogynous flowers fusion of the middle part of the petals as in
are very small (0.25-6mm), actinomorphic or Mesanthemum (often described as syngonanthoid
zygomorphic, shortly pedicellate or sessile. In fusion). In Lachnocaulon, the petals are com-
flowers with trimerous calyx, the median sepal pletely lacking or reduced to hairs (no ontogenetic
is in adaxial position, in flowers with dimerous studies available). The petals are hyaline or
 Eriocaulaceae
whitish or, when dried, pale brownish. Only a few
species of Syngonanthus have yellow petals during
anthesis. Before anthesis they are pale yellow to
whitish and after an thesis they become pale again.
In Eriocaulon and Mesanthemum, the petals
bear a black gland (Fig. 49E,I), which is usually
described as epipetalous, but is formed by the
apex of the petal primordium; the part distal to the
gland in mature petals represents a dorsal
effiguration of the petal (Stiitzel 1985).
In Eriocaulon, Paepalanthus and Lachnocaulon,
and Syngonanthus the petal whorl may be
dimerous or trimerous; in all other genera it is
trimerous. In Eriocaulon and Paepalanthus the
sepal and petal whorls may be isomerous or
heteromerous, in Lachnocaulon only L. digynum
is heteromerous. The stamen whorls and the petal
whorls are isomerous. The anthers are gen-
erally bithecous and 4-sporangiate, basifixed, or
dorsifixed and introrse. For Tonina, Philodice,
Lachnocaulon, and Blastocaulon monothecate 2-
sporangiate anthers are reported. In Tonina and
Philodice this is due to occasional and lateral re-
duction, in Lachnocaulon, Blastocaulon, and some
spp. of Paepalanthus, this is due to facial reduc-
tion (2-thecate 2-sporangiate). The anthers are
dark green or black in most species of Eriocaulon
and in Mesanthemum, yellow in the few species
of Syngonanthus with yellow petals (e.g., S.
anthemidiflorus), and white in all others. The
pollen is white in all genera except in species
with yellow anthers, where it may also be yellow.
The gynoecium is di- or trimerous. The style is
shorter or as long as the stylodia in all genera
except Leiothrix, where it is more than 2 times
longer. In Eriocaulon and Mesanthemum, the stig-
mas are in carinal (dorsal) position to the carpels,
in all other genera they are inserted commis-
surally and may be simple or 2-branched. If the
stigmas are in commissural position, there are
appendices in carinal position. The appendices
are homologous to the stigmas in Eriocaulon and
Mesanthemum and produce a lot of sucrose nectar
during anthesis. In some species they produce
odorous substances (Paepalanthus subgen.
Platycaulon and sect. Actinocephalus). In a few
species of Paepalanthus and Syngonanthus, the
appendices are reduced to nearly invisible rudi-
ments and produce no nectar.
EMBRYOLOGY. Complete data on embryology are
available only for Eriocaulon and Leiothrix
(Monteiro-Scanavacca and Mazzoni 1978; Arekal
and Ramaswamy 1980; Ramaswamy and Arekal
1982; Ramaswamy et al. 1983).
201
The anther wall consists of epidermis, endoth-
ecium, a middle layer, and a glandular tapetum.
One or 2 rows of archesporia! cells may occur.
The tapetum cells are 1-nucleate throughout their
existence or rarely may become 2-nucleate in
some cells, when meiosis starts. Microsporo-
genesis is successive. The pollen is 3-celled at
shedding. The ovules are atropous, pendulous,
bitegmic, and tenuinucellate. The micropyle is
formed by the inner integument only. Both
integuments are 2-layered. The outer layer of the
outer integument mostly degenerates while the
anticlinal walls of the inner layer of the outer
integument form the characteristic striate or
"hairy" -like surface. The funicle is very short. The
nucellus develops a hypodermal archesporia!
cell, which enlarges and functions directly as
megaspore mother cell. The megaspores form a
linear tetrad of which the proximal megaspore is
functional. The development of the female
gametophyte follows the Polygonum type. The
distal, single-layered part of the tenuinucellate
nucellus degenerates. The endosperm is at first
Nuclear, but after the first free divisions cell
wall formation starts at the chalazal end and
gradually extends to the micropylar end. Wall
formation may also start simultaneously from
the chalaza! and the micropylar end. The endo-
sperm consists of an outer layer of small cells
with few starch grains, a chalaza! region of small
cells with no or few starch grains, and a central
part of large cells filled with compound starch
grains.
The embryo is campanulate and attached to the
endosperm at the micropylar end. The embryo
shows no exomorphologic differentiation. Germi-
nation starts with the formation of at least 2leaves
before the first root is formed.
POLLEN MORPHOLOGY (Thanikaimoni 1965).
Pollen grains are dispersed singly. The pollen is
spheroid to ellipsoid, 25-35 [till in diameter,
spiraperturate (the aperture often runs around the
pollen grain as the suture does at a tennis ball).
The pollen surface is echinate to echinulate and
may have spinulae of different size (T. Stiitzel,
unpubl.).
KARYOLOGY. Chromosome numbers are known
from Eriocaulon and a few species of Pae-
palanthus. The base numbers 8, 9, 15, 20, and 25
have been reported, on which diplo-, tetra-, and
hexaploids are based. The chromosomes are very
small.
202 Eriocaulaceae

PoLLINATION. Eriocaulaceae are entomophilous. DISPERSAL. Within the family, different ways of
Flies (Diptera) and beetles have been observed as seed dispersal can be found. In Eriocaulon the
pollinators. Paepalanthus subgen. Platycaulon whole flowers of both sexes (including bracts)
and P. sect. Actinocephalus, at least in some loosen from the receptacle. They may be dispersed
habitats, seem to be pollinated mainly by Ceram- by wind and/or water. In Paepalanthus three dif-
bycidae. Most species enforce cross-pollination by ferent ways of dispersal are known. In one case,
separate flowering periods of male and female the diaspore, which consists of a whole female
flowers. In Paepalanthus, Syngonanthus, and flower except the sepal whorl, is catapulted up to
Leiothrix, there is a diurnal change between male 2m by the drying sepal whorl. The 3 seeds remain
and female phase. Flowering within 1 head may together or may be scattered singly during the
continue for nearly 1 year, but a single flower is flight. In other cases, the diaspore is elevated only
open for only a few hours in these genera. Male slightly above the surface of the head by the drying
and female phase are synchronous for all heads of sepal whorl and is dispersed by wind. Bracts as
one plant and for all plants of a population, prob- well as male flowers remain on the receptaculum
ably even for the species as whole (Stiitzel 1984). in both cases. Wind or catapult dispersal may be
In Eriocaulon, the anthesis of each flower may followed by water dispersal. In Paepalanthus sect.
last much longer (several days), male and female Actinocephalus the diaspore consists of the whole
stages are synchronized within the plant and head including peduncle and sheath. In this case,
generally within a population, but obviously not the plant produces several umbels of heads, each
within the species (Stiitzel 1984). During the short with 50 to 200 heads but only a few seeds per head.
period in which male and female phases are In Syngonanthus no elevating or catapulting effect
changing, neighboring flowers of the opposite sex can be found. The diaspores are dispersed by wind
may be open at the same time. In this case, self- or water. In Leiothrix flavescens, the seeds may be
pollination (geitonogamy) is possible and may scattered singly (Stiitzel1984).
even be an important factor in the reproductive
bioloy of Paepalanthus. PHYTOCHEMISTRY. The family is characterized
Little is known about the composition of the by oxalate crystals (druses and single crystals but
nectar and nothing about the various odorous no raphids). Silica bodies have been reported only
substances which are produced by the floral for Syngonanthus xeranthemoides, where they
glands in many species. occur together with sclereidal cells in the subepi-
dermal leaf tissue. Harris and Hartley (1980)
FRUIT AND SEED. In all genera the fruit is a locu- reported the presence ofUV-fluorescent constitu-
licidal capsule opening with a dorsal longitudinal ents in the cell walls of two species of the family,
slit. The single seed per locule is pendulous from indicating cell wall-bound ferulic, p-coumaric,
the placental top and fills the locule completely. and/or diferulic acids. The highly UV-absorbing
The opening of the locule is forced by the seed, flavonols quercetagetin and patuletin, but no
when the drying pericarp shrinks. other flavonolic compounds, were recorded by
The seed is elliptical or ovoid and has a charac- Bate-Smith (1968).
teristic sculpture. It may be reticulate or tubercu- Hoehne (1939) reported that heads and rhi-
late, often described as "hairy". This texture is zomes are used in folk medicine against several
formed by the anticlinal walls of the inner layer of kinds of diseases of the skin, but there is neither
the outer integument and may be more or less clinical proof nor any report on chemical data to
complicated. If only longitudinal walls remain support this.
prominent, the seed becomes striate. In ripe seeds,
the outer layer of the outer integument is often RELATIONSHIPS WITHIN THE FAMILY. Recent
completely obliterated. If the outer layer persists studies indicate that Rondonanthus is probably
in ripe seeds (some species of Eriocaulon and closest to the phylogenetic base of the family. An
Paepalanthus), the surface is very similar to that early bifurcation into 2 clades separates the genera
in Xyris (Xyridaceae), but in Xyridaceae the layer with syngonanthoid fusion of the petals including
forming the sculpture is the outer layer of the Eriocaulon from the rest of the family (Hensold
inner integument (Giulietti et al. 1984). For many and Giulietti 1991).
taxa, seed-coat characters seem to be more useful While most genera seem to be well defined,
for taxonomic purposes than floral characters Paepalanthus is a conglomerate of several, prob-
(Phillips 1996). ably not closely related groups of species of which
 Eriocaulaceae 203

some may even have their closest relatives in tropics; some species enter the humid temperate
the syngonanthoid clade. In its present circum- regions of N America and Europe, such as E.
scription Paepalanthus comprises primitive and septangulare (Ireland, Scotland, native) and
derived groups. The elevation of subgenera and E. cinereum (rice fields ofN Italy, anthropogenic).
sections of Ruhland's revision to distinct genera asIn E Asia Eriocaulon occurs in the Indo-Malayan
proposed sometimes is no solution, as "Eupae- region, in China, and also in Japan, and reaches
palanthus" sensu Ruhland in this case would form Sachalin. Syngonanthus and Paepalanthus are
a polyphyletic remain. mainly S American and with some species are
In older systems the family is generally divided represtented in Central America, Africa, and
into two subfamilies. In this case, the Eriocau- Madagascar. Syngonanthus has also a single
loideae (only Eriocaulon and Mesanthemum) species in the southern USA. Mesanthemum is
comprise the genera with 2 whorls of stamens and restricted to Africa and Madagascar, Leiothrix,
the Paepalanthoideae those with a single staminal Blastocaulon, Philodice to Central America.
whorl. This is rather artificial and separates Tonina occurs in northern S America, Central
Mesanthemum from the closely related Rondon- America, and Cuba. The species of Rondonanthus
anthus and Syngonanthus. Another classification are restricted to the Guayana Shield. Lach-
has been proposed by Stiitzel (1985), who distin- nocaulon is restricted to the east coast and the
guished three subfamilies, Eriocauloideae, Missisippi region of N America.
Syngonanthoideae, and Paepalanthoideae. This Eriocaulaceae occur mainly on poor sandy soils,
concept, which was based on the assumption that in marshy or boggy habitats, and in the moun-
Eriocaulon is the most primitive genus within the tains, even on rocks. Some species of Eriocaulon,
family, has now to be abandoned as well. Paepalanthus, and Leiothrix and Tonina grow
submersed in rivers, ponds, and in tempo-
AFFINITIES. Mainly on account of the unique rarily flooded places. Other species, mainly of
kind of inflorescences and the special kind of Paepalanthus, Leiothrix, and Syngonanthus, occur
spiraperturate pollen grains, the Eriocaulaceae in the mountains on rocks or on sandy soils in
have always been considered as a natural, mono- habitats which appear to be rather dry; they grow
phyletic group. Nevertheless, the Eriocaulaceae together with xerophytic plants such as Trimesia
are far from being isolated. Close and impor- and Melocactus. Several species are both annual or
tant relationships exist between Eriocaulaceae perennial according to climate and living condi-
and Xyridaceae, as evidenced by similarities in tions. The Eriocaulaceae have no spezialized
embryological features, echinate pollen, and the hibernating organs such as bulbs or underground
structure of seeds and inflorescences. Within the corms, they endure short dry periods with well-
Xyridaceae, subfam. Abolbodoideae shows more developed green leaves. Leiothrix niveus has hairy
and closer similarities with the Eriocaulaceae than green leaves in the summer but is completely
subfam. Xyridoideae. Subfam. Abolbodoideae covered with white hairs during the winter, so that
resembles especially Paepalanthus in pollen then only small wool cushions but no green parts
morphology and the peculiar appendices of the of the plant can be seen. Some species seem well
gynoecium, which are found in no other family. In adapted to periodic fires (such as the Velloziaceae
contrast, Xyris has atropous seeds similar to which cooccur with them in the same habitats)
Eriocaulaceae, while Abolboda has anatropous and the older parts of their elongated stem lack
seeds. leaves. The water cisterns may also be important
While in the past Eriocaulaceae have been in respect to fire protection.
placed together with Restionaceae close to Poales,
at present their relationship to Commelinales PARASITES. In the Eriocaulaceae the heads are
seems better substantiated and the family is often parasitized by larvae of Coleophoridae.
included in the Commelinales s.l or placed as Similar or the same species parasitize the heads of
a separate order Eriocaulales close to the Xyris and some Juncaceae in the same habitats.
Commelinales (Dahlgren and Clifford 1982; Caterpillars of different groups of Lepidoptera
Dahlgren et al. 1985). can be found on leaves of Eriocaulaceae. Larvae of
Diptera have been found mining in the rhizomes
DISTRIBUTION AND HABITATS. Eriocaulon has of Eriocaulon megapotamicum.
the widest distribution of all genera of the family Several species of smuts parasiting on Erio-
and is distributed throughout the tropics and sub- caulaceae have been described, e.g., Tolypospo-
204 Eriocaulaceae

rium hessii E. Muller on Eriocaulon lanatum, 9. Stamens 2-thecate, 4-sporangiate 9. Paepalanthus

Tolyposporium eriocauli Clint. on Eriocaulon - Stamens 2-thecate, 2-sporangiate 10. Blastocaulon
vittifolium, Sorosporidium mesanthemi E.
Muller on Mesanthemum radicans, and Ustilago 1. Rondonanthus Herzog
eriocauli Mass. Clint. on Syngonanthus
Rondonanthus Herzog, Feddes Rep. 29: 120 (1931); Hensold &
welwitschii. These smuts are all exclusive to Giuliettti, Ann. Mo. Bot. Gard. 78: 441-459 (1991), rev.
Eriocaulaceae, but this may be an effect of re- Wurdackia Moldenke (1957).
stricted collecting.
Perennial rosulate herbs; leaves alternate or disti-
PALEOBOTANY. Pollen has been recorded for E. chous. Flowers trimerous, unisexual, or hermaph-
septangulare from the Pleistocene of Ireland, and rodite and male (R. flabelliformis); female: sepals
for E. pellucid urn from the Pleistocene of Canada. free; petals fused only in the middle as in
Both species are sometimes regarded as synony- Syngonanthus or free, with 3 conspicuous, linear,
mous and cannot be distinguished on the basis of staminodes; gynoecium with appendices and bifid
pollen characters. stigmas, both inserted at the same level; male:
sepals free; petals fused into a tube; stamens 3,
EcoNOMIC IMPORTANCE. In Central Brasil bithecate, tetrasporangiate, basally fused with the
(Minas Gerais and neighboring areas) the heads petals or free; gynoecium rudimentary. Five spp.,
of several species (mainly Syngonanthus sect. restricted to the Guayana Shield in northern S
Eulepis) with their long peduncles are collected America.
and dried for room decoration. The value of The circumscription of Hensold and Giulietti
exported inflorescences, which are often colored (1991) comprises the monotypic genera Wur-
when marketed, has reached nearly 2 Mio. US $ dackia and Rondonanthus plus 2 spp. formerly
(A.M. Giulietti, pers. comm.) and several species belonging to Paepalanthus and 1 that had been
are endangered because of intensive collecting. placed in Syngonanthus. The genus is mainly
based on the presence of large staminodes in
KEY TO THE GENERA female flowers and is the only one which com-
1. Female flowers with conspicuous linear staminodes
prises species with and without the syngonanthoid
1. Rondonanthus fusion of the petals.
- Female flowers with inconspicuous tuberclelike staminodes
or without staminodes 2
2. Stamens in 2 whorls (i.e., 4 or 6); gynoecium without glands
2. Eriocaulon L. Fig. 49
(appendices); stigmas simple in carina! (dorsal) position of Eriocaulon L., Sp. Pl.: 87 (1753).
the carpels 3
- Stamens in only 1 whorl (i.e., 2 or 3 stamens),
oppositipetalous; gynoecium with carina! glands (appen- Mostly rosulate perennial or annual herbs, highly
dices); stigmas simple or 2-branched, commissural 4 variable in size (leaves 1-60 em long, 0.1-15 em
3. Petals free in male and female flowers 2. Eriocaulon wide, peduncles 1-120 em long) but very uniform
- Petals fused to a 3-lobed tube in male flowers; petals fused in habit. Submerged and floating species with
in the middle and free at base and apex in female flowers
3. Mesanthemum elongated leafy stems; terrestrial species with
4. Petals lacking in male flowers, reduced to hairs or lacking stout, usually horizontal rhizomes, sometimes
in female flowers 4. Lachnocaulon forming erect trunks up to 80 em tall. Inflores-
- Male and female flowers with petals 5 cence a single head or composed of 2-15 or more
5. Petals of female flowers free at base and apex but fused in
heads in the center of a rosette. Sometimes with
the middle 6
- Petals in female flowers free 7 proliferating heads and peduncles functioning
6. Peduncles with a closed sheath at the baseS. Syngonanthus like stolons. Flowers unisexual, mostly trimerous,
- Peduncles without closed sheath 6. Philodice more rarely dimerous or heteromerous, sub-
7. Style 3 times as long as the simple stigmas or longer; glands tended by bracts; male: sepals free or fused to an
(appendices) inserted on style distinctly below the stigmas
7. Leiothrix
adaxial spathelike organ; anthophore between
- Style as long as the stigmas or shorter; if present, glands sepals and petals sometimes present, as long as
(appendices) inserted at the same level as stigmas 8 sepals and longer than petals; petals free, with an
8. Persistent petals of the female flower reduced to small hairy "epipetalous" gland; 4 or 6 stamens in 2 whorls,
lobes as long as the fruit; peduncles subtended by an open bithecate, tetrasporangiate, the filaments in the
sheath, fused with the main axis 8. Tonina
- Persistent petals of female flowers longer than the fruit;
outer whorl free, in the inner whorl fused with
peduncles with closed sheath at the base or sheath lacking, petals up to the glands which, in color (brown,
but never open 9 dark green or black) and surface structure,
 Eriocaulaceae
resemble the glands formed by the rudimentary
carpels and mostly also the thecas of the stamens;
female: sepals free, sometimes fused to an adaxial
spathe; petals free, with "epipetalous" glands;
gynoecium with stigmas in carinal position; nectar
and scent production from epipetalous glands in
male and female flowers and from rudimentary
gynoecium in male flowers. About 400 spp., tro-
pical and subtropical regions of the world.
3. Mesanthemum Koern. Fig. 50R
Mesanthemum Koern., Linnaea 27: 527 (1856).
Rosulate plants of medium size (leaves 10-30cm,
peduncles 20-40 em long). Inflorescences single
heads or composed of few heads in the centre of a
rosette. Flowers unisexual, trimerous, bracteate;
male with basally fused sepals, with a short
anthophore between sepals and petals, petals
fused and with "epipetalous" glands which may
have some hairs; stamens 3 + 3, those of the outer
whorl free, filaments of the inner whorl fused to
the petals approximately half the distance to the
gland; female with free sepals and petals fused
only in the middle as in Syngonanthus; stigmas in
carinal position, appendices lacking. About 18
species in tropical or subtropical regions of Africa,
2 also on Madagascar. Records for S America
erroneous.
4. Lachnocaulon Kunth
Lachnocaulon Kunth, Enum. Pl. 3: 497 (1841).
Rosulate plants with 1-many heads. Flowers
trimerous except L. digynum with trimerous sepal
whorl and dimerous androecium and gynoecium;
in male flowers sepals free, petals completely lack-
ing; stamens inserted at an anthophore reaching
the length of the filaments; stamens 2-thecate 2-
sporangiate due to facial reduction (Stiitzel and
Gansser 1995); gynoecium reduced to glands; in
female flowers sepals free, petals lacking; gyno-
ecium with bifid stigmas and appendices (glands),
both inserted on same level. For L. digynum erro-
neously simple stigmas are reported. About eight
spp. in N America, Florida, and the Mississippi
region.
5. Syngonanthus Ruhl. Fig. 50J,K
Syngonanthus Ruhl. in Urban, Symb. Antill. 1: 487 (1900).
Comanthera L.B. Smith (1937).
Carptotepala Mold. (1951).
205
Rosulate or caulescent herbs, sometimes with !-
several elongated internodes between rosulate
parts. Scapelike stems in the vegetative part
without basal sheath, sometimes composed of
several internodes and then bearing a few reduced
leaves. Inflorescences in the center of the rosette,
single heads or composed of up to 150 heads, or
arranged in different ways to umbel-like inflores-
cences with more or less long, leafy stems; involu-
cra! bracts sometimes (sect. Eulepis) enlarged,
petaloid. Flowers unisexual, generally trimerous
and unisexual, in S. amazonicus and few related
spp. hermaphrodite and dimerous, usually sub-
tended by bracts, sometimes all but marginal
flowers without bracts; male flowers; sepals
free; petals fused to a 3-lobed, sometimes very
short tube; stamens bithecate, tetrasporangiate
with free filaments; gynoecium reduced to :±:
globose glands; female flowers: sepals free; petals
fused in the middle, apically and basally free.
Gynoecium with :±: globose appendices (glands),
and simple stigmas, both inserted at the same
level; appendices lacking in a few species. About
200 (80-250) spp., mainly S America but a few also
in Africa and Madagascar.
6. Philodice C. Mart. Fig. 50L-Q
Philodice C. Mart., Nova Acta Phys.-Med. Acad. Caes. Leop.-
Carol. Nat.-Cur. 17: 16 (1835).
Small rosulate or subcaulescent plants (stems 3-
17 em long); peduncles short (1-2cm), without
sheath; leaves filiform, 0.5 wide and 1.5 em long;
flowers as in Syngonanthus, but the petals of
female flowers not fading or shrinking after
anthesis, looking turgid even in dried speci-
mens; stamens reported as monothecate and
bisporangiate but often bithecate and tetra-
sporangiate. One (or 2?) spp., restricted to the
basins of the Amazonas and Orinoco rivers,
tropical S America.
7. Leiothrix Ruhl. Fig. 50A-D
Leiothrix Ruhl. in Engler, Pflanzenreich. V.30: 225 (1903);
Giulietti (1984) Estudos taxonomicos do g~nero Leiothrix.
Tese Livre-docencia, Univ. Sao Paulo (1984) (rev.)
Vegetative morphology and inflorescence struc-
ture similar to Eriocaulon. Peduncles 10-50cm
long, leaves 3-30 em long. Flowers trimerous;
male flowers: sepals free; petals fused at least
basally; stamens 3, bithecate and tetrasporangiate,
with free filaments; gynoecium reduced to glands;
206
female flowers: sepals free, petals free, gynoecium
with simple stigmas and appendices, the latter
inserted at the base or near the middle of the
style; style at least 3 times as long as the stigmas
(Fig. SOD). In subgen. Stephanophyllum heads al-
ways proliferating and peducles becoming thus
stolonlike, plants therefore may develop a tangle
of dry, dead peduncles covering rocks, in which
the rosettes (often without roots) are the only liv-
ing parts. About 37 spp., S America, mainly Brazil.
8. Tonina Aubl. Fig. 50E-I
Tonina Aub!., Hist. Pl. Gui.: 856 (1775).
Habit somewhat like Elodea canadensis, mostly
growing submersed. Leaves about 10 mm long and
2-4 mm wide, arranged spirally on the long erect
or floating stem. Inflorescences single, axillary
heads with short peduncles and open adaxial
sheaths; sheath and peduncle extraaxillary.
Flowers trimerous; male flowers: sepals at margin
fused half of their length; petals forming a tube;
anthophore between sepal and petal whorl, at
anthesis equalling or surpassing the petal tube;
stamens usually bithecate and tetrasporangiate,
sometimes and irregularly reduced to mono-
thecate and bisporangiate; filaments free. Only
one sp., T. fluviatilis Aubl., northern S America.
9. Paepalanthus Kunth
Paepalanthus Kunth, Enum. Pl. 3: 498 (1841).
Moldenkeanthus Morat (1976).
Rosulate, caulescent (erect or reptant) herbs or
rarely shrublike plants, from a few mm up to
about 4 m (P. hillarei) in size. Inflorescence a
single head or a group of heads terminating a
rosette, or single heads in the axil of the leaves of a
rosette, or inflorescences composed of up to more
than 1000 heads in different ways. Inflorescences
in sect. Actinocephalus formed by several umbels
of heads, each with about 50-200 heads; in subgen.
Platycaulon peduncles flattened and bearing a
group of 5 to 20 shortly stalked (sect. Divisi) or
sessile heads (sect. Conferti), peduncles bearing
such a group of heads with closed sheath at base,
the short peduncles of the single heads without
sheaths or bracts. Flowers trimerous, dimerous, or
heteromerous, male: sepals free or only basally
fused; petals fused; stamens bithecate and tetra-
sporangiate, rarely 2-sporangiate; filaments free;
gynoecium reduced to glands; female: sepals free
or only basally fused; petals free; gynoecium with
Eriocaulaceae
simple (subgen. Actinocephalus) or bifid (subgen.
Xeractis and Platycaulon) stigmas; subgen.
Xeractis with enlarged petaloid involucra! bracts
similar to Syngonanthus sect. Eulepis. About 400
(200-600) spp., most inS America, some in Cen-
tral America, Africa, and Madagascar. One record
from Japan due to confusion of labels.
Circumscription of Paepalanthus provisional;
some subgenera such as Platycaulon, Xeractis and
sect. Actionocephalus forming natural groups,
others are based on vegetative characters of little
significance.
10. Blastocaulon Ruhl.
Blastocaulon Ruh!. in Engler, Pflanzenreich IV.30: 223 (1903).
Small rosulate plants with very thin peduncles
and sometimes stolonlike branches. Flowers
trimerous, male flowers: sepals free; petals fused;
stamens 2-thecate, 2-sporangiate; filaments free;
gynoecium reduced to glands; female flowers:
sepals free; petals free; gynoecium with bifid
stigmas and appendices (glands). About 5 spp., all
in Minas Gerais, Brazil.
The genus is separated from Paepalanthus
exclusively because of the 2-thecate 2-sporangiate
anthers, but 2-sporangiate anthers occur also in
some species of Paepalanthus. Further informa-
tion is needed to decide whether Blastocaulon
should be recognized as distinct from Pae-
palanthus. In this respect it might be important to
realize that Blastocaulon (and some of the species
of Paepalanthus with 2-sporangiate anthers) grow
in extremely shady places together with filmy
ferns, while Eriocaulaceae normally prefer bright
places.
Selected Bibliography
Arekal, G.D., Ramaswamy, S.N. 1980. Embryology of
Eriocaulon hookerianum Stapf and the systematic position
of Eriocaulaceae. Bot. Not. 133: 295-309.
Bate-Smith, E. C. 1968. The phenolic constituents of plants and
their taxonomic significance. J. Linn. Soc. 60: 325-356.
Begum, M. 1968. Embryological studies in Eriocaulon quin-
quangulare L. Proc. Indian Acad. Sci. 67: 148-156.
Castro, N.M. 1986. Estudos morphologicos dos orgaos
vegetativos de especies de Paepalanthus KUNTH (Erio-
caulaceae). Thesis, Instituto de Biociencias, Univ. de Sao
Paulo, Sao Paulo.
Dahlgren, R.M.T., Clifford, H.T. 1982. See general references.
Dahlgren, R.M.T. eta!. 1985. See general references.
Engler, A. 1895. Eriocaulaceae. In: Engler, A. Die Pflanzenwelt
Ostafrikas und der Nachbargebiete, Theil C. Berlin: Dietrich
Reimer.
Erdtman, G. 1952. See general references.
 Eriocaulaceae 207

Giulietti, A.M. 1978. Modifica~oes taxonomicos no genero Stiitzel, Th. 1985. Die epipetalen Driisen der Gattung Erio-
Eriocaulon L. Bol. Bot. Univ. S. Paulo 6: 39-47. caulon (Eriocaulaceae). Beitr. Bioi. Pfianz. 60: 271-276.
Giulietti, A.M. 1984. Estudos taxonomicos no genero Leiothrix Stiitzel, Th. 1988. Untersuchungen zur Wurzelanatomie der
RUHL. These de Livre docencia, Univ. de Sao Paulo, Sao Eriocaulaceen. Flora 180: 223-239.
Paulo. Stiitzel, Th., Gansser, N. 1995. Floral morphology of North
Giulietti, A.M., Monteiro, W.R., Mayo, S.J., Stephens, J. 1984. A American Eriocaulaceae and its taxonomic implications.
preliminary survey of testa sculpture in Eriocaulaceae. Feddes Repert. 106: 495-502.
Beitr. Bioi. Pfianz. 62: 189-209. Takhtajan, A. 1980, 1982. See general references.
Harris, P.J., Hartley, R.D. 1980. See general references. Thanikaimoni, G. 1965. Contribution to the pollen mor-
Hensold, N. 1988. Morphology and systematics of Pae- phology ofEriocaulaceae. Pollen Spores 7: 181-191.
palanthus subgenus Xeractis (Eriocaulaceae). Syst. Bot.
Monogr., Vol. 23, ISO pp.
Hensold, N., Giulietti, A.M. 1991. Revision and redefinition of
the genus Rondonanthus Herzog. Ann. Mo. Bot. Gard. 78:
441-459.
Hoehne, F.C. 1939. Plantas e substancias vegetais t6xicas e
medicinais. Sao Paulo: Graphicars.
Koernicke, Fr. 1854. Eriocaulacearum monographiae sup-
plementum. Linnaea 27: 561-692.
Koernicke, Fr. 1863. Eriocaulaceae. In: von Martius, C.F.Ph.
(ed.) Flora Brasiliensis, Vol. 3, 1.
Koernicke, F. 1867. Eriocaulaceae. Ann. Mus. Bot. Lugd.-Bat.
3:162-164,238-241.
Kral, R. 1989. The genera ofEriocaulaceae of the southeastern
United States. J. Arnold Arbor 70: 131-142.
Malmanche, L.A. 1919. Contribution aI etude anatomique des
eriocaulonacees et des families voisines; restiacees, cen-
trolepidacees, xyridacees, philydracees, mayacacees. Paris:
St. Cloud.
Martius, C.F.Ph. von 1835. Die Eriocaulaceen als selbstiindige
Pfianzenfamilie, aufgestellt und erliiutert. Nova Acta Acad.
Nat. Cur. 17, 1: 1-72.
Monteiro-Scannavacca, W.R., Mazzoni, S.C. 1978. Embryo-
logical studies in Leiothrix fluitans (Mart.) Ruhl.
(Eriocaulaceae). Rev. Brasil. Bot. 1: 59-64.
Monteiro, W.R., Giulietti, A.M., de Moraes Castro, M. 1984.
Aspects of leaf structure of some species of Eriocaulon
(Eriocaulaceae) from Serra do Cip6 (Minas Gerais, Brasil).
Rev. Brasil. Bot. 7: 137-147.
Monteiro, W.R., de Moraes Castro, M., Giulietti, A.M. 1985.
Aspects of leaf structure of some species of Leiothrix Ruh!.
(Eriocaulaceae) from Serra do Cip6 (Minas Gerais, Brasil).
Rev. Brasil. Bot. 8: 109-125.
Morong, Th. 1891. Notes on the North American species of
Eriocauleae. Bull. Torrey Bot. Club 28: 351-362.
Phillips, S.M. 1996. Eriocaulon schimperi (Eriocaulaceae) and
some related species in eastern Africa. Kew Bull. 51: 333-
342.
Poulsen, V.A. 1888. Anatomiske Studier over Eriocaulaceerne.
Vidensk. Medd. Dan Naturhist Foren. 1888: 221-385.
Ramaswamy, S.N., Arekal, G.D. 1982. Embryology of
Eriocaulon xeranthemum Mart. (Eriocaulaceae). Acta Bot.
Neerl. 31: 41-54.
Ramaswamy, S.N., Arekal, G.D., Gaju, M.V.S. 1983. Develop-
mental anatomy of seed coat and pericarp in two species of
Eriocaulon L. (Eriocaulaceae). Bull. Torrey Bot. Club 110:
287-291.
Ruhland, W. 1903. Eriocaulaceae. In: Engler, A. (ed.) Pfian-
zenreich, Vol. 30. Leipzig: Engelmann, 294 pp.
Silveira, A.A. 1908. Flore de Serras Mineiras. Imp. Off. Belo
Horizonte.
Silveira, A.A. 1928. Eriocaulaceae. In: Floralia Montium. Imp.
Off. Belo Horizonte.
Stiitzel, Th. 1984. Bliiten- und infioreszenzmorphologische
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Bot., Vol. 71. Vaduz: Cramer, 108 pp.
208 Flagellariaceae

Flagellariaceae rations or slightly oblique with 1 to 10 trabeculae

0. APPEL and C. BAYER
Flagellariaceae, Dumort., Anal. Fam. Pl.: 59, 60 (1829), nom.
cons.
Perenniallianas from sympodial rhizomes; aerial
shoots solid, glabrous. Leaves alternate, dis-
tichously arranged, glabrous, rolled in bud; sheath
closed, distally with 2 lateral lobes; lamina
lanceolate-acuminate constricted at base, distal
part prolonged into a tendril. Inflorescence termi-
nal, paniculiform. Flowers usually hermaphroditic,
trimerous, (sub-)sessile, without prophylls. Tepals
in 2 whorls, whitish, membranous, subequal,
persistent, their bases shortly fused and some-
what saccate. Stamens in 2 whorls, occasionally
reduced to staminodia; filaments very short,
anthers linear, basifixed tetrasporangiate, laterally
dehiscent by longitudinal slits. Ovary syncarpous,
superior, 3-locular with axile placentation; style
short, divided into 3 spreading, linear stigmatic
branches with papillose upper surface; ovules soli-
tary in each locule, pendulous, bitegmic, almost
atropous, crassinucellate. Fruit drupaceous, red or
black, 1(-2)-seeded; seeds with abundant starchy
endosperm, embryo minute, lenticular.
A monogeneric family with about four species,
widespread throughout the Palaeotropis including
Australia and the Pacific Islands.
(Smithson 1956; Cheadle and Kosakai 1980).
When tips of mature leaves come in contact with
a support, they differentiate into coiled, fibrous
tendrils, otherwise they abscise. In contrast to
many other tendril-bearing plants, the sensitive
portion of the leaf tip is located on the adaxial face
(du Sablon 1887; Arber 1925). Stomata are distrib-
uted on both surfaces of the leaves. They have 2
subsidiary cells and have been referred to the grass
type (Chadefaud 1995; Smithson 1956; Stebbins
and Khush 1961), but lack the dumb-bell-shaped
cells and the perforations between adjacent guard
cells typical of the Poaceae (Sack 1994). Frolich
and Barthlott ( 1988) observed scalelike, unordered
epicuticular wax crystals exclusively adjacent to
the stomatal grooves, similar to those occurring in
Restionanceae. According to Tomlinson (1969)
and Mary et al. (1985), the hypodermis is 1-layered
and indistinct; near the midrib its cells exhibit
sclerotic outer cell walls. Distinct palisade cells are
lacking, the vascular bundles are surrounded by a
fibrous sheath. The mesophyll includes cells con-
taining calcium oxalate crystals. Associated with
vascular bundles, cells with granular silica bodies
and secretory canals are found (Solereder and
Meyer 1929). Sieve-tube plastids contain cuneate
protein crystals and additional less densely packed
crystals and correspond to the subtype Pile', which
is apparently restricted to the order Poales
(Behnke 1981).

VEGETATIVE STRUCTURES. The aerial shoots

(Fig. 51) reach up to 20m in length (Larsen 1972).
According to the interpretation of Tomlinson and
Posluszny (1977), true dichotomy occurs in
Flagellaria indica and axillary branching is
lacking in vegetative aerial shoots. However, the
occurrence of dichotomy was negated by Kircher
(1986), who interpreted this special branching as
due to suppression of the prophyll and the basal
internode of a lateral shoot.
The epidermis cells never contain silica bodies.
The 1- or 2-layered hypodermis consists of slightly
sclerotic cells. A peripheral mechanical layer
between cortex and central cylinder is formed by
vascular bundles embedded in sclerotic tissue
(Tomlinson 1969; Linder 1987). Vessels are found Fig. SlA-D. Flagellariaceae. A-C Flagellaria guineensis. A
in the stems, roots and leaves. The cross-walls of Habit. B Flower. C Part of infructescence. D F. indica, auricu-
vessel elements are horizontal with simple perfo- late leaf base. (After Takhtajan 1982; D orig.)
 Flagellariaceae 209

REPRODUCTIVE STRUCTURES. The inflorescence non-ephemeral antipodals m mature embryo

of Flagellaria indica resembles a panicle (Fig. 51). sacs.
Nevertheless, its main axis and lateral branches of
various order are not terminated by single flowers, POLLEN MORPHOLOGY. The pollen grains of
but by more or less condensed spikes or racemes Flagella ria are spheroidal, 18-22 f!m in diameter,
with short-stalked flowers. In this respect, these and ulcerate with a pore of 3-4 f!m in diameter
inflorescences exhibit basically the same pattern (Fig. 52). The exine is microperforate. The pollen
as those of Poaceae. As in some of the latter, a closely resembles that of Poaceae, from which it
more or less conspicuous thickening may be differs mainly in the somewhat less well-defined
present adaxially on the base of each lateral axis, annulus. The "graminoid" aperture, covered by a
possibly serving to spread the branches. Abaxially, thin endexine layer that extends to some distance
a weaker serial branch is sometimes present. under the margin, is common to Poales. In con-
Usually, the first 2 bracts of lateral axes may be trast to Poaceae, Ecdeiocoleaceae and Anarth-
placed transversally at almost one level, thus re- riaceae, an operculum seems to be absent in
sembling the prophylls of many dicotyledons. Flagellariaceae (Erdtman 1994, 1952; Chanda
In some flowers of Flagellaria indica the num- 1965, 1966; Chanda and Rowley 1967; Ladd 1977;
ber of stamens or carpels is reduced. Occasionally, Linder and Ferguson 1985).
tri-, hexa- or heptasporangiate anthers have been
observed; Subramanyam and Narayana (1972) in- KARYOLOGY. 2n = 38 has been reported for
terpreted them as the result of fusion or reduction various species of Flagellaria (Mangenot and
of microsporangia. Stigmas are Dry with multi- Mangenot 1958, 1962; Chuang et al. 1962; Shetty
cellular papillae (Heslop-Harrison and Shivanna and Subramanyam 1964; Briggs 1966).
1977). Flowers of Flagellaria indica are odorous
(Backer 1951). PHYTOCHEMISTRY. Flagellaria indica is known to
The endocarp of the drupe is composed of stone contain the cyanogenetic glycoside triglochinin
cells forming a homogeneous lignified layer. The (Hegnauer 1986), kaempferol, saponins, ~­
mature seed coat consists of a single layer that sitosterol, and traces of alkaloids (Mary et al. 1985;
corresponds to the thickened, although not ligni- Williams and Harborne 1971, 1988). Harris and
fied, peripheral walls of the outer epidermis of the Hartley (1980) observed the characteristic UV
outer integument; this layer is fused with the fruit fluorescence of cell walls, indicating the presence
wall. Compound starch grains are present in the of bound p-coumaric, ferulic and diferulic acids.
endosperm (Tillich 1996). Flagellaria has been
considered to be wind-pollinated; the fruits are AFFINITIES. Besides Flagellaria, the family was
probably dispersed by birds (Linder 1987). originally considered to include Hanguana and

EMBRYOLOGY. The embryology of Flagellaria

indica was studied by Subramanyam and
Narayana (1972) and Rudall and Linder (1988).
The anther wall includes 2 middle layers; fibrous
thickenings of the endothecium are restricted to
few cells. As in Eriocaulaceae and most Poaceae,
the endothecial thickenings have a complete base
plate (Manning and Linder 1990). The tapetum is
mostly unilayered and corresponds to the secre-
tory type. Microsporocyte division is simulta-
neous; occasionally, large pollen grains (30 X
25 fl) are formed through fusion of2 nuclei. Pollen
is shed at the 2-celled stage (Chadefaud 1995;
Shetty and Subramanyam 1964).
Fast growth of the inner integument leads to
the formation of an endostome. The embryo sac
is deep-seated, free of starch, and develops
Fig. 52. Flagellariaceae. Flagellaria indica, distal pollen view
according to the Allium type. In contrast to the showing raised margin of "graminoid" aperture and indis-
descriptions by Subramanyam and Narayana tinctly scobriculate (perforated) exine. SEM X3600. (Photo
(1972), Rudall and Linder (1988) observed 3 H.P. Linder)
210 Flagellariaceae
]oinvillea, both of which are now generally ac-
cepted as constituting separate families. The rela-
tionships with Hanguanaceae (see there) seem to
be remote at best. In contrast, both Joinvilleaceae
(see there) and Flagellariaceae seem to be best
placed in the Poales. Nevertheless, due to their
crassinucellate ovules, binucleate pollen and lack
of the 6- and 28-kb inversions of the chloroplast
genome (Doyle et al. 1992; Kellogg and Linder
1995), the Flagellariaceae appear to occupy a
somewhat isolated and basal position (Rudall and
Linder 1988; Linder and Rudall 1993; Davis 1995;
Tillich 1996). This is also implicit in most cla-
dograms based on structural and/or molecular,
and combined data (Duvall et al. 1993; Chase et al.
1995; Graham and Barrett 1995; Hahn et al. 1995;
Kellogg and Linder 1995; Linder and Kellogg
1995; Stevenson and Loconte 1995).
DISTRIBUTION AND HABITATS. Flagella ria spe-
cies are distributed throughout tropical Asia,
Africa, N Australia and the Pacific Islands. They
occur in coastal regions and humid, but not con-
stantly swampy lowland forests, forest margins
or mangroves (Linder 1987), occasionally up to
1500 m above sea level.
PALAEOBOTANY. The fossil record of the Poales
starts in the Maestrichtian and suggests a
Gondwanan origin of the order. Due to the simi-
larity of poalean pollen, it is difficult to refer
fossils with certainty to one of the component
families. Flagellaria-type pollen is known from
the Upper Miocene of Borneo (Muller 1981).
Linder {1987) stated that some of the Eurasian
Tertiary material most likely belongs to Fla-
gellaria.
EcoNOMIC IMPORTANCE. This is very minor. The
shoots are used for basket work and to construct
thatches. Some medical uses were summarised by
Mary et al. (1985).
Only one genus:
Flagellaria L. Fig. 51
Flagellaria L., Sp. Pl.: 333 (1753); Backer, Fl. Males. I, 4: 245-
248 (1951), reg. rev.; Smith, Flora Vitensis Nova 1: 283-289
(1970), reg. rev.
Characters as for family.
About four spp., of which F. indica L. is the most
common.
Selected Bibliography
Arber, A. 1925. See general references.
Backer, C.A. 1951. Flagellariaceae. In: Van Steenis, C.G.G.J.
(ed.); Flora Malesiana I, 4. Djakarta: Noordhoff-Kolff, pp.
245-250.
Behnke, H.-D. 1981. Sieve-element characters. Nord. J. Bot. 1:
381-400.
Briggs, B.G. 1966. Chromsome numbers of some Australian
monocotyledons. Contrib. N. S. W. Nat!. Herb. 4: 24-34.
Chadefaud, M. 1955. Remarques sur quelques pollens de
plantes tropicales, particulierement interessants des points
de vue palynologique ou systematique. Rev. Gen. Bot. 62:
641-660.
Chanda, S. 1965. On the pollen morphology of the Flagel-
lariaceae with reference to taxonomy. Trans. Bose Res. Inst.
Calcutta 8: 53-55.
Chanda, S. 1966. On the pollen morphology of the
Centrolepidaceae, Restionaceae and Flagellariaceae, with
special reference to taxonomy. Grana 6: 355-415.
Chanda, S., Rowley, S. 1967. Apertural types in pollen of the
Restionaceae and Flagellariaceae. Grana 7: 16-36.
Chase, M.W. eta!. 1995. See general references.
Cheadle, V.I., Kosakai, H. 1980. Occurrence and specialization
of vessels in Commelinales. Phytomorphology 30: 98-117.
Chuang, T.-1., Chao, C.Y., Hu, W.W.L., Kwan, S.C. 1962. Chro-
mosome numbers of the vascular plants of Taiwan I.
Taiwania 8: 51-66.
Coode, M.J.E. 1978. Flore des Mascareignes 187: Flagel-
lariacees. In: Bosser, J., Cadet, T., Julien, H.R., Marais, W.
(eds.) Flore des Mascareignes: La Reunion, Maurice,
Rodrigues. Mauritius: The Sugar Research Institute.
Davis, G.L. 1966. See general references.
Davis, J.I. 1995. A phylogenetic structure for the monocotyle-
dons, as inferred from chloroplast DNA restriction site
variation, and a comparison of measures of clade support.
Syst. Bot. 20: 503-527.
Doyle, J.J., Davis, J.I., Soreng, R.J., Garvin, D., Anderson, M.J.
1992. Chloroplast DNA inversions and the origin of the
grass family (Poaceae). Proc. Nat!. Acad. U.S.A. 89: 7722-
7726.
Duvall, M.R., Clegg, M.T., Chase, M.W., Clark, W.D., Kress,
W.J., Hills, H.G., Eguiarte, L.E., Smith, J.P., Gaut, B.S.,
Zimmer, E.A., Learn, G.H., Jr. 1993. Phylogenetic hypoth-
eses for the monocotyledons constructed from rbcL se-
quence data. Ann. Mo. Bot. Gard. 80: 607-619.
Erdtman, G. 1944. Pollen morphology and plant taxonomy. II.
Sven. Bot. Tidskr. 38: 163-168.
Erdtman, G. 1952. See general references.
Friilich, D., Barthlott, W. 1988. Mikromorphologie der
epicuticularen Wachse und das System der Monokotylen.
Trop. Subtrop. Pflanzenwelt 63: 1-135.
Graham, S.W., Barrett, S.C.H. 1995. Phylogenetic systematics
of Pontederiales: implications for breeding-system evolu-
tion. In: Rudall, P.J., Cribb, P.J., Cutler, D.P., Humphries,
C.J. (eds.) Monocotyledons: systematics and evolution.
Royal Botanic Gardens, Kew, pp. 415-441.
Hahn, W.J., Givnish, T.J., Sytsma, K.J. 1995. Evolution of the
monocot chloroplast inverted repeat: I. Evolution and phy-
logenetic implications of the ORF 2280 deletion. In: Rudall,
P.J., Cribb, P.J., Cutler, D.P., Humphries, C.J. (eds.) Mono-
cotyledons: systematics and evolution. Royal Botanic Gar-
dens, Kew, pp. 579-587.
Harris, P.J., Hartley, R.D. 1980. See general references.
Hegnauer, R. 1986. See general references.
 Flagellariaceae 211

Heslop-Harrison, Y., Shivanna, K.R. 1977. The receptive Humphries, C.J. (eds.) Monocotyledons: systematics and
surface of the angiosperm stigma. Ann. Bot. (Lond.) 41: evolution. Royal Botanic Gardens, Kew. pp. 543-578.
1233-1258. Subramanyam, K., Narayana, H.S. 1972, pub!. 1975. Some
Kellogg, E.A., Linder, H.P. 1995. Phylogeny of Poales. In: aspects of the floral morphology and embryology of
Rudall P.J., Cribb, P.J., Cutler, D.F., Humphries, C.J. (eds.) Flagellaria indica Linn. In: Murty, Y.S., Johri, B.M., Mohan
Monocotyledons: systematics and evolution. Royal Botanic Ram, H.Y., Varghese, T.M. (eds.) Advances in plant mor-
Gardens, Kew, pp. 511-542. phology. Neerut: Trakashan pp. 211-217.
Kircher, P. 1986. Untersuchungen zur Bliiten- und In- Tillich, H.-J. 1996. Seeds and seedlings in Hanguanaceae and
floreszenzmorphologie, Embryologie und Systematik Flagellariaceae (monocotyledons). Sendtnera 3: 187-197.
der Restionanceen im Vergleich mit Gramineen und Tomlinson, P.B. 1969. Flagellariaceae. In: Metcalfe, C.R. (ed.)
verwandten Familien. Diss. Bot. 94. Anatomy of the monocotyledons 3. Oxford: Clarendon
Ladd, P.G. 1977. Pollen morphology of some members of the Press, pp. 69-82.
Restionaceae and related families, with notes on the fossil Tomlinson, P.B., Posluszny, U. 1977. Features of dichotomiz-
record. Grana 16: 1-14. ing apices in Flagellaria indica (Monocotyledones). Am. J.
Larsen, K. 1972. Flagellariaceae. Flora Thailand 2 (2): 162-166. Bot. 64: 1057-1065.
Linder, H.P. 1987. The evolutionary history of Poales/ Williams, C.A., Harborne, J.B. 1971. Flavonoid patterns in the
Restionales- a hypothesis. Kew Bull. 42: 297-318. monocotyledons. Flavonols and flavons in some families
Linder, H.P., Ferguson, I.K. 1985. On the pollen morphology associated with the Poaceae. Phytochemistry 10: 1059-
and phylogeny of the Restionales and Poales. Grana 24: 65- 1063.
76. Williams, C.A., Harborne, J.B. 1988. Distribution and evolu-
Linder, H.P., Kellogg, E.A. 1995. Phylogenetic patterns in the tion of flavonoids in the monocotyledons. In: Harborne, J.B.
Commelinid clade. In: Rudall, P.J., Cribb, P.J., Cutler, D.F., (ed.) The flavonoids: advances in research since 1980.
Humphries, C.J. (eds.) Monocotyledons: systematics and London: Chapman and Hall, pp. 505-524.
evolution. Royal Botanic Gardens, Kew, pp. 473-496.
Linder, H.P., Rudall, P.J. 1993. The megagametophyte in
Anarthria (Anarthriaceae, Poales) and its implications for
the phylogeny of the Poales. Am. J. Bot. 80: 1455-1464.
Mangenot, S., Mangenot, G. 1958. Deuxieme liste de nombres
chromosomiques nouveaux chez diverses dicotyledones et
monocotyledones d' Afrique occidentale. Bull. Jard. Bot.
Etat 28: 315-329.
Mangenot, S., Mangenot, G. 1962. Enquete sur les nombres
chromosomiques dans une collection d'especes tropicales.
Rev. Cytol. Bioi. Veg. 25: 411-447.
Manning, J.C., Linder, H.P. 1990. Cladistic analysis of patterns
of endothecial thickenings in the Poales/Restionales. Am. J.
Bot. 77: 196-210.
Mary, Z., Pattan Shetty, J.K., Yoganarasimhan, S.N. 1985.
Pharmacognostical studies on Flagellaria indica L.
(Flagellariaceae). J. Econ. Taxon. Bot. 6: 1-8.
Muller, J. 1981. Fossil pollen records of extant angiosperms.
Bot. Rev. (Lancaster) 47: 1-142.
Rudall, P.J., Linder, H.P. 1988. Megagametophyte and nucellus
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1786.
Sablon, L. du 1887. Recherches sur l'enroulement des vrilles.
Ann. Sci. Nat. VII, 5: 5-50, pl. 1-3.
Sack, F.D. 1994. Structure of the stomatal complex of the
monocot Flagellaria indica. Am. J. Bot. 81: 339-344.
Shetty, B.V., Subramanyam, K. 1964. Cytology of Flagellaria
indica Linn. Curr. Sci. 33: 279-280.
Smith, A.C. 1979. Flagellariaceae. In: Smith, A.C. (ed.) Flora
Vitensis Nova. A new flora of Fiji 1. Honolulu: SB Printers,
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Stebbins, G.L., Khush, G.S. 1961. Variation in the organization
of the stomatal complex in the leaf epidermis of mono-
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212
Haemodoraceae
M.G. SIMPSON
Haemodoraceae R. Br., Prodr. 1: 299 (1810), nom. cons.
Erect to decumbent perennials with a sympodial
rhizome or a stolon, corm or bulb. Roots and
subterranean stems often red or reddish. Leaves
basal or cauline, distichous, generally ascending,
straight, rarely bent or twisted; base sheathing and
equitant, blades ensiform, unifacial, lanceolate,
narrowly linear or acicular, hollow-tubular in
Tribonanthes, entire, acuminate, mucronate, or
aristate, glabrous or pubescent, flat, plicate in
Barberetta and Wachendorfia. Inflorescence ter-
minal on short to elongate aerial shoots, scapose
to subscapose, bracteate, consisting of a simple
raceme (Barberetta), a raceme or panicle of 1, 2,
or 3-many flower clusters (Haemodorum), or a
corymb, raceme, panicle or capitulum of simple,
bifurcate, or trifurcate helicoid cymes. Inflores-
cence axes and bracts glabrous or canescent to
tomentose. Flowers variously colored, pedicellate
to subsessile, bisexual, ascending, divergent or
pendulous, actinomorphic or zygomorphic,
trimerous; perianth membranous to coriaceous,
either composed of 3 outer and 3 inner imbri-
cate, ascending to spreading, distinct or basally
fused similar tepals (outer median tepal poste-
riorly positioned in zygomorphic taxa) or (in
Anigozanthos, Blancoa, and Conostylis) perianth
short to long tubular (tube rarely absent) with 6
monocyclic and valvate lobes, the tube splitting at
anthesis along mid-anterior line in Anigozanthos;
tepals or tepal lobes glabrous (Haemodorum,
some Phlebocarya spp.) or canescent, hispidulous
or tomentose externally. Trichomes tapering,
pilate or branched. Stamens 6 or 3, then
opposite inner tepals, distinct, inserted or ex-
serted, free or adnate to perianth tube, equal or
unequal in length and/or position; staminodia
present in Schiekia and Pyrrorhiza; filaments
terete to basally thickened or flattened; anthers
oblong, straight or arcuate (apically mucronate,
apiculate or laminar in some taxa), basifixed, sub-
basifixed or dorsifixed, 2-thecate, 4-sporangiate,
longitudinally and introrsely dehiscent. Gynoe-
cium of 3 fused carpels; ovary superior or
inferior, glabrous or pubescent; ovary 3-locular or
Haemodoraceae
(Phlebocarya) basally 3- and apically 1-locular
or (Barberetta) 1-locular by abortion of latero-
anterior carpels; placentation axile (basal in
Phlebocarya); ovules 1-7 or numerous (ca. 20-50)
per carpel, anatropous or atropous, hypotropous,
pleurotropous or irregularly positioned; style ter-
minal (subapical in Barberetta), terete or flattened
on one side, straight (or curved), stigma oblong,
ovoid or rudimentary, minutely papillate; septal
nectaries present in most taxa. Fruit a !-many-
seeded, loculicidal to apically poricidal capsule,
sometimes indehiscent (in Anigozanthos ful-
iginosus dehiscing along septae into 3 single-
seeded mericarps). Seeds discoid, ellipsoid, ovoid
or globose, smooth to longitudinally ridged,
glabrous or hairy; endosperm starchy, embryo
minute, positioned at micropylar end.
A tropical to temperate family of 13 genera and
about 100 species distributed in eastern and
southeastern N America, Cuba, southern Mexico,
Mesoamerica, northern S America, S Africa, New
Guinea and Australia.
VEGETATIVE MORPHOLOGY. The main subterra-
nean stems bear distichous scalelike or photosyn-
thetic leaves with axillary buds developing into
lateral, propagative, or vegetative branches (Figs.
53, 54). They are rhizomatous (often with prolif-
erative stolons), bulbous (in Haemodorum) or
cormose (in Pyrrorhiza and Tribonanthes). The
underground stems (and sometimes roots) of
Dilatris, Haemodorum, Lachnanthes, Pyrrorhiza,
Schiekia, Wachendorfia and Xiphidium xan-
thorrhizon have a red or orange-red to maroon
color. Aerial stems are often cespitose and un-
branched, terminating in a single scapose to
subscapose inflorescence. In some species of
Conostylis a rhizome gives rise to several sympo-
dial above-ground stolons, each consisting of an
elongate, virgate stem (an internode) terminating
in a cluster of photosynthetic leaves.
Leaves arising from nodes of subterranean
stems are bifacial, distichous, sheathing, scarious
to fibrous and generally rudimentary. Aerial pho-
tosynthetic leaves are basal to sub-basal in all taxa
(except the species of Conostylis with aerial
branches). In all Haemodoraceae the photosyn-
thetic leaves are distichous, unifacial, ascending,
generally straight (distinctly cernuous and dor-
siventral in Xiphidium and tortuous in Conostylis
spp.). The leaf blades are flat, except in
Wachendorfia and Barberetta, where they are pli-
cate, and Tribonanthes, where the leaves are tubu-
lar. Some species of Haemodorum and Conostylis
have subterete or acicular leaves. The margins are
 Haemodoraceae
\~
c~
~
~
F
G~
B
Fig. 53A-K. Haemodoraceae-Haemodoroideae. A Dilatris
pilansii, flower in longitudinal section. B-G Haemodorum
spicatum. B Habit. C Flower, three tepals removed. D Locule
with ovary wall removed to show two ovules on placenta. E
Mature capsule with the persistent tepals and stamens. F Seed,
abaxial view. G Seed in cross-section with endosperm
(stippled) and embryo (white). H Wachendorfia thyrsiflora,
flower, showing one of the two basal pockets. I-K Xiphidium
caeruleum. I Partial inflorescence. J Ovary in longitudinal
section. K Flower. (Orig. M.G. Simpson)
213
~~ B
Fig. 54A-L. Haemodoraceae-Conostylidoideae. A, B Tribo-
nanthes variabilis. A Flower. B Stamen, abaxial (left) and
adaxial view, showing lobed extension of connective. C-E
Conostylis aurea. C Habit. D Flower. E Ovary, longitudinal
section, showing pendulous ovules on apical placentae. F C.
aculeata, ovary in longitudinal section. G Anigozanthos
humilis, flower, abaxial view. H A. manglesii, partial inflores-
cence. I A. fuliginosus, longitudinal section of ovary. J-L A.
flavidus. J Capsule. K Seed. L Same, longitudinal section, with
endosperm (stippled) and embryo (white). (Orig. M.G.
Simpson)
214 Haemodoraceae
entire and ciliate or nonciliate, apices are acute
to aristate, and vestiture ranges from glabrous to
densely tomentose.
VEGETATIVE ANATOMY. (Based on Schulze 1893;
Arber 1925; Stenar 1927, 1938; Green 1959;
Cheadle 1968; Simpson and Dickison 1981; and
Simpson 1990). The sieve-tube plastids contain
cuneate protein bodies but lack starch and
filamental protein. Oxalate raphides are common
in all parts of the plant. Vessels are present in the
roots but absent in the stems of all genera except
Lachnanthes, which has stem vessels with scalari-
form perforation plates. The leaves lack vessels.
Mature roots consist of a uniseriate exodermis, a
parenchymatous cortex, a sclerified endodermis, a
1-celled continuous pericycle, and a relatively
small vascular cylinder. Root hairs arise only from
short cells of the rhizodermis, which alternate
with long cells. The vascular cylinder contains few
(generally 4-5) xylem and phloem strands, the lat-
ter usually containing a single sieve-tube member.
The root pith is sclerified. The vessels have simple
perforation plates, except in Dilatris and Tri-
bonanthes, which have scalariform plates.
The stem contains a !-layered epidermis oflon-
gitudinally elongate cells, a parenchymatous outer
region and a central parenchymatous ground tis-
sue. The rhizomes and stolons - as well as the
aerial stems - have a heavily sclerified cylindrical
sheath (mechanical cylinder, Schulze 1893),
within which is a cylinder of numerous vascular
strands. Stem vascular bundles are collateral to
amp hivasal.
The leaf epidermis is composed of 1 layer of
thin- to thick-walled, longitudinally elongate cells
with straight to undulate anticlinal walls and a
cuticle of variable thickness. Leaf trichomes occur
on the margins and/or surfaces in some taxa and
are mostly similar to floral trichomes (see Floral
Anatomy, below). Stomata are paracytic.
In Dilatris there are spherical epithelial-lined
mucilage chambers in the leaves and ovary. All
(and only) members of Conostylidoideae have dis-
tinctive tannin cells scattered in various plant
organs. These tannin cells are isodiametric to
elongate with a thin tannin-impregnated wall and
numerous, minute tanniniferous spherules just
interior to the wall.
INFLORESCENCE STRUCTURE. The inflorescences
mostly consist of 1 or more helicoid cymes ar-
ranged and aggregated in various patterns. They
are bracteate, the bracts in cymose components
generally appearing to be opposite or perpen-
dicular to the pedicels. Within the subfamily
Haemodoroideae the inflorescence may be a con-
gested corymb of several bifurcate or trifurcate
helicoid cymes (Dilatris and Lachnanthes); a
corymb of simple helicoid cymes (Pyrrorhizia,
Schiekia and Wachendorjia); or a thyrse with lat-
eral helicoid cymes (Xiphidium). The inflores-
cence of Haemodorum consists of a panicle or
simple to branched raceme of either 1, 2, or 3-
many flower clusters, each flower being generally
subtended by 2 bracteoles. The inflorescence of
Barberetta is a "determinate raceme", which
may have been derived by the reduction of a
panicle with lateral helicoid cymes to single
flowers.
In the subfamily Conostylidoideae Conostylis
and Tribonanthes have a congested corymb or ca-
pitulum of a single bifurcate helicoid cyme; in
T. uniflora the inflorescence consists of a single
flower (apparently by reduction, as several vesti-
gial bracts are present). Phlebocarya has a panicu-
late inflorescence consisting of a central main axis
bearing short, lateral axes which terminate in an
ascending bifurcate helicoid cyme pair. Blancoa
has a thyrse with lateral bifurcate pairs of helicoid
cymes having pendulous flowers. Anigozanthos
has either a single terminal simple or bifurcate
helicoid cyme or a panicle of bifurcate helicoid
cymes.
FLOWER STRUCTURE. Flower color ranges from
red, red-orange, yellow-orange, yellow, to green,
white, or black. Flower size varies from 5 mm wide
and long in Barberetta to over 9 em long in
Anigozanthos. The flowers are bisexual, short-
pedicellate to subsessile and may be either actino-
morphic and erect (pendulous in Blancoa) or
zygomorphic and divergent. The external floral
surface ranges from pubescent to tomentose (gla-
brous in Haemodorum and Phlebocarya) with
dendritic, short to long tapering, and/or pilate tri-
chomes. The perianth is trimerous and consists of
6 similar tepals, which are distinct to incompletely
basally connate, or united into a tube with 6 distal
lobes. With the exception of 3 genera (see below),
the perianth is bicyclic and imbricate with 3 outer
and 3 inner lobes. In species with zygomorphic
flowers with this perianth type the outer posterior
tepal is median in position. All but two members
of subfamily Haemodoroideae have free tepals
(Fig. 53). Wachendorfia and Schiekia are similar in
that the outer posterior tepal is basally fused both
to the outer latero-anterior tepals (forming dis-
tinctive pockets or pouches at their junction, Fig.
53H) and to the 2 inner latera-posterior tepals. In
 Haemodoraceae 215

subfamily Conostylidoideae, Phlebocarya and one inner tepal, the inner a stamen. Mature tepals con-
species of Conostylis have free tepals, and all other tain 3 main traces; stamens only a single, generally
genera possess a short to elongate tube (Fig. 54). amphicribral trace. The remaining central vascu-
Anigozanthos, Blancoa, and most Conostylis have lature enters the placentae, forming either a
a perianth consisting of 6 triangular monocyclic branched plexus or 3 pairs of placental bundles.
and valvate lobes both at maturity and during Septal commissures commonly occur near the
early development. The perianth lobes of Tri- apex of the ovary. The style contains both a 3-
bonanthes superficially appear valvate in open lobed stylar canal (contiguous with the locules)
flowers but are clearly imbricate in bud. Stamen and the dorsal carpellary bundles, which traverse
number in the Haemodoroideae is 3 (these oppo- the ovary apex to enter the style.
site the inner tepals) or 1 in Pyrrorhiza (opposite Septal nectaries, which occur in 11 of the 13
the inner anterior tepal). All members of the genera, consist of a flattened or convoluted canal,
Conostylidoideae have 6 stamens. The stamens lined with a uniseriate endothelium. Septal necta-
are always distinct and either free or, in taxa with ries of inferior ovaries open at the ovary apex;
a perianth tube, adnate to the tepals. Narrowly in superior ovaries, septal nectaries are either
lanceolate staminodes occur in both Schiekia, vestigial (Pyrrorhiza), open at the extreme ovary
which has 2 staminodes opposite the outer latera- base (Barberetta), or open in the receptacular
anterior tepals, and in Pyrrorhiza, which has 2 region within specialized basal pockets termed
staminodes opposite the inner latera-posterior "perianth apertures" ( Wachendorfia and Schiekia).
tepals. Anthers are equal or unequal in size and Xiphidium and Phlebocarya lack septal nectaries;
position, generally oblong, basifixed or dorsifixed, Dilatris has only vestigial, apical septal nectaries.
with 2 thecae, 2 locules, and dehisce introrsely by Cells containing oxalate raphide crystals are
longitudinal slits. In some taxa of the Conostyl- common throughout all flower parts. All members
idoideae, the anther is apically mucronate or of the subfamily Conostylidoideae (but no mem-
laminar. Tribonanthes is distinctive in having a bers of the Haemodoroideae) have tanniniferous
flattened, lobed flange extending from the stamen idioblasts, mainly in the placental tissues. The tan-
connective. nin cells are isodiametric to elongate with a thin,
The gynoecium is syncarpous and the ovary is lignified wall and numerous lignified granular
either superior, inferior or half-inferior. The ovary spherules located just interior to the cell wall.
is usually 3-locular but 1-locular in Barberetta by Dilatris is unique in the family in having spherical
abortion of the latera-anterior carpels or basally endothelial-lined mucilage chambers in the tepals
3- and apically 1-locular in Phlebocarya. Placenta- and ovary wall.
tion is axile, but basal in Phlebocarya. Placentae
may be rudimentary or quite large and either EMBRYOLOGY. (Data from Stenar 1927, 1938;
basally surrounding the ovule or peltiform. The Dellert 1933; de Vos 1956; Simpson 1988; and
ovules range in number from 1 to many, are ellip- Steinecke and Hamann 1989). Only a few genera-
soid to ovoid, and in several taxa have a narrow Anigozanthos, Conostylis, and Tribonanthes of the
micropylar extension. The style is terminal (sub- Conostylidoideae and Dilatris, Haemodorum,
apical in Barberetta) and terete, with a minutely Lachnanthes, Wachendorfia and Xiphidium of the
papillate rudimentary, ovoid, or globose stigma. Haemodoroideae - have been investigated em-
bryologically. Anther-wall development is of the
FLORAL ANATOMY. (Data from Simpson and monocotyledonous type. The tapetum is amoe-
Dickison 1981; Simpson 1990, 1993, 1998, boid. Radially oriented, lignified cell wall thicken-
unpubl.). Floral vasculature is basically similar in ings occur in the endothecial and middle layer
all investigated taxa. The pedicel contains numer- cells. Microsporogenesis is successive, resulting in
ous vascular strands arranged in a circle, with or decussate to tetragonal tetrads. Pollen grains are
without a central group of bundles. Three major binucleate at anthesis. Mature ovules are either
strands arise from either the central or peripheral anatropous or atropous and vary in position and
strands of the pedicel. Each bundle divides into 2 shape. The ovules are bitegmic with integuments
radially aligned strands, the inner becoming the generally composed of 2 cell layers and cras-
dorsal carpellary bundle and the outer supplying sinucellate. A parietal cell is present, with a nucel-
the outer tepal and (in taxa with 6 stamens) a lar cap developing in some taxa. Embryo-sac
stamen. Slightly distal to the first 3 bundles, a development follows the Polygonum type. The 3
second set of 3 major bundles branch off; each antipodals are often large and either obpyriform
bundle divides radially, the outer entering an or globose. Endosperm formation is Helobial,
216 Haemodoraceae
with early cell formation in the chalaza} chamber.
The endosperm of ripe seeds has both starch and
protein bodies. Embryo development is probably
of the Asterad or Onagrad type, with the embryo
small relative to the endosperm.
POLLEN MORPHOLOGY. (Data from Erdtman
1952, and Simpson 1983). All eight genera of sub-
family Haemodoroideae have boat-shaped, sul-
cate, heteropolar pollen grains with a verrucose
(in Schiekia foveolate) exine sculpturing. In con-
trast, the pollen grains in all six genera of
Conostylidoideae have a characteristic rugulate
sculpturing and circular apertures. They are either
fusiform with 2 apertures (Anigozanthos, Blancoa,
Conostylis spp., Macropidia), cylindric with 2 ap-
ertures (Phlebocarya), triangular with 3 apertures
(Conostylis spp.), or globose with 7-8 apertures
(Tribonanthes). Haemodoraceae all lack a typical
tectate-columellate exine, a specialized feature for
the family (Simpson 1983). A distinctive 2-layered
exine, lacking a foot-layer, occurs in all Conostyl-
idoideae and in four genera of the Haemo-
doroideae (Barberetta, Dilatris, Wachendorfia,
and Xiphidium). Lachnanthes and Haemodorum
possess a 1-layered exine; Pyrrorhiza has a 2-
layered exine with a granular inner layer; and
Schiekia has a 3-layered exine with a granular
middle layer. An intergradation of exine-wall ar-
chitectural types is evident in all family members
(Simpson 1983). The apertural in tine wall is gener-
ally 2-layered, with a thin inner cellulose layer and
a thicker, largely pectic outer layer traversed by
numerous radially aligned channels or vesicles.
Pollen-wall development was studied for Xiphi-
dium by Simpson (1989).
In Lanaria, Lophiola, and Pauridia, referred
to Haemodoraceae in some recent treatments
(Melchior 1964; Geerinck 1969; Hutchinson 1973),
the pollen grains have a foveolate to reticulate wall
sculpturing in a tectate-columellate exine. Thus,
palynological evidence supports the placement
of these three genera outside Haemodoraceae
(Simpson 1983).
KARYOLOGY. To date, chromosome counts have
been determined for 12 genera and ca. 33 species
of the family. In subfamily Haemodoroideae,
chromosome counts include: n = 12 in Haemo-
dorum; n = 15 in Barberetta and Wachendorfia;
n = 19 in Xiphidium; n = 19-21 in Dilatris; and
n = 24 in Lachnanthes (monotypic). Chromosome
numbers of Pyrrorhiza and Schiekia have not been
determined. In the subfamily Conostylidoideae
chromosome numbers include: n = 6 in Anigo-
zanthos; n = 11 in Tribonanthes and n = 7 in
Phlebocarya; n = 8 in Blancoa; and n = 8 (4, 5, 6,
7, 14, 16, 21, 28) in Conostylis.
POLLINATION. Species of the Australian Anigo-
zanthos, Blancoa, and Macropidia are pollinated
by honey-eater birds of the family Meliphagidae
(Hopper and Burbidge 1978). Conostylis, also
Australian, is pollinated indiscriminately by bees
(Halictus sp. and Apis mellifera L.) and hybridiza-
tion in sympatric populations is common.
A weak incompatibility system seems to occur
in the S African Wachendorfia paniculata, which
is apparently insect-pollinated (Ornduff and
Dulberger 1978).
Buchmann (1980) observed that Xiphidium
caeruleum in Panama is visitated by euglossine
bees, which utilize a "buzz" vibrational mecha-
nism to remove pollen.
Stylar enantiomorphy, in which the style is
strongly oriented to the left or right, occurs in the
S African Barberetta, Dilatris, and Wachendorfia,
in the N American Lachnanthes, and in the S
and Central American Schiekia and Xiphidium
(Ornduff 1974; Ornduff and Dulberger 1978;
Simpson 1990). Buchmann (1980) speculates that
such floral enantiomorphy "may promote greater
outcrossing between flowers (intermorphs) on
different plants, due to increased pollinator
movements."
FRUIT AND SEED. The fruit is usually a dry, locu-
licidal capsule or is indehiscent. It may be oblong,
ellipsoid or ovoid in shape, often strongly 3-lobed
and glabrous to pubescent. In Anigozanthos spp.
the fruit splits partially along the septae into 3
mericarps with the incomplete loculicidalline of
dehiscence resulting in apical pores. In Anigo-
zanthos fuliginosus it dehisces along the septae
into 3 mericarps (cocci) with a hard endocarp,
each containing a single seed. In Haemodorum the
apex of the inferior ovary swells during fruit
development, resulting in what superficially
appears to be a fruit derived from a superior
ovary.
In Dilatris, Haemodorum, Lachnanthes, and
Schiekia (Haemodoroideae) the seeds are gla-
brous, marginally winged and discoid, convex/
concave with the hilum positioned centrally on the
concave side. Pyrrorhiza has discoid seeds with a
central, proximal hilum but with coarse, marginal
trichomes. Seeds of Wachendorfia are globose and
dark-tomentose, those of Xiphidium are globose
and papillate and in Barberetta the seeds are
ovoid, glabrous, and minutely papillate.
 Haemodoraceae 217

In the Conostylidoideae the seeds are small, posed different combinations of genera within
ovoid, or globose and generally glabrous (longitu- tribes or subfamilies. In the present treatment, the
dinally ridged in Anigozanthos). Phlebocarya has Haemodoraceae are divided into two subfamilies:
only a single, globose seed per fruit. Conostylidoideae and Haemodoroideae.
The seeds of all family members have a starchy The subfamily Conostylidoideae is distin-
endosperm and a minute, generally ovoid embryo guished by the occurrence of a rootstock and roots
positioned at the micropylar end. The seed coat that lack a red pigmentation, 6 stamens, isopolar,
is derived from both integuments, each of which rugulate pollen grains with 2-7 circular apertures,
contributes 2 cell layers (additional layers formingand distinctive tannin idioblasts (see Vegetative
near the micropyle). In Wachendorfia the inner Anatomy and Floral Anatomy, above). In contrast,
integument has a water-soluble violet-purple pig- all genera of the Haemodoroideae (except for
ment as in Tacca. Barberetta) possess a rootstock and roots that
have a red pigmentation, 3 stamens, unisulcate,
DISPERSAL AND REPRODUCTIVE BIOLOGY. Fruit heteropolar, and generally verrucate pollen
and seed dispersal are virtually unknown. The grains, and lack tannin idioblasts.
small, dry seeds suggest dispersal by wind and Within Conostylidoideae, Anigozanthos, Blan-
water. Seeds of several genera in the Haemo- coa, and Conostylis presumably make up a
doroideae have a discoid, convex/concave shape monophyletic group, which has been treated as a
and marginal wings, features which may promote separate tribe, Conostylidoideae (Macfarlane et al.
dispersal by water. The fact that almost all genera 1987). In this group Anigozanthos has a unique,
of Haemodoroideae occur in wet or seasonally wet apomorphic floral zygomorphy, caused by split-
habitats may support this hypothesis. ting of the perianth tube at an thesis. Anigozanthos
probably arose from an ancestor of the large genus
PHYTOCHEMISTRY. The Haemodoraceae are Conostylis, which would then be paraphyletic.
chemically unique in being the only family of Also Blancoa seems to have evolved from a para-
vascular plants to possess perinaphthenone pig- phyletic Conostylis. Four species of Conostylis
ments. These compounds are responsible for the share with Blancoa a long, tubular perianth.
red coloration which is prominent in the roots or The relationships of the other two genera of the
rootstocks of some genera. The occurrence of Conostylidoideae, Phlebocarya, and Tribonanthes
perinaphthenone compounds was first reported are unclear. Each of these has been treated as
in Haemodorum corymbosum by Cooke and Segal separate tribes, Phlebocaryeae and Tribonan-
{1955), who named it haemocorin. Subsequently, theae, respectively (Macfarlane et al. 1987).
eight genera and ten species in both subfamilies The relationships within the Haemodoroideae
have been found to contain such pigments are not entirely clear. Based on the most recent
(Hegnauer 1986). phylogenetic analysis (Simpson 1990), Haemo-
Chelidonic acid is reported in Dilatris, Lach- dorum is the most basal genus, sharing some
nanthes, Wachendorfia, Anigozanthos, Blancoa primitive features with members of the subfamily
and Conostylis (Ramstad 1953). Oxalate raphides Conostylidoideae (e.g., in trichome anatomy, simi-
occur in all Haemodoraceae. lar to that of Phlebocarya ciliata). The other genera
Species of Anigozanthos and Wachendorfia were of the Haemodoroideae appear to be united by de-
found to have cell wall-bound p-coumaric and rived floral trichomes, having a basal rosette of 3-4
diferulic acids, thus flourescent under UV light epidermal cells (or evidence of such a rosette).
(Harris and Hartley 1980). The presence of these Dilatris, Wachendorfia, Barberetta, Schiekia,
acids has been shown to be nearly consistent in Pyrrorhiza, and Xiphidium are united in having
taxa outside Liliiflorae proper. They are, for distinctive, derived pilate trichomes. Within this
example, lacking in Cyanella and Tecophilaea, group, all but Dilatris are united in having a supe-
which argues against a close relationship between rior ovary and (with the exception of Xiphidium),
Tecophilaeaceae and Haemodoraceae but in favor infralocular septal nectaries (Simpson 1993).
of a close relationship between Haemodoraceae, Within this superior-ovaried group, Wachendorfia
Philydraceae and Pontederiaceae. and Barberetta are united in having a similar pol-
len ultrastructure (Simpson 1983, 1990) and
SUBDIVISION AND RELATIONSHIPS WITHIN THE Schiekia and Pyrrorhiza are united in having
FAMILY. As reviewed by Geerinck {1968), the staminodes and similarities in ovule anatomy
classification of the Haemodoraceae has been (M.G. Simpson, 1990, unpubl.). The exact relation-
variable and uncertain. Several authors have pro- ship of Xiphidium to these genera is unclear.
218 Haemodoraceae

The genera of Hagenbachia, Lanaria, Lophiola, All Conostylidoideae are restricted to SW Aus-
and Pauridia, which were classified in the tralia. Finally, Haemodorum is widespread in
Haemodoraceae in some recent family treatments Australia (incl. Tasmania) with its limit in New
(see Melchior 1964; Geerinck 1969; Hutchinson Guinea.
1973) are excluded from the family in the present Xiphidium occurs in moist soils, often along
treatment. Hagenbachia is a synonym of Chloro- watercourses. Schiekia is found in mountain
phytum (Anthericaceae) (Ravenna 1977). Lanaria savannas and woodlands, and Pyrrorhiza in
differs from the Haemodoraceae in having tectate- cloud forests. Dilatris and Wachendorjia occur
columellate exine (Simpson 1983) and agrees in sandy, sometimes wet habitats. Members
better with Tecophilaeaceae in embryology of the Conostylidoideae and Haemodorum
(Steinecke and Hamann 1989) and seed morphol- generally occur in open to closed, winter-wet,
ogy and chemistry (Huber 1969). Lophiola lacks periodically burned sandy heath vegetation or
perinaphthenones and has tectate-columellate woodlands.
exine (Simpson 1983). It is now shown to be a
member of Melanthiaceae. Also Pauridia differs EcoNOMIC IMPORTANCE. Underground parts of
from the Haemodoraceae palynologically several Australian species were roasted and eaten
(Simpson 1983) and is retained in Hypoxidaceae by aborigines (Millspaugh 1887). Narcotic effects
(Geerinck 1968, 1969). have been attributed to the eastern N American
Lachnanthes caroliniana (red root), from the
AFFINITIES. Recent evidence supports the very roots of which an invigorating tonic was made by
close relationship between Haemodoraceae, the Aborigines (Millspaugh 1887). The Australian
Philydraceae, Pontederiaceae, and possibly Com- Haemodorum corymbosum produces a red pig-
melinaceae. All investigated members of Phily- ment (called hemocorin), which is said to have
draceae and the subfamily Conostylidoideae of antitumor (Schwenk 1962) and antibacterial activ-
Haemodoraceae and most members of Ponte- ity (Narasimhachari et al. 1968).
deriaceae have tannin idioblasts, a possible Several Australian species of Haemodoraceae,
apomorphy uniting them (see Vegetative including species of Blancoa, Conostylis,
Anatomy and Floral Anatomy, above). Simpson Haemodorum, Tribonanthes and, especially,
( 1983) demonstrated a striking similarity in pollen Anigozanthos are grown horticulturally. Finally,
exine structure between members of the Haemo- Lachnanthes caroliniana is listed as an aggressive
doraceae and Pontederiaceae, arguing for their weed in cranberry bogs (Robertson 1976).
close relationship. In addition to palynological
similarities, these two families are similar (along
with other monocotyledon families, particularly KEY TO THE GENERA
the Philydraceae) in other features. However, the 1. Stamens 3 or 1 (Haemodoroideae) 2
Haemodoraceae and Philydraceae are similar in - Stamens 6 (Conostylidoideae) 9
having derived, ensiform leaves, yet the Phily- 2. Ovary superior 3
draceae and Pontederiaceae are similar in seedling - Ovary inferior 7
3. Stamen 1 5. Pyrrorhiza
morphology (Tillich 1995). Recent molecular - Stamens 3 4
studies (Chase et al. 1993; Duvall et al. 1993; Davis 4. Corolla actinomorphic; inflorescence a simple raceme or a
1995) shows varying interrelationships of the thyrse with lateral cymes 5
Haemodoraceae, Philydraceae, Pontederiaceae, - Corolla zygomorphic; inflorescence a congested corymb of
and Commelinaceae. helicoid cymes or sometimes (Wachendorfia) a thyrse with
lateral cymes 6
5. Inflorescence a simple raceme; functional carpel1; ovule 1;
DISTRIBUTION AND HABITATS. Lachnanthes style subterminal 1. Barberetta
is found in N America, ranging from Nova - Inflorescence an elongate thyrse with lateral helicoid
Scotia in Canada, extending along the coastal cymes; functional carpels 3; ovules 20-30 per carpel; style
terminal 8. Xiphidium
plain of the southeastern USA to Cuba. Xiphidium
6. Stamens unequal, the 2 latera-posterior anthers reduced;
occurs from S Mexico to northern S America, with ovules 3-4 per carpel 6. Schiekia
one species in Cuba. Schiekia occurs in northern - Stamens equal; ovule 1 per carpel 7. Wachendorfia
S America from Venezuela to Guiana. Pyrrorhiza 7. Ovules 1 per carpel 2. Dilatris
is restricted to the isolated Mt. Neblina, - Ovules 2 or more per carpel 8
8. Ovules 2 per carpel; flowers glabrous
Amazonas, Venezuela. Barberetta occurs in 3. Haemodorum
Transkei and Natal, S Africa, while Dilatris and - Ovules 5-7 per carpel; flowers tomentose
W achendorfia are endemic to the Cape, S Africa. 4. Lachnanthes
 Haemodoraceae
9. Perianth dicyclic; tepals imbricate (may superficially
appear valvate in open flowers); flowers glabrous or
white-lanate 10
- Perianth mostly monocyclic; tepals valvate; flowers to-
mentose with dendritic trichomes 11
10. Leaves tubular; inflorescence 1-fiowered or a capitulum of
a bifurcate helicoid cyme; flowers white-lanate
10. Tribonanthes
- Leaves fiat to acicular; inflorescence an open, much-
branched panicle of helicoid cymes; flowers essentially
glabrous 9. Phlebocarya
11. Flowers actinomorphic; perianth tube, if present, not
splitting along mid-anterior line 12
- Flowers zygomorphic; perianth tube splitting along
mid-anterior line 13. Anigozanthos
12. Flowers pendulous; perianth pink-orange 12. Blancoa
- Flowers generally ascending; perianth mostly yellow or
whitish, rarely orange to purplish 11. Conostylis
Subfamilies and Genera of the Haemodoraceae
I. Subfamily Haemodoroideae Dahlgren,
Clifford & Yeo (1985).
Tribe Haemodoreae Benth. (1883}, as Euhaemodoreae.
Rhizomatous, cormose, or bulbous herbs, mostly
with red pigmentation in roots and rootstock;
leaves flat to plicate; flowers actinomorphic or
zygomorphic; stamens 3 or 1; staminodes present
or absent; ovary inferior or superior; functional
locules 3 or 1; ovule 1, 2, 5-7, or numerous per
carpel; seeds glabrous, papillate, or hairy.
1. Barberetta Harv.
Barberetta Harv., Gen. South Afr. Pl., ed 2: 377 (1868}.
Rhizomatous to cormose herbs, ca. 20 em tall;
cormels present in axils of scale leaves; leaves lan-
ceolate, glabrous, plicate; inflorescence a simple
raceme; axis and bracts pilate-hairy; flowers acti-
nomorphic; tepals orange, distinct, imbricate,
spreading, cuspidate; stamens 3, free; ovary supe-
rior functionally unilocular; posteriorly swollen,
glabrous; ovule 1 per ovary, style anteriorly sub-
apical; fruit an irregularly dehiscent, glabrous cap-
sule; seed ovoid, glabrous, minutely papillate; x =
15. Only one sp., B. aurea Harv., Transkei, Natal.
2. Dilatris Bergius Fig. 53A
Dilatris Bergius, Descr. Pl. Cap.: 9 (1767}.
Rhizomatous herbs to 1 m tall; roots and rhizome
red; leaves linear, glabrous; inflorescence a
corymb of 3-5 bifurcate cymes; axes and bracts
219
canescent; flower + actinomorphic; tepals
orange-yellow or mauve, distinct, imbricate,
externally with minute, dark brown, circular
papillae on apical parts; stamens 3, free, unequal,
one anther large on short filament, 2 smaller on
long filaments; ovary inferior, with long
velutinous, and short, pilate hairs; style slightly
curved; ovules 1 per carpel; seeds discoid, convex-
concave, minutely papillose; x = ca. 19-21. Four
spp., Cape S Africa.
3. Haemodorum J.E. Smith Fig. 53B-G
Haemodorum J.E. Smith, Trans. Linn. Soc. (Lond.) 4: 213
(1798}, Macfarlane, Fl. Australia45: 134-148 (1987}.
Bulbous (to cormose) herbs to 1m tall; subterra-
nean stems red; leaves narrowly lanceolate to
acicular with sheath sometimes extending nearly
to apex, glabrous; inflorescence a glabrous panicle
or simple to branched raceme of 1, 2, or 3-many
flowers; flower dark maroon, red to yellow, or
greenish, actinomorphic, glabrous; tepals distinct,
imbricate, ascending, ± coriaceous; stamens 3,
free; ovary inferior; ovules 2 per carpel; fruit 3-
lobed, glabrous, seeds discoid, glabrous, with a
membranous, marginal wing; x = 12. Twenty
spp., Australia (including Tasmania) and New
Guinea.
4. Lachnanthes S. Elliott
Lachnanthes S. Elliott, Sketch Bot. South Carol. Georgia 1: 47
( 1816), nom. cons.; Robertson, J. Arnold Arbor. 57: 205-216
(1976}.
Rhizomatous and stoloniferous herbs to 1m tall;
roots and rhizomes orange-red; leaves linear;
inflorescence a pubescent corymb of trifurcate or
bifurcate cymes; flowers yellow-green to yellow-
white, actinomorphic, externally pubescent; outer
tepals narrowly triangular, with a dark aristate tip,
inner tepals oblong, involute; stamens 3; ovary
inferior; ovules 5-7 per carpel, laterally exserted,
enantiostylous; fruit globose, 3-lobed, pubescent
to basally tomentose; seeds reddish, discoid, con-
vex/concave, minutely scabrate; x = 24. Only one
sp., L. caroliniana (Lam.) Dandy, Cuba, south-
eastern USA and Nova Scotia.
5. Pyrrorhiza Maguire & Wurdack
Pyrrorhiza Maguire & Wurdack, Mem. N.Y. Bot. Gard. 9: 318
(1957}; Maas & Maas-van de Kamer, Fl. Neotr. Monogr. 61
(1993}.
220
Herbs to 1.5m tall with orange-red corms aggre-
gated in clusters; leaves narrowly oblanceolate,
glabrous, coriaceous; inflorescence a corymb of
2-3 ascending helicoid cymes, axes and bracts
tomentose; flowers zygomorphic; perianth basally
orange-red, apically white; tepals distinct, the
outer externally velutinous with pilate hairs; the
inner glabrous; stamen 1, free, opposite inner an-
terior tepal; staminodes 2; ovary superior; ovules
2 per carpel; fruit ellipsoid, flattened on 3 sides,
glabrous; seeds discoid, concave and glabrous on
distal surface but with dense marginal hairs. Only
one sp., P. neblinae Maguire & Wurdack, Mt.
Neblina, Amazonas, Venezuela.
6. Schiekia Meisn.
Schiekia Meisn., Pl. Vase. Gen. 1: 397, 2: 300 (1842); Maas &
Maas-van de Kamer, Fl. Neotr. Monogr. 61 (1993).
Rhizomatous herbs to ca. 1m tall; leaves linear
to narrowly lanceolate, glabrous; inflorescence a
corymb of simple helicoid cymes, axes and bracts
canescent; flowers cream-white; zygomorphic;
tepals imbricate; staminodia 2; stamens 3, free, the
2 latero-posterior stamens with short filaments
and small anthers, anterior stamen relatively long
with filament twisted and anther locules facing the
curved stigma; ovary superior, glabrous; ovules
3-4 per carpel, anatropous; style flattened pos-
teriorly, curved 90° (dextrorse in all material
examined), with stigma laterally facing anterior
anther; capsule widely obovoid, 3-lobed, glabrous;
seeds dark brown, discoid-peltate. Only one sp., S.
orinocensis (Kunth) Meisn., northern S America,
Guyana Highland.
7. Wachendor.fia Burm. Fig. 53H
Wachendorfia Burm., L., Syst. Nat., ed 10, 2: 864 (1759); Helme
& Linder, Bothalia 22: 59-75 (1992).
Rhizomatous herbs to 2.5 m tall, rhizomes red;
leaves linear, entire, glabrous to hirsute, plicate;
inflorescence a raceme of numerous, widely
spaced 3-4-flowered helicoid cymes, axes and
bracts cane scent; flowers ca. 2 em long, zygomor-
phic; tepals yellow to purplish, imbricate; stamens
3, free, anterior stamen curved to side opposite
style; anthers sub-basifixed; ovary superior,
densely pubescent; ovule 1 per carpel, anatropous,
hypotropous; style posteriorly curved, enantio-
morphic; fruit 3-lobed, ovoid, canescent; seeds
ovoid, with a dark brown tomentum. n = 15. Four
spp., sensu Helme, Linder (1992), endemic to the
Cape region of S Africa.
Haemodoraceae
8. Xiphidium Aubl. Fig. 531-L
Xiphidium Aubl., Hist. Pl. Guiane: 33 (1775); Maas & Maas-
van de Kamer, Flora Neotr. Monogr. 61 (1993).
Rhizomatous and stoloniferous herbs to 0.5 m tall;
leaves cernuous, lanceolate, with sheathing base,
glabrous, margins minutely spinulose; inflores-
cence a raceme of helicoid cymes, axes and bracts
pubescent with pilate trichomes; flowers ca.
10 mm wide, actinomorphic; tepals white, distinct,
imbricate, membranous; outer tepals sparsely pu-
bescent at apices and along lower midrib; stamens
3, free; anthers basifixed; ovary superior, very
sparsely pubescent; ovules numerous (c. 30) per
carpel, lanceoloid, atropous, generally hypotro-
pous; style bent anteriorly near apex; capsule
obloid-ovoid, orange, glabrous; seeds dark brown,
globose or irregularly tetrahedric, papillate. n =
19. Two spp., Mexico, Meso-America, northern S
America, and Cuba.
II. Subfamily Conostylidoideae (Lindley)
MacFarlane & Hopper (1987).
Tribe Conostylideae Lindley (1847), as Conostyleae.
Rhizomatous to cormose herbs, without red pig-
mentation in roots and rootstock; leaves flat to
tubular; flowers actinomorphic or zygomorphic;
stamens 6; staminodes absent; ovary inferior
or half-inferior; functionallocules 3; ovules 1, 2, or
numerous per carpel; seed glabrous, smooth or
with longitudinal ridges.
9. Phlebocarya R. Br.
Phlebocarya R. Br., Prodr. 1: 301 (1810); Macfarlane eta!., Fl.
Australia 45: 55-148 (1987).
Rhizomatous cespitose herbs, ca. 0.5 m tall; leaves
linear to acicular, straight to slightly flexuous,
eciliate or ciliate, glabrous or pilose; inflorescence
a panicle of bifurcate helicoid cymes; axes and
bracts glabrescent or lanuginose, trichomes uni-
cellular or dendritic; flower ca. 8-9 mm wide, acti-
nomorphic; tepals white, distinct, imbricate,
glabrous; stamens 6, free; anthers basifixed,
apically mucronate to apiculate; ovary inferior,
ovoid, glabrous; locules 3 below, 1 above by sepa-
ration of septae; placentation basal; ovules 1 per
carpel; capsule indehiscent, ellipsoid, glabrous,
with a hard endocarp; seed 1 per fruit, globose,
glabrous. n = 7. Three spp., endemic to SW
Australia.
 Haemodoraceae
10. Tribonanthes Endl. Fig. 54A,B
Tribonanthes End!. in End!. & Fenzl, Nov. Stirp. Dec.: 27
(1839); Macfarlane et al., Fl. Australia 45: 131-134 (1987).
Cormose herbs, to 40 em tall; leaves narrowly lin-
ear, hollow-tubular, with long open sheath; inflo-
rescence 1-ftowered or a capitulum of a bifurcate
pair of helicoid cymes, axes lanate, bracts gla-
brous; flowers actinomorphic; perianth short-
tubular, white-lanate; perianth lobes imbricate,
divergent, ovate; stamens 6, anthers dorsifixed,
with apical laminar, cernuous appendage; connec-
tive extending as an erect, apically lobed flange;
ovary inferior, globose; ovules numerous (ca. 50
per carpel), anatropous, heterotropous; stigma
ovoid, basally pubescent, with 3 linear sclerous
lobes; capsule indehiscent, widely ellipsoid, white-
lanate; seeds globose-rectanguloid, glabrous. n =
11. Five spp., endemic to SW Australia.
11. Conostylis R. Br. Fig. 54C-F
Conostylis R. Br., Prodr. 1: 300 (1810); Macfarlane et a!., Fl.
Australia 45: 57-110 (1987).
Rhizomatous (some spp. stoloniferous) to erect,
cespitose herbs; leaves basal or cauline, fiat, or
subterete, entire to bristly-ciliate; inflorescence a
capitulum or corymb of bifurcate helicoid cymes;
axes and bracts canescent to tomentose; flowers
yellow, ca. 5-25 mm long, actinomorphic, gener-
ally tomentose with dendritic trichomes; perianth
with a short to elongate tube (tube absent in C.
breviscapa); tepals mostly valvate; stamens 6;
anthers dorsifixed; ovary inferior; placentae oc-
cupying most of locule volume and covered by
numerous ovules or restricted to apex oflocule and
bearing a few pendulous ovules; stigma 3-lobed;
capsule generally ovoid. n = 8 (4, 5, 6, 7, 14, 16, 21,
28). Approximately 45 spp., in six subgenera (sensu
Macfarlane et al. 1987), endemic to SW Australia.
12. Blancoa Lindl.
Blancoa Lind!., in Edw. Bot. Reg., (1839); Macfarlane eta!., Fl.
Australia 45: 110-112 (1987).
Rhizomatous, cespitose herbs to ca. 25 em tall;
leaves narrowly linear to linear-falcate, ciliate,
mucronate, canescent; inflorescence a raceme of
bifurcate helicoid cymes, axes and bracts canes-
cent; flowers pink-orange, 3.5-4cm long, actino-
morphic, tomentose with dendritic trichomes;
perianth tube long, cylindrical; perianth lobes val-
vate; stamens 6; anthers dorsifixed; ovary half-
221
inferior, ovoid; placentae rudimentary at apex of
septae; ovules ca. 3 per carpel; stigma globose,
slightly 3-lobed; capsule globose, tomentose. n =
8. Only one sp., B. canescens Lindl., endemic to SW
Australia.
13. Anigozanthos Labill. Fig. 54G-M
Anigozanthos Labill., in Rel. Voy. Rech. La Perouse 1: 409
(1800) [including Macropidia fuliginosa ]. Drumm ex
Harv.]; Macfarlane eta!., Fl. Australia 45: 112-128 (1987);
Anderberg & Eldenas, Aust. Syst. Bot. 4: 655-664 (1991).
Rhizomatous, often cespitose herbs; leaves lan-
ceolate to linear, mucronate, glabrous to canes-
cent or glaucescent; inflorescence a terminal
simple or bifurcate helicoid cyme or a loose
panicle of generally bifurcate helicoid cymes, axes
and bracts canescent to tomentose; flowers ca. 4-
9cm long, zygomorphic, outer surface with den-
dritic trichomes, inner surface glabrous; perianth
tubular, longitudinally splitting along mid-
anterior line during anthesis, cylindrical to fun-
nelform or flattened by virtue of reftexed tepal
lobes; perianth lobes valvate; stamens 6; ovary in-
ferior, subglobose; placentae ellipsoid to rudimen-
tary; ovules numerous or (in A. rufus and A.
pulcherrimus) 2 per carpel or (in A. fuliginosus) 1
per carpel; stigma globose; capsule ellipsoid, with
transverse wrinkles, splitting along septae and
apically opposite locule; in A. fuliginosus, capsule
dehiscing into 3 1-seeded mericarps surrounded
by hard endocarp; seeds ovoid and glabrous or
ellipsoid with longitudinal ridges. n = 6. Twelve
spp., endemic to SW Australia.
Selected Bibliography
Arber, A. 1925. Monocotyledons. A morphological study.
Cambridge: Cambridge University Press.
Buchmann, S.L. 1980. Preliminary anthecological observa-
tions on Xiphidium caeruleum Aubl. (Monocotyledoneae:
Haemodoraceae) in Panama. J. Kans. Entomol. Soc. 53:
685-699.
Chase, M.W. eta!. 1993. See general references.
Cheadle, V.I. 1968. Vessels in Haemodorales. Phytomor-
phology 18: 412-420.
Cooke, R.G., Segal, W. 1955. Colouring matters of Australian
plants IV. Haemocorin: a unique glycoside from Haemo-
dorum corymbosum Vahl. Aust. J. Chern. 8: 107.
Cronquist, A. 1981. See general references.
Dahlgren, R. 1980. See general references.
Davis, ].I. 1995. A phylogenetic structure of the monocotyle-
dons, as inferred from chloroplast DNA restriction site
variation, and a comparison of measures of clade support.
Syst. Bot. 20: 503-527.
Dellert, R. 1933. Zur systematischen Stellung von Wach-
endorfia. bsterr. Bot. Z. 82: 335-345.
222 Haemodoraceae

De Vas, M.P. 1956. Studies on the embryology and relation- floral ontogeny. International Journal of Plant Sciences. In
ships of South African genera of the Haemodoraceae: press.
Dilatris Berg. and Wachendorfia Burm. J. S. Afr. Bot. 22:41- Simpson, M.G., Dickison, W.C. 1981. Comparative anatomy of
63. Lachnanthes and Lophiola (Haemodoraceae). Flora 171:
Duvall, M.R. et al. 1993. See general references. 95-113.
Erdtman, G. 1952. See general references. Steinecke, H., Hamann, U. 1989. Embryologisch-syste-
Geerinck, D. 1968. Considerations taxonomiques au sujet des matische Untersuchungen an Haemodoraceen. Bot. Jahrb.
Haemodoraceae et de Hypoxidaceae (Monocotyledones). Syst. Ill: 247-262.
Bull Soc. Bot. Belg. 101: 265-278. Stenar, H. 1927. Zur Entwicklungsgeschichte der Gattung
Geerinck, D. 1969. Genera of Haemodoraceae and Hypoxida- Anigozanthos Labill. Bot. Not. 1927: 104-114.
ceae. Bull. Jard. Bot. Nat!. Belg. 39: 47-82. Stenar, H. 1938. Die systematische Stellung der Gattung
Green, J.W. 1959. The genus Conostylis R. Br. I. Leaf anatomy. Xiphidium Loefl. Sven. Bot. Tidskr. 32: 274.
Proc. Linn. Soc. N. S. W. 85: 334-373. Takhtajan , A. 1980. See general references.
Harris, P.J., Hartley, R.D. 1980. Phenolic constituents of the Thorne, R.F. 1981. See general references.
cell walls of monocotyledons. Biochem. Syst. Ecol. 8: 153- Tillich, H.-J. 1995. Seedlings and systematics in mono-
160. cotyldeons. In: Rudall, P.J., Cribb, P.J., Cutler, D.F.,
Hegnauer, R. 1986. See general references. Humphries, C.J. (eds.) Monocotyledons: systematics and
Hopper, S.D., Burbidge, A.H. 1978. Assortative pollination by evolution, Vol. 1. Royal Botanic Garden, Kew.
red wattlebirds in a hybrid population of Anigozanthos
Labill. (Haemodoraceae). Aust. J. Bot. 26: 335-350.
Huber, H. 1969. See general references.
Hutchinson, J. 1973. The families of flowering plants, 3rd edn.,
val. 2, Monocotyledons. Oxford: Clarendon Press.
MacFarlane, T.D., Hopper, S.D., Purdie, R.W., George, A.S.,
Patrick, S.J. 1987. Flora of Australia. Haemodoraceae. Vol.
45: 55-148. Canberra: Australian Government Publishing
Service.
Melchior, H. 1964. Haemodoraceae. In: Engler's Syllabus der
Pflanzenfamilien, 12th edn., Vol. 2. Berlin: Borntraeger.
Millspaugh, C.F. 1887. American medicinal plants. Philadel-
phia: Caxton Press of Sherman.
Narasimhachari, N., Joshi, V.B., Krishnan, S., Panse, M.V.,
Wamburkar, M.N. 1968. Antibacterial properties of
perinaphthenone derivatives. Current Sci. (India) 37: 288.
Ornduff, R. 1974. Heterostyly in South African flowering
plants: a conspectus. J. S. Afr. Bot. 40: 169-187.
Ornduff, R., Dulberger, R. 1978. Floral enantiomorphy and the
reproductive system of Wachendorfia paniculata. New
Phytol. 80: 427-434.
Ramstad, E. 1953. Dber das Vorkommen und die Verbreitung
der Chelidonsaure in einigen Pflanzenfamilien. Pharm. Act.
Helv. 28: 45-57.
Ravenna, P. 1977. Neotropical species threatened and endan-
gered by human activity in Iridiaceae, Amaryllidaceae, and
allied bulbous families. In: Prance, G.T., Elias, T.S. (eds.)
Extinction Is Forever. Bronx, N. Y.: N. Y. Bot. Gard.
Robertson, K.R. 1976. The genera of Haemodoraceae in the
southeastern United States. J. Arnold Arbor. 57: 205-216.
Schulze, R. 1893. Beitrage zur vergleichenden Anatomie der
Liliaceen, Haemodoraceen, Hypoxidoideen und Vellozia-
ceen. Bot. Jahrb. Syst. 17: 295-394.
Schwenk, E. 1962. Tumor action of some quinoid compounds
in the Cheekpouch Test. Arzneim. Forsch. 12: 1143.
Simpson, M.G. 1983. Pollen ultrastructure of the Haemo-
doraceae and its taxonomic significance. Grana 22: 79-103.
Simpson, M.G. 1988. Embryological development of Lach-
nanthes caroliniana (Haemodoraceae). Am. J. Bot. 75:
1394-1408.
Simpson, M.G. 1989. Pollen wall development of Xiphidium
coeruleum (Haemodoraceae) and its systematic implica-
tions. Ann. Bot. II, 64: 257-269.
Simpson, M.G. 1990. Phylogeny and classification of the
Haemodoraceae. Ann. Mo. Bot. Gard. 77: 722-784.
Simpson, M.G. 1993. Septal nectary anatomy and phylogeny in
the Haemodoraceae. Syst. Bot. 18: 593-613.
Simpson, M.G. 1998. Reversal in ovary position from
inferior to superior in the Haemodoraceae: evidence from
 Hanguanaceae
Hanguanaceae
C. BAYER, 0. APPEL, and P.J. RuDALL
Hanguanaceae Airy Shaw, Kew Bull. 18: 260 {1965).
Perennial, dioecious herbs, when young covered
with short branched hairs; aerial shoots ascending
to erect, solid, terete; stolons creeping, sometimes
floating, encircled by appressed, disintegrating
leaf sheaths (Hanguana malayana) or plants
dump-forming with the young shoots directly on
the old stem. Leaves spiralled, with linear to lan-
ceolate blades, at least lower ones long-petioled,
the base forming an open, stem-clasping sheath;
venation of the blade pinnate-parallel, with
tesselate venules; ptyxis supervolute. Inflorescence
terminal, paniculate or sometimes whorled, com-
posed of spikelike partial inflorescences. Flowers
(sub-)sessile, actinomorphic, trimerous, hypogy-
nous. Tepals bractlike, pale yellowish or greenish,
sometimes red-dotted, persistent, shortly connate
at base, arranged in 2 whorls, the outer smaller
than the inner, hood-shaped ones. Staminate flow-
ers with rudimentary ovary which is surrounded
by a 6-lobed, disklike structure; stamens 6, about
as long as inner tepals or longer; filaments filiform
with broader base, inserted on the perianth base;
anthers with short connective, basifixed, introrse,
tetrasporangiate, longitudinally dehiscent. Pistil-
late flowers with 6 staminodes, the inner minute;
ovary ovoid-globose, 3-locular with axile placen-
tation; style very short or stigma sessile, deeply
divided into 3 short, spreading lobes; ovules
solitary in each locule, almost atropous, with
funicular obturator. Fruit a 1( -3)-seeded berry;
seed bowl-shaped, arching over the globose pla-
centa; seed coat sclerified; endosperm copious,
starchy, embryo small.
A monogeneric family with probably five or
more species from tropical SE Asia, Micronesia
and Australia.
VEGETATIVE MORPHOLOGY. Individuals of H.
malayana are highly variable in size and morphol-
ogy (Fig. 55), indicating that there are certainly
more than the two species currently recognised (P.
Boyce, pers. comm.). The taxonomy of this genus
needs further attention, although Backer (1951)
considered the variation as not meriting taxo-
223
nomic rank. Aquatic or swamp-dwelling forms are
robust and up to 2m tall, whereas terrestrial plants
are smaller, up to 1m tall (Larsen 1972, 1983;
Hewson 1986). The capacity to float is due to the
presence of aerenchyma ducts (Backer 1951).
Leaves are densely hairy when very young, with
branched hairs, but in older leaves the hairs re-
main only along the midrib and petiole. Leaves are
petiolate with broad leaf blades which have a char-
acteristic venation pattern, numerous close-set
thin cross-veins linking the more prominent par-
allel longitudinal veins (tesselate venation). The
seedlings have a compact, achlorophyllous cotyle-
don and a foliation sequence starting with reduced
leaves (cataphylls) on the developing shoot, char-
acters considered to be primitive by Tillich (1996).
VEGETATIVE ANATOMY. Branched multicellular
trichomes are present on the leaf and stem sur-
faces. Stomata are tetracytic and more common
on the abaxial than adaxial surface (Smithson
1956; Tomlinson 1969; Baranova 1975). A hypo-
dermis is present beneath both adaxial and abaxial
epidermises, but thicker (1-2-layered) on the
adaxial side. The mesophyll includes a chloren-
chyma of weakly elongated cells and large tannin-
idioblasts. Axially oriented vascular bundles in the
lamina are in 2 layers, a larger adaxial layer and
smaller abaxial layer, with numerous transverse
.
B
c
~D
Fig. SSA-E. Hanguanaceae. Hanguana malayana. A Habit. B
Staminate flower. C Young fruit. D Fruit, longitudinal section.
E Seed, longitudinal section. (After Takhtajan 1982 and
Dahlgren et a!. 1985}
224 Hanguanaceae

veins. In the sheath there is a third adaxial layer of
small inverted bundles (Sill1997). Larger (adaxial)
veins have a parenchymatous or fibrous inner
bundle sheath and endodermoid outer bundle
sheath with unevenly thickened walls. Silica bod-
ies are present in H. malayana but are absent in
specimens of a yet undescribed species (Sill1997).
They are found mainly in the unevenly thickened
outer bundle-sheath cells of larger veins
(Tomlinson 1969) and in the adaxial hypodermal
layer (Solereder and Meyer 1929; Sill1997), not in
the epidermis. The sheath and petiolar region
contain large mucilage channels (Sill1997).
Cross-sections of axes exhibit a loose central
tissue with starch and fibrovascular bundles, an
irregular pericycle, a distinct endodermis of
thick-walled cells similar to those surrounding
single vascular bundles, a narrow parenchymatic,
starch-containing cortex with scattered bundles,
and an epidermis without stomata. Vessels are
present in the roots, but lacking from the aerial
parts (Tomlinson 1969). Perforation plates are
scalariform (Cheadle and Kosakai 1980).
Mucilage hairs are present within ovary locules,
especially surrounding the ovules (Rudall et al.
1997). Ovules are atropous and bitegmic.
EMBRYOLOGY. The tapetum is amoeboid, with a
periplasmodium (C.A. Furness and P.J. Rudall,
in prep.). Microsporogenesis is successive (P.J.
Rudall, unpubl.). Ovules are probably tenuinu-
cellate (i.e. lacking a parietal cell). Megasporogen-
esis is unknown.
PoLLEN MoRPHOLOGY. The pollen grains of
Hanguana are more of less spheroidal, about
19 f.tm in diameter, and inaperturate. The thin,
hyaline exine is provided with small spinules
(Erdtman 1952; Linder and Ferguson 1985).
KARYOLOGY. G. Petersen (unpubl.) found in H.
malayana from Ceylon a large number of chromo-
somes (2n = more than 90), and in an unpub-
lished new species from Sarawak (material from
J. Bogner) 2n = ca. 48. Chromosomes are very
small (less than 1 f.tm).

INFLORESCENCE MORPHOLOGY. The inflores- PoLLINATION. Nectar secretion, which occurs in

cence is a panicle (Backer 1951), composed of 1 both male and female flowers, indicates insect pol-
terminal and several lateral, slender, spikelike lination. According to Maury ( 1888), insects might
partial inflorescences. Each of these basic units be attracted by the more conspicuous yellowish
includes numerous axillary clusters that represent nectaries of male flowers and by the pink ovary of
few-flowered monochasia (H.-J. Tillich, unpubl.). female flowers.
Most of the lateral partial inflorescences are likely
to arise from accessory buds, since they are borne FRUIT AND SEED. The fruit is a 1-seeded berry, up
collaterally in the axil of each subtending leaf. to 2 em long. There is some variation in fruit
The subtending leaves are sessile and gradually colour, from whitish or reddish to almost black.
reduced upward. During seed development the inner integument
disappears and the mature seed coat is a sclerified
FLORAL STRUCTURE. The flowers are small, dioe- meso-endotesta. The starch-containing endo-
cious, actinomorphic and hypogynous (Fig. 55). sperm is enclosed by a single peripheral layer of
Both male and female flowers have 6 tepals in 2 aleuron cells (Tillich 1996).
whorls, each with 3 vascular bundles (P.J. Rudall,
unpubl.). Tepals of female flowers are thicker than AFFINITIES. Hanguana has been associated with
those of male flowers. Filaments of male flowers a wide range of monocotyledons (Maury 1888).
each have a single vascular bundle, and an anther Dahlgren et al. (1985) placed it in Asparagales, but
with 4 locules. Staminodes of female flowers despite some similarities with Astelia (branched
are much reduced, the inner 3 flattened and hairs, short broad styles and mucilage-filled ova-
nectariferous (Maury 1888; Endress 1995; P.J. ries), it now seems certain that Hanguana belongs
Rudall, unpubl.). The gynoecium in male flowers in the commelinoid clade (Chase et al. 1995,
is much reduced, the upper part in 3 separate 1995a). This is strongly supported by molecular
lobes, the basal part lacking locules or ovules, with data from rbcL (Chase et al. 1995), together with
nectariferous lobes opposite the inner stamens the presence of silica bodies in leaves and the pres-
(P.J. Rudall, unpubl.). In female flowers the gyno- ence of ferulic acid in cell walls (Harris and
ecium is large, with a broad trilobed stigma and Hartley 1980; Rudall and Caddick 1994), both con-
very short broad style. The ovary is unilocular in sistent commelinoid characters. Amongst the
the upper part, trilocular in the lower part, with commelinoids, Hanguana has long been associ-
a single ovule per locule and axile placentation. ated with Flagellaria and]oinvillea, more recently
 Hanguanaceae
in a family Flagellariaceae (Backer 1951). How-
ever, as Airy Shaw (1965) pointed out, from pollen
morphology, leaf anatomy and many other char-
acters, a relationship with Flagellaria is unlikely.
A relationship with either palms (Maury 1888;
Airy Shaw 1978, 1981; Chase et al. 1995b) or
Zingiberales (Stevenson and Loconte 1995) is
possible, and these associations are currently
being explored (D.W. Stevenson, H.P. Linder and
P.J. Rudall, unpubl.).
DISTRIBUTION AND HABITATS. Hanguana
ranges from Sri Lanka to SE Asia, Micronesia
(Palau) and N Australia. The plants grow along
rivers and lakes or in lowland swamps and humid
forests up to 1500m, or in freshwater, sometimes
forming floating islands. In some morasses, H.
malayana suppresses all other plants except for
some orchids (Backer 1951).
EcoNOMIC IMPORTANCE. According to Maury
(1888), the roots of "Susum anthelminthicum
Blume", the aquatic form of H. malayana, have
been used as a vermifuge in veterinary medicine.
However, this use has never been confirmed ac-
cording to Backer (1951), who reported that raw
stems and runners of this form are said to be
edible.
Only one genus:
Hanguana Blume Fig. 55
Hanguana Blume, Enum. Pl. Java: 15 (1827).
Susum Blume (1830).
Characters as for family.
Probably five or more spp.
Selected Bibliography
Airy Shaw, H.K. 1965. Diagnoses of new families, new names,
etc., for the seventh edition of Willis's Dictionary. Kew Bull.
18: 261
Airy Shaw, H.K. 1978. Three interesting plants from the
Northern Territory of Australia (Thymelaeaceae, Flacour-
tiaceae and Hanguanaceae). Kew Bull. 33: 1-5.
Airy Shaw, H.K. 1981. A new species of Hanguana from
Borneo. Kew Bull. 35: 819-821.
Backer, C.A. 1951. Flagellariaceae. In: Van Steenis, C.G.G.J.
(ed.) Flora Malesiana I, 4. Djakarta: Noordhoff Kolff, pp.
245-250.
Baranova, M.A. 1975. Stomatographic study of the Flagel-
lariaceae. Bot. Zh. SSSR 60: 1690-1697.
Chase, M.W. et al. 1995, 1995a. See general references.
Cheadle, V.I., Kosakai, H. 1980. Occurrence and specialization
of vessels in Commelinales. Phytomorphology 30: 98-117.
225
Dahlgren, R. et al. 1985. See general references.
Endress, P.K. 1995. Major evolutionary traits of monocot flow-
ers. In: Rudall, P.J., Cribb, P.J., Cutler, D.F., Humphries, C.J.
(eds.) Monocotyledons: systematics and evolution. Royal
Botanic Gardens, Kew, pp. 43-79.
Erdtman, G. 1952. See general references.
Harris, P.J., Hartley, R.D. 1980. See general references.
Hewson, H.J. 1986. Hanguanaceae. In: George, A.S. (ed.) Flora
of Australia 46. Netley: Griffin Press, pp. 172-173.
Larsen, K. 1972. Flagellariaceae; Hanguanaceae. Flora Thai-
land 2(2): 162-166.
Larsen, K. 1983. Hanguanaceae. In: Leroy, J.F. (ed.) Flore du
Cambodge, du Laos et du Viet-nam, part 20. Paris: Museum
National d'Histoire Naturelle, pp. 129-132.
Linder, H.P., Ferguson, I.K. 1985. On the pollen morphology
and phylogeny of the Restionales and Poales. Grana 24: 65-
76.
Maury, P. 1888. Surles affinites du genre Susum. Bull. Soc. Bot.
Fr. 35: 410-417.
Rudall, P.J., Caddick, L.R. 1994. Investigation of the presence
of phenolic compound in monocotyledonous cell walls,
using UV fluorescence microscopy. Ann. Bot. 74: 483-491.
Rudall, P.J., Prychid, C., Jones, C. 1997. Intra-ovarian tri-
chomes in monocotyledons. In: Owens, S.J., Rudall, P.J.
(eds.) Reproductive biology. Royal Botanic Gardens,
Kew.
Sill, E. 1997. Systematische Anatomie der Vegetationsorgane
der Hanguanaceae Airy Shaw (1965) (Monocotyledoneae).
Diploma Thesis. Inst. Syst. Botanik, Universitat Miinchen.
Smithson, E. 1956. The comparative anatomy of the Flagel-
lariaceae. Kew Bull. 11: 491-501.
Solereder, H., Meyer, F.J. 1929. Systematische Anatomie der
Monokotyledonen: 4. Farinosae. Berlin: Borntraeger.
Stevenson, D.W., Loconte, H. 1995. Cladistic analysis of
monocot families. In: Radall, P.J., Cribb, P.J., Cutler, D.F.,
Humphries, C.J. (eds.) Monocotyledons: systematics and
evolution. Kew: Royal Botanic Gardens, Kew, pp. 543-578.
Tillich, H.-J. 1996. Seeds and seedlings in Hanguanaceae and
Flagellariaceae (Monocotyledons). Sendtnera 3: 187-197.
Tomlinson, P.B. 1969. Commelinales-Zingiberales. In:
Metcalfe, C.R. (ed.) Anatomy ofthe monocotyledons, vol. 3.
Oxford: Clarendon Press.
226 Heliconiaceae

Heliconiaceae often becoming cormlike at the soil surface. The

short rhizome branches typically give the plants a
L. ANDERSSON caespitose habit, but some species have rhizome
branches developed as runners. Starch is com-
monly stored in the ground parenchyma of the
rhizome (Tomlinson 1969), but no specialised
storage structures are known.
In most species the only aerial stem is the inflo-
rescence scape. Some species in sects. Zingi-
Heliconiaceae (A. Rich.) Nakai, J. Jpn. Bot. 17: 201 (1941). berastrum, Cannastrum and subgen. Griggsia,
Musaceae tribe Heliconieae K. Schum. (1900). have an elongated, herbaceous stem also in the
Musaceae subfam. Heliconioideae Lane (1955). vegetative phase (Kress 1984; Andersson 1985a).
The aerial shoot is hapaxanthic and ends as a
Rhizomatous herbs, often very tall. Aerial stems strictly terminal inflorescence.
either very short and leaf sheaths then forming a The leaf is usually differentiated into sheath,
pseudostem, or elongate with distinct internodes, petiole and blade, although a distinct petiole is
tightly enveloped by leaf sheaths; buds axillary. sometimes missing. All divisions of the leaf are
Leaves distichous, differentiated into sheath, peti- derived from the proximal (Unterblatt) part of the
ole (sometimes indistinct or lacking) and blade; primordium, the distal (Oberblatt) part giving rise
sheath without distinct ligule; blade entire, with only to a precursory tip, which is usually not
coarse midrib and closely set, parallel, slightly sig- indentifiable in the mature leaf (Kaplan 1973). The
moid lateral veins fusing near margin; lateral leaf sheaths are erect and overlap tightly to form a
veins interconnected by tertiary cross-veins. Inflo- false trunk. Even in those species that have an
rescence terminal, usually on leafy shoots but elongated vegetative axis, the leaf sheaths give the
sometimes on specialized shoots bearing only chief mechanical support to the aerial shoot.
bladeless sheaths, thyrsiform, bearing cincinnate The ptyxis is supervolute. The leaf blade has a
flower clusters in axils of spathaceous bracts on an massive compound midrib from which closely set,
indeterminate main axis. Flowers subtended by parallel secondary veins diverge, which have a
carinate bracteoles, perfect, trimerous and 5- slightly sigmoid course and fuse near the leaf
cyclic, heterochlamydeous; perianth zygomor- margin. Secondary veins are interconnected by
phic, all elements fused to form a basal tube, lat- parallel tertiary ones, which are perpendicular to
eral sepals and petals fused above tube, median the secondary.
sepal free; androecial elements six, the outer me-
dian stamen staminodial, scalelike, inserted on VEGETATIVE ANATOMY. Vegetative anatomy was
top of the basal perianth tube opposite the median described in detail by Tomlinson {1969). Structure
sepal, the remaining 5 stamens fertile, with fila- and development of stomata were described by
ments adnate to the tubular part of the perianth, Tomlinson (1974) and Olatunji (1980). Napp-Zinn
anthers basifixed, elongate, 2-thecate, 4- and Franz ( 1980), and Andersson ( 1981) described
sporangiate. Ovary inferior, 3-locular; placenta- the trichomes.
tion axile; ovules anatropous, subbasal and single Root stele polyarc. Rhizomes with the central
in each locule; style 1, filiform, stigma capitate. cylinder demarcated by an endodermis. Hairs
Septal nectaries present. Fruit drupaceous with 3 often present on aerial parts, multicellular, uni-
operculate pyrenes; seeds exarillate, with thin seriate, often branched. Leaf axis with a main
aleurone-storing perisperm and copious, starch- arc of vascular strands, which are continuous with
and oil-storing endosperm. the major lateral veins of the lamina, several arcs
A family of a single genus, Heliconia, with of smaller and shorter strands, some entirely fi-
perhaps 200 species, occurring throughout the brous, present on the abaxial side of the main arc,
American tropics and Melanesia. a layer of small vascular and fibre bundles present
close to the adaxial surface; with 1 separate system
VEGETATIVE MORPHOLOGY. The plants have a of air canals on each side of the main arc of vascu-
sympodial rhizome with branches terminating in lar bundles. Lamina: epidermal cells with sinous
aerial shoots, and with renewal branches arising anticlinal walls; stomata predominantly polycytic,
near the base of the aerial shoot (Tomlinson 1962). guard cells symmetrical in cross-section; hypoder-
Vegetative buds are axillary (Fisher 1978). Rhi- mis !-layered; mesophyll homogeneous and com-
zome branches are usually short and massive, posed of compact or lobed cells; vascular strands
 Heliconiaceae 227

with a sheath of fibres on either side, only major

vascular bundles connected to the adaxial side.
Silica cells associated with vascular bundles
present in all parts of the plant except the roots;
silica bodies trough-shaped. Raphide sacs present
in the ground tissue of all organs. Tannin present
in morphologically unspecialised cells in all parts
of the plant.

INFLORESCENCE. The main inflorescence axis

is indeterminate. It bears spathaceous bracts
("spathes"), in the axils of which partial inflores-
cences arise (Fig. 56). The spathes are primarily
distichous, but may become spuriously helical due
to twisting of the rachis. They are normally persis-
tent, but caducous or precociously necrotic in a
few species. The flowers are bracteolate.
The partial inflorescences have been variously
interpreted. The classical (and still most com-
monly accepted) interpretation, suggested by
Eichler (1875), is that it is a cincinnus. The whole
inflorescence is then a thyrse with monchasial par-
tial inflorescences. Thompson (1933) and Lane
(1955), however, considered the partial inflores-
cences to have spicoid structure. The classical
view has been supported by Fahn (1953), who con-
cluded that Thompson's interpretation was based
on erroneous observations.

FLORAL STRUCTURE AND ANATOMY. The flowers

have a basic trimerous, pentacyclic organization,
with 2 whorls of tepals, 2 of stamens, and 1 of
carpels. They are oriented with the unpaired sepal
in the anterior position. All the tepals are fused
basally mutually and with the filament bases, to
form a tube, which is usually rather short. Above
this, the median outer tepa! and the filaments are
distinct, whereas the other 5 tepals are further
fused to form a sheath. The median member of the
outer androecial whorl is staminodial and scale-
like and fits into the slit in this sheath, opposite the
free tepa!. In this way, a tube is formed which can
be dilated by a pollinator without being ruptured.
The morphological interpretation of the sta-
minode in Heliconia was questioned by Mello Fig. 56A-E. Heliconiaceae. Heliconia solomonensis. A Inflo-
Filho (1972), who regarded it as a vestigial flower rescence, note pollinating macroglossine bat. B Cincinnal
(!), and by Barreiros, who termed it a ligule bract cut away to show flower at anthesis. (a ovary; b free
(1972a) or apendice liguloide (1972b). Its sepal; c fused sepals). C Ovary with style, stigma, and
staminode. D Staminode. E Pyrenes in ventral and lateral
androecial nature is supported by the occasional view. (Kress 1990b)
formation of abortive sporangia (L. Andersson,
unpubl.).
The gynoecium is formed by 3 carpels. The wards. They have the shape of a triradiate crater,
ovary is inferior and trilocular, with a single basal which is closed apically and opens with 3 pores at
ovule in each locule. Septal nectaries are present the base of the style (Pai and Tilak 1965; Kirchoff
in the axile tissue from the sublocular region up- 1992).
228 Heliconiaceae
EMBRYOLOGY. The embryology of Heliconia was
studied by Mauritzon (1936), and scattered infor-
mation was also reported by Humphrey (1896).
The microsporogenesis has not been described.
The ovule is anatropous, with 2 integuments, the
outer with 6-9, and the inner one with 2 cell layers.
The micropyle is formed by both integuments.
The nucellus is crassinucellate. Megasporogenesis
follows the Polygonum type. No parietal cell
seems to be formed. A circular thickening from
the outer integument, the so-called MikropyHir-
kragen is intiated by periclinal, subepidermal divi-
sions in the outer integument. The nucellar
epidermis becomes palisadelike when the em-
bryosac is mature. No aril is discernible at any
stage. Endosperm formation is Nuclear. The ma-
ture embryo is cylindrical.
POLLEN MORPHOLOGY. Pollen grains are me-
dium-sized {50-IOOj.!m equatorial diameter)
and strongly heteropolar (Fig. 57). They are chara-
cterised by a very reduced exine and elaborate
intine (Kress et al. 1978; Stone et al. 1979; Kress
and Stone 1983). The exine is continuous on the
proximal hemisphere and absent from or discon-
tinuous on the distal one. It is endexinous in na-
ture and varies in sculpturing from psilate to
spinulose. Sculpturing is mostly different on the
proximal and distal hemispheres. The intine is
thick and 2-layered. On the distal hemisphere it
consists mainly of a thick, channelled exintine,
with thin and homogeneous endintine. On the
proximal hemisphere it is unstratified and mainly
endintine. Germination always occurs from the
distal hemisphere, although there is no structur-
ally distinct aperture.
Shape and sculpture are quite variable between
species and very useful taxonomically (e.g. Kress
1984; Andersson 1992).
KARYOLOGY. The chromosome number 2n = 24
was found to be common to 30 taxa of Heliconia
representing all major subdivisions of the genus
(Andersson 1984). Other numbers reported for
this genus seem unreliable. The chromosomes
range between 0.7 and 1.7 j.!m. One pair of chro-
mosomes has satellites.
POLLINATION AND REPRODUCTIVE SYSTEMS.
New World Heliconia are pollinated by humming-
birds (Linhart 1973; Stiles 1975, 1979; Kress
1985a). Plants that grow in large monoclonal
stands in open habitats are mainly visitid by terri-
torial, non-hermit hummingbirds (Trochilinae).
They have short perianths and produce many
Fig. 57. Heliconiaceae. Pollen of Heliconia spathocircinata,
nearly disk-shaped grain, distal hemisphere up. SEM. Bar
10 11m (Photo T. Bergquist)
flowers with copious nectar each day. Plants that
grow in closed forest habitats are visited by
traplining hermit hummingbirds (Phaethor-
ninae). These plants tend to have long, curved
perianths, and to produce few flowers per day with
small amounts of dilute nectar. The different for-
aging behaviours of the two types of humming-
birds result in predominant self-fertilisation in
the former pollination system and predominant
cross-fertilisation in the latter.
The Old World Heliconia solomonensis has been
shown to be pollinated by the macroglossine bat
Melonycteris woodfordi (Kress 198Sb). The inflo-
rescence of this species, like those of all Old World
species, is green, and the flowers are green and
white, odourless, and open in the evening.
Both self-compatibility and self-incompatibility
have been demonstrated in Heliconia (Kress
1983a,b, 198Sb). Of the species examined, the
majority was found to be self-compatible.
FRUIT AND SEED. Fruit and seed structure was
examined by Desler {1994}. The fruit is a drupe
with a rather thin pericarp and a thick, stony en-
docarp formed by sclerenchyma fibres. The
pyrene has a germinal aperture at the mycropylar
end, closed by an operculum. The mature seeds
lack a micropylar collar as well as hypostase. Both
integuments are degraded and compressed. A !-
layered nucellus remnant persists in the mature
seed and contains aleurone but no starch. Both
 Heliconiaceae
starch and oil are stored in the endosperm, starch
in the form of concentric single grains.
DISPERSAL. The fleshy drupes of Heliconia are
dispersed by birds in the neotropics and a wide
variety ofbirds have been reported (Stiles 1979) to
feed on them. Dispersal of fruits of palaeo tropical
species has not been documented; possibly they
are eaten by fructivorous bats. The drupes are pro-
tected by the cincinnal bract until maturity, when
they become exposed by elongation of the pedicel.
At maturity they shift to metallic blue in neo-
tropical species, to red in palaeotropical ones.
RELATIONSHIPS. Heliconia has traditionally been
included in a widely circumscribed Musaceae,
together with Musa, Ensete, Strelitzia, Ravenala,
Phenakospermum and Orchidantha (e.g.,
Schumann 1900; Winkler 1930; Lane 1955). The
relationships of these genera have been differently
judged by different authors (for a summary, see
Tomlinson 1962). In modern literature the most
common approach has been to distribute
them among four families, the Musaceae s. str.
(Musa, Ensete), Strelitziaceae (Phenakospermum,
Ravenala and Strelitzia), Heliconiaceae, and
Lowiaceae (Orchidantha) (e.g. Tomlinson 1962;
Cronquist 1981; Dahlgren et al. 1985). Recent cla-
distic analyses of morphological data (Kress
1990a) are in favour of this. In Kress' analyses,
these four families form a basal grade on the
Zingiberales tree, and Heliconia appears basal in
a clade containing the Cannaceae, Marantaceae,
Zingiberaceae and Costaceae.
DISTRIBUTION. Heliconia is centred inS and Cen-
tral America, with its variation centre along the
Andes from Peru and northwards to Costa Rica.
Four of the five subgenera (Heliconia, Griggsia,
Stenochlamys, and Taeniostrobus) are confined to
the neotropics. The fifth subgenus, Heliconiopsis,
occurs in the western Pacific (Fiji, Samoa, the
Solomon Islands; Kress 1990b).
ECONOMIC IMPORTANCE. Several species and cul-
tivars of Heliconia are popular ornamentals in the
tropics and have, in later years, become commer-
cially important as cut flowers in the international
florist trade (Kress 1990b; Berry and Kress 1991).
In Melanesia, immature, boiled flowers are eaten
in times of food shortage, and the leaves are some-
times used for wrapping and cooking food such as
fish and puddings.
Only one genus:
229
Heliconia L. Fig. 56, 57
Heliconia L. Mant. 2: 147 (1771), nom. cons.; Andersson,
Nord. J. Bot. 1: 218-245 (1981), rev. sect. Heliconia, Opera
Bot. 82: 1-123 (1985), rev. subgen. Stenochlamys, Fl. Ecua-
dor 22: 1-86 (1985), reg. rev., Opera Bot. 111: 1-98 (1992),
rev. subgenera Taeniostrobus and Heliconia; Daniels &
Stiles, Brenesia 15, suppl.: 1-149 (1979), reg. rev. Costa Rica;
Kress, J. Arnold Arbor. 65:429-532 (1984), reg. rev. subgen.
Griggsia, Allertonia 6: 1-58 (1990), rev. subgen.
Heliconiopsis. Heliconiopsis Miq. (1859).
Characters as for family.
About 200 spp. (other estimates occur), most
in tropical America, especially northern parts of
the Andes and southern Central America. Six spp.
indigenous in Melanesia. Several spp. are orna-
mental, and occasionally naturalised.
Selected Bibliography
Andersson, L. 1981. Revision of Heliconia sect. Heliconia.
Nord. J. Bot. 1: 218-245.
Andersson, L. 1984. The chromosome number of Heliconia
(Musaceae). Nord. J. Bot. 4: 191-193.
Andersson, L. 1985a. Revision of Heliconia subgen. Steno-
chlamys. Opera Bot. 82: 1-123.
Andersson, L. 1985b. Musaceae. In: Harling, G., Sparre, B.
(eds.) Flora of Ecuador 22: 1-86.
Andersson, L. 1992. Revision of Heliconia subgen. Taenia-
strobus and subgen. Heliconia (Musaceae-Heliconioideae ).
Opera Bot. 111: 1-98.
Barreiros, H. de Souza 1972a. Heliconia nova brasiliana et
varietas. Morfologia e ecologia - dispersao e poliniza<;ao
(Heliconiaceae (End.) Nakai). Rev. Bras. Biol. 32: 205-208.
Barreiros, H. de Souza 1972b. Especies criticas de Heliconia
L.I: ecologia e morfologia (Heliconiaceae ). Rev. Bras. Biol.
32: 297-302.
Berry, F., Kress, W.J. 1991. Heliconia. An identification guide.
Washington: Smithsonian Institution Press.
Cronquist, A. 1981. See general references.
Dahlgren, R. et al. 1985. See general references.
Daniels, G.S., Stiles, F.G. 1979. The Heliconia taxa of Costa
Rica. Keys and descriptions. Brenesia 15 (Suppl.): 1-149.
Desler, B. 1994. Vergleichende Untersuchung der Anatomie
und Morphologie von Scitamineen-Samen. Unpubl. Di-
ploma Thesis. Kaiserslautern: University of Kaiserslautern.
Eichler, A.W. 1875. Bliithendiagramme construiert und erHiu-
tert Vol. 1. Leipzig: Engelmann.
Fahn, A. 1953. The origin of the banana inflorescence. Kew
Bull. 1953: 299-306.
Fisher, J.B. 1978. Leaf-opposed buds in Musa: their develop-
ment and a comparison with allied Monocotyledons. Am. J.
Bot. 65:784-791.
Humphrey, J.E. 1896. The development of the seed in
Scitamineae. Ann. Bot. 10: 1-40.
Kaplan, D.R. 1973. The problem of leaf morphology and
evolution in monocotyledons. Q. Rev. Biol. 48: 437-457.
Kirchoff, B.K. 1992. Ovary structure and anatomy in the
Heliconiaceae and Musaceae (Zingiberales). Can. J. Bot. 70:
2490-2508.
Kress, W.J. 1983a. Self-incompatibility in Central American
Heliconia. Evolution 37: 735-744.
230 Heliconiaceae

Kress, W.J. 1983b. Crossability barriers in neotropical Heli- Winkler, H. 1930. Musaceae. In: Engler, A., Prantl, K. (eds.)
conia. Ann. Bot. 52: 131-147. Die natiirlichen Pflanzenfamilien, 2nd edn. 15a: 505-541.
Kress, W.J. 1984. Systematics of Central American Heliconia Leipzig: W. Engelmann.
(Heliconiaceae) with pendent inflorescences. J. Arnold
Arbor. 65: 429-532.
Kress, W.J. 1985a. Pollination and reproductive biology of
Heliconia. In: D'Arcy, W.G., Correa, M.D. (eds.) The botany
and natural history of Panama. St. Louis: Missouri Botani-
cal Garden pp. 267-271.
Kress, W.J. 1985b. Bat pollination of an Old World Heliconia.
Biotropica 17: 302-308.
Kress, W.J. 1990a. The phylogeny and classification of the
Zingiberales. Ann. Mo. Bot. Gard. 77: 698-721.
Kress, W.J. 1990b. The taxonomy of Old World Heliconia
(Heliconiaceae). Allertonia 6: 1-58.
Kress, W.J. 1995. Phylogeny of the Zingiberanae: morphology
and molecules. In: Rudall, P.J., Cribb, P.J., Cutler, D.F.,
Humphries, C.J. (eds.) Monocotyledons: systematics and
evolution vol. 2. Royal Botanic Gardens Kew pp. 443-460.
Kress, W.J., Stone, D.E. 1983. Morphology and phylogenetic
significance of exine-less pollen of Heliconia (Heli-
coniaceae). Syst. Bot. 8: 149-167.
Kress, W.J., Stone, D.E., Sellers, S.C. 1978. Ultrastructure of
exine-less pollen: Heliconia (Heliconiaceae). Am. J. Bot. 65:
1064-1076.
Lane, I.E. 1955. Genera and generic relationships in Musaceae.
Mitt. Bot. Staatssamml. Miinch. 13: 114-131.
Linhart, Y.B. 1973. Ecological and behavioral determinants of
pollen dispersal in hummingbird-pollinated Heliconia. Am.
Nat. 107:511-523.
Mauritzon, J. 1936. Samenbau und Embryologie einiger
Scitamineen. Lunds Univ. Arsskr. N.F. Adv. 2 31: 1-31.
Mello Filho, L.E. de 1972. Uma nova interpreta'riio da
morfologia floral de Heliconia L.f. (Musaceae). An. Acad.
Bras. Cienc. 44: 608.
Napp-Zinn, K., Franz, A. 1980. Vergleichend-anatomische
Untersuchungen an petaloiden HochbHittern von Heliconia
revoluta (Griggs) Standley (vorHiufige Mitteilung). Flora
170: 565-569.
Olatunji, O.A. 1980. The structure and development of sto-
mata in some Zingiberales. Notes R. Bot. Gard. Edinb. 38:
499-515.
Pai, R.M., Tilak, V.D. 1965. Septal nectaries in the Scitamineae.
J. Bioi. Sci. 8: 1-3.
Schumann, K. 1900. Musaceae. In: Engler, A. (ed.) Das
Pflanzenreich IV.45: 1-45. Leipzig: Engelmann.
Stiles, F.G. 1975. Ecology, flowering phenology, and hum-
mingbird pollination of some Costa Rican Heliconia spe-
cies. Ecology 56: 285-301.
Stiles, F.G. 1979. Notes on the natural history of Heliconia
(Musaceae) in Costa Rica. Brenesia 15 (Suppl.): 151-180.
Stone, D.E., Sellers, S.C., Kress, W.J. 1979. Ontogeny of exine-
less pollen in Heliconia, a banana relative. Ann. Mo. Bot.
Gard. 66: 701-730.
Takhtajan, A.L. 1982. See general references.
Thompson, J. McLean 1933. Studies in advancing sterility VI.
The theory of Scitaminean flowering. Pub!. Hartley Bot.
Labor. 11: 1-144.
Tomlinson, P.B. 1962. Phylogeny of the Scitamineae - mor-
phological and anatomical considerations. Evolution 16:
192-213.
Tomlinson, P.B. 1969. Commelinales-Zingiberales. In:
Metcalfe, C.R. (ed.) Anatomy of the monocotyledons, Vol.
III. Oxford: Clarendon Press.
Tomlinson, P.B. 1974. Development of the stomatal complex
as a taxonomic character in the monocotyledons. Taxon 23:
109-128.
 Hydatellaceae 231

Hydatellaceae phloem and an endodermis. In the basal part of

the stem vascular bundles are irregularly dis-
u. HAMANN 1 persed in a parenchymatous matrix. In the centre
of the scape 2 small vascular bundles line a central
canal with their xylem portions; likewise, in the
scape of Trithuria submersa 3 collateral bundles
surround a much reduced pith.
Leaf anatomy is similar in both genera. The
leaves have a single central vascular bundle, a
Hydatellaceae Hamann, N.Z.J. Bot. 14: 195 (1976). narrow inner and wider outer parenchymatous
bundle sheath, radiating plates of mesophyll cells
Minute aquatic or subaquatic herbs, annual or alternating with wide air cavities, a 1-layered hy-
rarely perennial. Roots numerous, unbranched. podermis, and an epidermis of elongated cells.
Stem very short, branching to form a leafy tuft, Anomocytic stomata are recorded in Trithuria
often with minute multicellular hairs in the leaf submersa and T. lanterna, but the submerged
axils. Leaves basal, linear to filiform, lacking species of Hydatella apparently lack stomata.
distinct sheaths or ligules, 1-veined, glabrous. Vessels with scalariform perforations occur in
Inflorescence a heterogamous or homogamous the roots and stems. Calcium oxalate and silica
capitulum, terminal, sessile, or scapose, with an seem to be absent.
involucre of 1-4 pairs of opposite, membranous,
glabrous, 1-veined bracts often forming an appar- INFLORESCENCE STRUCTURE. Usually each plant
ent whorl. Flowers unisexual, numerous, minute, bears several flowering scapes each ending in a
lacking perianth and bracteoles. Male flower a terminal capitulum (Fig. 58); rarely there is only a
solitary stamen; anther elliptic to linear, basifixed, single scape. The scapes are usually much shorter
bithecate, tetrasporangiate, dehiscing by lateral than the leaves but elongate when the fruits ripen;
slits. Female flower a solitary unicarpellate pistil; in Trithuria lanterna, T. bibracteata and T.
ovary stalked, unilocular, utriculate, with konkanensis they are suppressed.
(Trithuria) or without (Hydatella) 3 prominent Each capitulum is surrounded by an involucre
ribs; ovule 1, pendulous, anatropous; stigma ter- of 2-4 (rarely more) hyaline or herbaceous, acute
minal, sessile, with 2-10 hairs each a single row of or acuminate bracts. They enclose a variable num-
distended cells. Fruit membranous, dehiscent or ber of male and/or female flowers, each repre-
indehiscent. Seed solitary, ovoid; embryo minute, sented by a single stamen or pistil, respectively. In
peripheral, covered by an operculum; endosperm species with bisexual inflorescences the number
replaced by starchy perisperm. of female flowers exceeds that of male flowers.
A family of two genera and ten species in In species with mainly unisexual inflorescences
Australia, New Zealand and India. sometimes bisexual ones are found. Hydatella
dioica is dioecious, and in the monoecious H.
VEGETATIVE MORPHOLOGY. The germination of inconspicua male flowers are so rare that plants
Trithuria submersa is hypogeal, with the cotyle- may be wholly female.
don remaining inside the testa and not becoming The structure of the capitulum is not completely
green; the primary leaf and the radicle erupt clear. Apart from the involucre there are no bracts,
through the weaker zone that surrounds the and the sequence of floral development needs
operculum of the testa (Cooke 1983b ). reinvestigation. In Hydatella the stamens be-
The plants are completely glabrous with excep- come exserted one after another. According to
tion of the stem base, where between the leaves in Hieronymus {1887), in Trithuria submersa groups
most species characteristic hairs consisting of a of 1 male flower and several female flowers are
single row of thin-walled cells are present. arranged in cincinnate structures.

VEGETATIVE ANATOMY. (From Solereder and
Meyer 1929; Cutler 1969; Cooke 1981). In Hy-
datella filamentosa the vascular tissue of the root
consists of a solitary central vessel surrounded by
1 Deceased.
FLOWER STRUCTURE. Whether the ovary is
monomerous or composed of more than 1 carpel
and hence pseudomonomerous is open to ques-
tion. In Hydatella sometimes besides the main
vascular bundle, which supplies the ovule, 1 or 2
relictual bundles were observed (U. Hamann pers.
observ). The 3 prominent ribs on the ovary wall in
232 Hydatellaceae
Fig. 58A-E. Hydatellaceae. A-C Trithuria submersa. AHabit.
B Capitulum. C Dehiscing fruit with seed visible. D Hydatella
dioica, staminate capitulum. E H. .filamentosa, pistillate ca-
pitulum. (Cooke 1987)
Trithuria and the 3-valved fruit in T. submersa
may also indicate a basically trimerous
gynoecium.
EMBRYOLOGY. Most data are available for
Hydatella inconspicua (Hamann 1975). The en-
dothecium is well developed beneath the fiat, per-
sisting epidermis of the anther. Pollen grains in H.
australis are 2-celled. In both genera the ovule is
bitegmic, crassinucellate, anatropous and pendu-
lous; the raphe is dorsal and the micropyle di-
rected upwards. Both integuments consist of 2 cell
layers and form the micropyle. During seed devel-
opment the apical cells of the inner layer of the
Fig. 59. Hydatellaceae. Trithuria macranthera, sulcate pollen
with spinulate ornamentation, SEM X 3700 (Photo H.P.
Linder)
inner integument give rise to an operculum (see
Fruit and Seed below). The pluricellular chalaza}
part of the nucellus persists and enlarges to form a
starchy perisperm. A small, non-persisting nucel-
lar cap is formed by single periclinal divisions of
the apical nucellus epidermis. The small, roundish
embryo sac is restricted to the apical part of the
nucellus; it has presumably a 3-celled egg appara-
tus and probably 2 polar nuclei, but perhaps no
(or early degenerating) antipodal cells.
Endosperm development is very probably ab
initio cellular. The ripe endosperm is only few-
celled and has no storage function. It surrounds
the minute, lens-shaped, incompletely developed
embryo. A suspensor (if any) is weakly developed.
POLLEN MORPHOLOGY. The pollen grains in
Hydatella dioica (Bortenschlager et al. 1966), H.
australis and Trithuria submersa (Hamann 1975)
are sulcate with a baculate exine (Fig. 59). In
Hydatella inconspicua the aperture is not clear
and the grains frequently adhere in tetrads, but are
often shrunken; this may point to pollen sterility
in this species, which has few male inflorescences
(U. Hamann, pers. observ).
PoLLINATION. Pollen grains germinate on the
sticky surface of the long stigmatic hairs and
the pollen tubes grow inside the hair through the
hollow cells (Yadav and Janarthanan 1995). This
mechanism may facilitate pollination in the sub-
merged species. Anemophily may occur in sub-
aquatic species, e.g. Trithuria submersa. It is
unknown whether autogamy plays a major role,
or if Hydatella inconspicua is apomictic. Pledge
 Hydatellaceae
(1974) found this species flowering 1.15m under
water; according to him and to Edgar (1966),
among several hundred female plants only few
with male or rarely bisexual inflorescences were
found.
FRUIT AND SEED. The minute, ovoid, brownish
seeds are 0.3-0.6 mm long. The main part of the
seed consists of the starchy perisperm, the outer
layer of which has smaller starch grains and is
clearly differentiated from the rest. Between the
perisperm and the small, lenticular embryo lies a
few-celled endosperm. The endotegmen contains
tannin and at the micropylar region forms an
operculum of enlarged cells with thickened walls.
The exotegmic and endotestal layers collapse,
while the exotestal layer assumes a structural
function, with thickened periclinal cell walls in
Hydatella inconspicua and H. filamentosa, and
thickened anticlinal walls in Trithuria submersa.
REPRODUCTIVE BIOLOGY. The seeds of Trithuria
submersa require a wet period in order to break
dormancy, but no stratification; this is typical of
a winter annual adapted to summer-dry climate
(Cooke 1983b).
AFFINITIES. The Hydatellaceae were formerly
included as a tribe - Diplanthereae - in the
Centrolepidaceae (Hieronymus 1887; Gilg-
Benedict 1930), but the profound differences
between them made it necessary to treat these
families separately (Hamann 1975, 1976). The
Hydatellaceae differ conspicuously from the
Centrolepidaceae, i.a., in having dithecate and
basifixed instead of monothecate and dorsifixed
anthers, in having anatropous instead of atropous
ovules, cellular and few-celled instead of nuclear
and pluricellular endosperm, in the presence of
perisperm instead of endosperm, in an exotestal
seed coat with operculum instead of a tegmic
and inoperculate seed coat, in sulcate instead of
ulcerate pollen, in anomocytic instead of paracytic
stomata, in hypogeal instead of epigeal ger-
mination.
Whereas the Centrolepidaceae s. str. have their
nearest relatives evidently in the Restionaceae
and Poaceae, the affinities of the Hydatellaceae are
still obscure, although recently an affinity with
the Typhales has been proposed (Stevenson and
Loconte 1995). Their combination of embryologi-
cal characters - especially the pendulous anatro-
pous ovule, the ab initio cellular endosperm, the
starchy perisperm as storage tissue and the oper-
233
culate seed - is unique among monocotyledons.
The strong structural reduction of the family
makes a phylogenetic placement very difficult,
and molecular evidence is highly desirable.
DISTRIBUTION AND HABITATS. The family is re-
corded from Australia, New Zealand and India,
and may also occur in the intervening Malesian
region. The plants grow submerged or partly sub-
merged in seasonally wet habitats, in swamps, or
on sandy shores of lakes. In W and S Australia
they are adapted to a summer-dry climate by their
winter-annual life cycle. The tropical Trithuria
lanterna is also annual, growing in seasonal
swamps during the wet season and flowering at
the beginning of the dry season (Cooke 1981). In
New Zealand Hydatella inconspicua may grow in
lakes at a depth of more that 1m (Pledge 1974).
KEY TO THE GENERA
1. Capitula heterogamous, with both male and female flowers;
fruit with 3 prominent ribs 1. Trithuria
- Capitula homogamous, with either male or female flowers;
fruit without prominent ribs 2. Hydatella
Genera of the Hydatellaceae
1. Trithuria J.D. Hook. Fig. 58A -C, 59
Trithuria J.D. Hook., Fl. Tasmania 2: 78 (1858); Cooke, Fl.
Austr. 45: 3-6 (1987).
]uncella F. Muell. (1887).
Very small, (sub )aquatic, annual herbs with radi-
cal, linear to filiform leaves, and several capitulate
inflorescences; inflorescences heterogamous;
flowers minute; stigmatic hairs 2-6, attached to
apex of ovary; fruit with 3 prominent ribs. Five
spp., Australia, India.
2. Hydatella Diels Fig. 58D,E
Hydatella Diels, Bot. Jahrb. Syst. 35: 93 (1904); Cooke, Fl.
Austr. 45: 1-3 (1987).
Minute, generally annual, submerged aquatic
herbs with many radical, filiform leaves and
several slender scapes; each scape with a single
headlike terminal inflorescence; inflorescences
homogamous; flowers minute, unisexual; plants
monoecious except H. dioica; stigmatic hairs 4-10,
caducous, attached to persistent boss on apex
of ovary; fruit terete, never ribbed. Five spp.,
Australia, New Zealand.
234 Hydatellaceae/H ydrocharitaceae
Selected Bibliography
Bortenschlager, S., Erdtman, G., Praglowski, J. 1966. Pol-
lenmorphologische Notizen tiber einige Bliitenpflanzen
incertae sedis. Bot. Not. 119: 160-168.
Cooke, D.A. 1981. New species of Schoenus (Cyperaceae) and
Trithuria (Hydatellaceae). Muelleria 4: 299-303.
Cooke, D.A. 1983a. Two Western Australian Hydatellaceae.
Muelleria 5: 123-125.
Cooke, D.A. 1983b. The seedling of Trithuria (Hydatellaceae).
Victorian Nat. 100: 68-69.
Cooke, D.A. 1987. Hydatellaceae. In: George, A.S. (ed.) Flora of
Australia, vol. 45. Canberra: Austral. Governm. Publishing
Service. pp. 1-5.
Cutler, D.A. 1969. Juncales. In: Metcalfe, C.R. (ed.) Anatomy of
the monocotyledons, vol. IV. Oxford: Clarendon Press.
Edgar, E. 1966. The male flowers of Hydatella inconspicua
(Cheesem.) Cheesem. (Centrolepidaceae). N.Z.J. Bot. 4:
153-158.
Edgar, E. 1970. Centrolepidaceae. In: Moore, L.B., Edgar, E.
(eds.) Flora of New Zealand, 2. Wellington.
Gilg-Benedict, Th. 1930. Centrolepidaceae. In: E.-P., Nat.
Pflanzenfam 2nd edn., 15a: 27-33. Leipzig: W. Engelmann.
Hamann, U. 1975. Neue Untersuchungen zur Embryologie
und Systematik der Centrolepidaceae. Bot. Jahrb. Syst. 96:
154-191.
Hamann, U. 1976. Hydatellaceae - a new family of Mono-
cotyledoneae. N.Z.J. Bot. 14: 193-196.
Hamann, U., Kaplan, K., Riibsamen, T. 1979. Dber die
Samenschalenstruktur der Hydatellaceae (Monocotyle-
doneae) und die systematische Stellung von Hydatella
filamentosa. Bot. Jahrb. Syst. 100: 555-563.
Hieronymus, G. 1887. Centrolepidaceae. In: E.-P., Nat.
Pflanzenfam. II, 4: 11-16. Leipzig: W. Engelmann.
Pledge, D.H. 1974. Some observations on Hydatella incon-
spicua (Cheesem.) Cheesem. (Centrolepidaceae). N.Z.J. Bot.
12: 559-561.
Soleder, H., Meyer, F.J. 1929. Systematische Anatomie der
Monokotyledonen. 4. Farinosae. Berlin: Borntraeger.
Stevenson, D.W., Loconte, H. 1995. Cladistic analysis of
monocot families. In: Rudall, P.J., Cribb, P.J., Cutler, D.P.,
Humphries, C.J. (eds.) Monocotyledons: systematics and
evolution, vol. 2. Royal Botanic Gardens, Kew, pp. 543-578.
Yadav, S.R., Janarthanam, M.K. 1995. Trithuria konkanensis
(Hydatellaceae), eine neue Art aus Indien. Aqua-Planta 3:
91-101.
Hydrocharitaceae
C.D.K. CooK
Hydrocharitaceae Juss., Gen. Pl.: 67 (1789), excluding
Nymphaea, Nelumbium [sic], Trapa, Proserpinaca and
Pistia, nom. cons.
Monoecious, dioecious or trioecious, perennial or
annual, aquatic herbs. Roots simple or, in two
genera, branched. Stems monomorphic, dimor-
phic or polymorphic, elongate or contracted,
simple or branched, when branched, then often
complex. Leaves heteroblastic or ontogenetically
constant, spiral, spirodistichous or spiro-
tristichous, distichous or whorled, scalelike or
leafiike, simple; stipules, when present, membra-
nous, translucent, either single and median or
paired and lateral, united below to the petiole;
petioles, when present, short or elongate, rigid or
fiexuose, sometimes sheathing below, winged or
unwinged, armed with spines or unarmed, some-
times not clearly delimited from the leaf blade; leaf
blades, when present, membranous to coriaceous,
submerged, floating or rarely emergent, linear to
orbicular or cordate; venation acrodromous or
reduced to 1 or 3 simple longitudinal veins, cross-
veins often present; margin entire with or without
spines, fiat to undulate or crisped; apex with or
without spines. Squamulae intravaginales usually
2-10 or more, or rarely 1 in each leaf axil. Inflores-
cence scapose or sessile, developing within 1 or 2,
free or united, prophylls (bracts) collectively
called a spathe; spathes submerged or emergent,
membranous to coriaceous, armed or unarmed,
winged or unwinged, persistent or evanescent,
containing 1-100 or more flowers, arranged in 1-
3 monochasia or with more complex systems be-
low the terminal flower; scape when present often
curving or spiralising and thus pulling the devel-
oping fruits under water. Flowers bisexual or uni-
sexual; actinomorphic or with a tendency to
zygomorphy in ontogeny (clear in Nechamandra,
Maidenia and Vallisneria); male flowers borne on
evanescent pedicels or abscising from parent be-
fore anthesis; bisexual flowers often cleistoga-
mous. Sepals 3 or very rarely absent, free,
caducous or persisting in fruit. Petals 3 or less or
occasionally absent, minute to large and showy,
not persisting. Stamens fertile or staminodial, in
 Hydrocharitaceae
up to ± 6 whorls of3 or sometimes more or less, in
some plants reduced to 1; staminodia very vari-
able in number, form and function depending on
pollination mechanism; anthers basifixed or
dorsifixed, with 1-4 microsporangia or by fusion
apparently 6-locular. Nectaries, when present,
usually 3, rarely more, borne at the bases of the
styles or stylodia (probably androecial in origin).
Ovary inferior, sometimes apocarpous, 3-20-(or
more)-carpellate; placentas parietal with or with-
out dissepiments; styluli or style branches 3-20 or
more, simple or forked; stigmas Dry, papillose;
ovules few to numerous; amphitropous, anatro-
pous or atropous, placentation laminar, the ovule
position often varies from ovule to ovule within a
carpel; pistillodium of male flowers when present
usually 3-lobed, the lobes often divided, in ento-
mophilous species often nectarial. Fruit a berry or
a somewhat fleshy or membranous to coriaceous
capsule dehiscing regularly or breaking up at ma-
turity. Seeds ellipsoidal to cylindrical or fusiform,
smooth or with highly complex outer testa, exal-
buminous, or (Ottelia) with scanty endosperm; at
time of germination the embryo often with several
leaves; at germination the seedling usually with a
well-developed hypocotyl and radicle; hypocotyl
often with unicellular hairs.
A cosmopolitan family with 17 genera, of which
3 are marine, and about 76 species.
VEGETATIVE MORPHOLOGY. Seeds of all species
germinate under water; the seedling morphology
has been reviewed by Tillich (1995). Considering
that all members of the family are aquatic, there is
an enormous diversity in growth form. The mor-
phology of the shoot is frequently complex and
has led to considerable controversy (Montesantos
1913; Wilder 1974a,b, 1975; Brunaud 1976, 1977;
Tomlinson 1982; Cook and Ltiond 1982a,b,c, 1983;
Cook and Urmi-Konig 1983a,b,c; Cook et al. 1983;
Yeo et al. 1984; Cook 1985). The genera Apalan-
the, Appertiella, Egeria, Elodea, Hydrilla and
Lagarosiphon (Fig. 60) are essentially similar in
possessing erect, elongate stems with simple,
estipulate and apetiolate leaves borne at more or
less constant distances along the stem. Some
species show stem and leaf polymorphism and
develop horizontal runners which may bear re-
duced, scalelike leaves; in addition some develop
specialised hibernacula which are modified stems
and/or leaves. Nechamandra, Maidenia and some
species of Vallisneria (Fig. 61) and Blyxa are es-
sentially similar to the above group but the leaves
are often unequally spaced along the stem and
there is a tendency to have truncated internodes
235
associated with flowering. This trend is taken to
an extreme in Enhalus, Halophila, Hydrocharis,
Stratiotes, Thalassia and some species of Blyxa
and Vallisneria, where the leaves are borne in con-
tracted rosettes and the rosettes are joined by
leafless or almost leafless runners. This latter
growth form is probably near to the primitive
condition in the Hydrocharitaceae. In most
species of Ottelia (Fig. 62) and some of Blyxa the
stem is monocarpic and remains contracted and
cormlike without developing runners. Thalassia,
Enhalus and some species of Ottelia develop
persistent rhizomes.
The leaves are usually spirodistichous or disti-
chous and usually 1 row is barren while the other
is capable of developing vegetative or floral buds.
The arrangement of branches on species with
whorled leaves is complex and suggests that the
whorled condition is derived from a spirodis-
tichous one. The leaves may be simple and
bandlike or complex with median or lateral
stipules, petioles and blades which vary from lin-
ear to orbicular or cordate. Most species show a
distinct heteroblastic development with linear
juvenile leaves. However, in some genera such
Hydrocharis and Limnobium, the very first leaves
show clearly differentiated blades and traces of
stipules in the petiolar region.
VEGETATIVE ANATOMY. The vegetative anatomy
has been reviewed by Tomlinson (1982), largely
based on the work of Ancibor (1979). Although
relatively few species have been studied in detail,
outstanding features are: the presence of vessels
with oblique scalariform plates in the roots of
some species, the presence of specialised and en-
larged root-hair initial cells and the presence of
inverted vascular bundles in the leaves of some
species. The latter discovery has led to discussion
on the nature of the leaf in monocotyledonous
plants. Contrary to Dahlgren et al. (1985), not
all genera have trichoblasts. Limnobium
possesses strange, layered root caps which are
readily shed.
INFLORESCENCE STRUCTURE. The inflorescence
structure has been reviewed by Kaul (1970) and
Wilder (1974a,b). Limnobium is usually consid-
ered to represent the archetype of the Hydro-
charitaceae: the main axis terminating in a flower
bears 2 (or occasionally 3 or rarely 1) monochasia.
Within the family trends are seen in reduction to
single flowers and to increasing sexual dimor-
phism. A frequent pattern in the family is to have
solitary female flowers and numerous male
236 Hydrocharitaceae

~ c
A

' • F

tJfi
l 0
N 2
~
p

Fig. 60A-P. Hydrocharitaceae. A-C Egeria densa. A Flower-
ing shoot. B Staminate flower. C Pistillate flower. D-F Elodea
canadensis. D Leafy shoot with a staminate flower. E Stami-
nate flower. F Pistillate flower. G-1 Hydrilla verticillata. G
Leafy shoot with pistillate flowers. H Staminate flower
after shedding pollen. I Pistillate flower. J-L Lagarosiphon
muscoides. J Leafy shoot. K Staminate flower at anthesis. L
Pistillate flower. M-P Nechamandra a/ternifolia. M Leafy
shoot with pistillate flowers. N Pistillate flower. 0 Staminate
flower. P Two spathes of staminate flowers showing the
squamulae intravaginales. (Orig. C.D.K. Cook)
 H ydrocharitaceae 237

Fig. 61A-N. Hydrocharitaceae. A-C Maidenia rubra. A
Plant with pistillate flowers. B Staminate inflorescence. C Pis-
tillate flower. D-G Vallisneria spiralis. D Pistillate plant. E
Staminate flower. F Pistillate flower. G Staminate inflores-
cence releasing flower buds. H Enhalus acoroides, pistillate
J.
plant with flower and young fruit. 1-L Thalassia testudinum. I
Sterile shoot. JStaminate flower. K Pistillate flower. L Fruit. M,
N Halophila ovalis. M Plant with pistillate flower. N Staminate
flower. (Orig. C.D.K. Cook)
238 Hydrocharitaceae
Fig. 62A-G. Hydrocharitaceae. A Ottelia a/simoides, habit. B,
C Hydrocharis morsus-ranae. B Habit. C Branched stamens. D,
E Limnobium laevigatum. D Habit. E Staminate flower at an-
thesis. F, G Blyxa aubertii. F Habit. G Inflorescence. (Orig.
C.D.K. Cook)
flowers arranged in complex inflorescences.
Within the inflorescence the individual flowers
are sometimes subtended by bracteoles or the
bracteoles are lacking. The total inflorescence is
borne within a spathe. In Hydrocharis the spathe
of the female inflorescence is reduced to a single
bract, a similar situation is sometimes found in
Limnobium. Normally, the spathe consists of 2
bracts, in Halophila, Hydrocharis, Limnobium and
Stratiotes they are free; in all other genera they are
congenitally united (in Enhalus they separate later
and appear to be free at anthesis).
FLOWER STRUCTURE AND FLORAL ANATOMY.
The family shows a wide range of floral types, re-
flecting a spectacular diversity of pollination
mechanisms which will be dealt with later. Of par-
ticular interest is the morphological and physi-
ological elaboration of staminodia which may play
 Hydrocharitaceae
roles in the visual or olfactory attraction of insects,
the feeding of by nectar secretion or they may
act as sails which aid the transportation of male
flowers. The anthers may be introrse, latrorse or
extrorse; sometimes the anther wall disintegrates;
endothecial fibres are often absent altogether. In
many genera the female flowers are separated
from the gynoecium by a long hypanthium. The
hypanthium often also assumes the function of the
pedicel. There is some evidence, in spite of epi-
gyny, that the carpels may in some genera be
apocarpous. Important accounts of floral anatomy
have been written by Singh {1966a,b), Kaul {1968),
Tomlinson {1969), Cook et al. {1981), McConchie
(1983), McConchie and Kadereit {1987). The seed
coat structure of several genera has been investi-
gated by Shaffer-Fehre {1991).
EMBRYOLOGY. The division of the pollen mother
cell is of the successive type. The pollen grains are
generally 3-celled, more rarely 2-celled when shed.
The ovule is anatropous, bitegmic and crassinu-
cellar (atropous in Nechmandra and Vallisneria)
and the embryo sac of the Polygonum type.
Emdosperm formation is Helobial and free-
nuclear divisions take place in the micropylar
chamber, which, after wall formation, gives rise to
the endosperm proper (Davis 1966).
PoLLEN MoRPHOLOGY. The pollen of all genera
has been recently examined (Cook and Liiond
1982a,b,c, 1983; Cook and Urmi-Konig 1983a,b,c,
1984, 1985; Cook et al. 1983; Cook 1985; and
unpubl.; Takahashi 1994); contrary to some other
published accounts, the pollen is inaperturate,
spherical or nearly so, smooth or sculptured. The
exine is often very thin and reduced. The pollen of
insect-pollinated species is echinate, of sub-
merged-pollinated species smooth and of water-
surface-pollinated species variable but usually
somewhat verrucose. In some species of Elodea
the pollen is liberated in tetrads. In Halophila and
Thalassia it is liberated in moniliform chains held
together by mucilage.
KARYOLOGY. In general, the karyotypes are rela-
tively asymmetrical and usually have large and
small chromosomes. There is considerable polyp-
loidy 2n = 14 to 2n = 132 but also frequent aneu-
ploidy. It is therefore difficult to determine exact
base numbers: x = 7? Enhalus; x = 8 Egeria,
Elodea, Hydrilla; x = 9? Halophila; x = 11
Lagarosiphon, Ottelia. The genus Blyxa probably
has both x = 8 and 9 and V allisneria perhaps x =
8 and 10. For the monotypic genera Nechamandra
239
2n = 16 and 2n = 99, 100 and 101 have been
reported while for Stratiotes 2n = 24, 40-50, 48,
60-64 have been recorded. Within single, well-
defined species there is often considerable varia-
tion, for example, Hydrocharis dubia 2n = 14, 16,
20 and 22; the most extreme species is Ottelia
alismoides, which has 2n = 22, 28, 38, 42, 44, 48,
50, 52, 55, 60, 64, 66, 68, 72, 110 and 132; however,
the somatic numbers 66 (12 counts) and 44 (7
counts) and haploid numbers 33 (8 counts) and 22
(5 counts) are the most frequent. All attempts at
correlating chromosome number with morpho-
logical, ecological or geographical features have
failed (Cook and Urmi-Konig 1983b). Hydrilla
verticillata has 2n = 16,24 and 32. Nakamura and
Kadono {1993), after investigating 263 strains, re-
ported that dioecious strains had either 16 or 24
chromosomes (the triploid was more common);
the monoecious strains were all tetraploid (2n =
24) in Japan; in SE Asia, India and the USA diploid
monoecious strains have been reported. Early
reports of sex chromosomes in Elodea and
Hydrilla are not substantiated by more recent
work.
POLLINATION. The floral biology of the Hydro-
charitaceae is fascinating and sometimes spec-
tacular, Cook {1982) reviewed the subject. There
are 5 major mechanisms:
1. Insect-pollinated with enticement. The flowers
have large showy petals, osmophores and necta-
ries; the male and female flowers are essentially
alike. The genera Apalanthe, Egeria, Hydrocharis
and Ottelia (Fig. 62) have sweet-smelling flowers
while Stratiotes has a foul smell not unlike rotting
meat. Hydrocharis is described in detail by
Scribailo and Poluszny {1983).
2. Insect-pollinated but without enticement. This
mode of pollination is found in some species of
Blyxa, described in detail by Cook et al. {1981).
The flowers lack scent and nectar and show ex-
treme sexual dimorphism. In the male flowers the
pollen is deposited on the adaxial surface of cylin-
drical petals. The female flowers superficially
resemble those of the male but the petals are
reduced to filamentous threads and the stigmas
resemble the petals of the male flowers in form,
colour and position. The petals in the males and
the stigmas in the females exude a fluid and the
wetted pollen is transferred from male to the fe-
male on the feet of carnivorous or scavenging in-
sects. The insect density by pools in the tropics is
often very high and the flowers of Blyxa offer con-
240 H ydrocharitaceae
venient perching places for them. Pollination is
efficient in spite of the fact that the flowers are not
attractive and offer insects no enticement and no
reward.
3. Pollinated at the water surface. This group can
be further subdivided.
3a. Pollen deposited on the water surface. In the
species of Elodea with unisexual flowers the pollen
grains are unwettable and are liberated from the
anthers either singly or in tetrads and come to
float on the surface of the water (Fig. 60D-F). The
female flowers lie on the surface of the water. The
sepals are water-repellent, which causes local
changes in the surface tension, with the result that
pollen is drawn towards the stigmas lying on the
surface. The marine genus Halophila is usually
considered to be pollinated under water but Cox
and Knox (1989) have shown in H. ovalis that the
strings of pollen, although liberated under water,
are negatively buoyant and may be pollinated at
the surface.
3b. Pollen transferred directly from the male
flower to the female. In Appertiella, Enhalus,
Lagarosiphon, Maidenia, Nechamandra and
Vallisneria the male flowers are liberated as buds
from the parent plant. The pedicels of the male
flowers have a well-differentiated abscission layer.
After liberation, the buds float to the surface and
then open in air on the surface of the water. The
sepals and in some species also the petals are
unwettable and reflexed, forming the "hull" of a
small boat. In Appertiella, Lagarosiphon (Fig. 60L)
and Nechamandra the anthers are dorsifixed to
relatively long filaments which are held horizon-
tal. The anther wall is very thin and disintegrates
before anthesis; the pollen grains stick loosely to
the filament and are thus exposed for contact with
the stigma. In the female flowers the perianth seg-
ments are spreading and unwettable and float on
the surface of the water; the stigmas are held above
the surface of the water. Pollination takes place
when the anther comes in contact with a stigma. In
Lagarosiphon erect and papillose staminodia
function as sails transporting male flowers over
considerable distances. In the genera Maidenia
and Vallisneria (Fig. 61E) the male flowers are
very small with 1 or 2 erect stamens. The female
flowers lie horizontally on the surface of the water.
In many botanical texts they are erroneously illus-
trated in a vertical position. Through locally in-
duced changes in the surface tension brought
about by wettable and unwettable parts, they
attract male flowers to the stigmatic region. The
marine genus Enhalus is, in principle, like
V allisneria, except that the petals in the female
flowers are relatively large and water-repellent.
The male flowers are attracted to the water-
repellent petals and as the tide rises or a wave
passes, the male flowers are tipped onto the stig-
mas and enclosed by the unwettable petals within
an air bubble. In Vallisneria americana the male
flowers are also tipped into the females as waves
pass, for details see Svedelius (1932).
4. Pollen transferred through air. In Limnobium
(Fig. 62D,E), described in detail by Cook and
Urmi-Konig (1983c), the male flowers are held
above the females. Pollen is liberated from the
anthers, where it is either directly blown away or
falls onto the spreading sepals where it may be
further transported by wind or fall as the sepals
wither. In Hydrilla (Fig. 60G-I) the pollination is
very specialised and, in detail, very complex (Cook
and Li.iond 1982a). The male flower buds are liber-
ated from the parent plant and, under the right
conditions, open explosively on the water surface.
The pollen is scattered in the air and comes to lie
in a circle of about 20 em or more in diameter on
the water surface. The female flowers look like
inverted bells or wine glasses which are under
water below and open to the air above. The inside
is hydrophobic and the outside is hydrophilic. The
perianth segments are limp which means if the
water level rises or a wave passes, the flower
closes, embracing an air bubble only to open again
later. The stigmas are very small and remain dry.
Only pollen that is released from the anther and
reaches the stigma directly is effective in pollina-
tion. The pollen grains which land on the surface
of the water are lost for reproductive purposes.
5. Pollinated below the water surface. The marine
genera Halophila and Thalassia have wettable pol-
len. In comparison with other Hydrocharitaceae,
the flowers are very reduced with uniseriate peri-
anths or perianth rudiments. The anthers are rela-
tively large and almost sessile. The pollen grains
are spherical but embedded in a gelatinous mass,
apparently of thecal origin and liberated in
moniliform chains. The process of pollination has
been documented for Thalassia testudinum (Fig.
611-M) by Cox and Tomlinson (1988) and for
Halophila ovalis (Fig. 61M,N) by Cox and Knox
(1989); the latter species may also be pollinated at
the surface.
FRUIT AND SEED DISPERSAL. In some species of
Ottelia the fruit dehisces regularly and the seeds
are liberated in a pulpy mass (not unlike a small
 Hydrocharitaceae
banana); this pulpy mass is readily eaten by fish,
who perhaps play or role in dispersal. In most
other genera the fruit dehisces irregularly or is
indehiscent. In Halophila, Hydrocharis, Limno-
bium, and Stratiotes the fruit is somewhat fleshy
and is perhaps eaten by animals. In Hydrilla and
perhaps also in Apalanthe, Appertiella, Elodea and
Lagarosiphon the long, thin few-seeded fruits be-
come separated from the mother plant and repre-
sent the dispersal units. In the remaining genera
the dispersal units are seeds. The seeds may be
smooth or have a very complex testa (Shaffer-
Fehre 1991). Within some species, such as Blyxa
aubertii or B. japonica, the seeds are highly poly-
morphic (Cook and Liiond 1983) and it is interest-
ing that the smooth seeds are dispersed singly
while the spiny seeds are bound together and dis-
persed in groups. In Hydrilla, where the fruit is the
dispersal unit, there is considerable polymor-
phism with smooth, spiny or hooked fruits (Cook
and Liiond 1982a). The embryo is relatively large
with a massive hypocotyl and in some genera,
such as Ottelia and Stratiotes, has several foliage
leaves already developed before germination; it is
usually more or less straight but distinctly curved
in Halophila.
REPRODUCTIVE BIOLOGY. There is an unconvinc-
ing suggestion that Halophila stipulacea may be
agamospermous (Cook 1982). The rest of the fam-
ily is almost certainly amphimictic. Among the
monoecious species there is no evidence of genetic
incompatibility, in fact the opposite seems to be
the case, as many species are highly autogamous
and some even develop specialised cleistogamous
flowers. The number of perfectly dioecious species
is unknown. It is difficult to ascertain, as many
monoecious species develop male and female
flowers on separate branches or at different times.
For example, in Limnobium the 1st foliar inter-
node in a rosette has the epigenetic competence
for developing female inflorescences, the 2nd
internode is barren and the 3rd internode may
develop female or male inflorescences. In appar-
ently perfect dioecious species, occasional bi-
sexual flowers are found, and in some species such
as Ottelia alismoides there is polygamy with fe-
male, male or bisexual plants. In some dioecious
species, such as Blyxa octandra (see Cook et al.
1981) there are often at least 10 male plants to each
female in natural populations.
Several species develop specialised hibernacula
or turions which may allow cold or dry periods to
be bridged; these organs may also play an impor-
tant role in dispersal. However, even without
specialised organs, several species show efficient
241
vegetative spread. Some invading species are rep-
resented by a single sex outside their native range,
for example: Egeria densa - male, Elodea cana-
densis - female, Lagarosiphon major - female,
Stratiotes aloides - female, and Vallisneria spiralis
-female.
The seeds of most species germinate under
water only and have simple, linear or triangular
juvenile leaves; the hypocotyl often possesses spe-
cialised anchoring hairs. The seeds of Hydrocharis
and Limnobium germinate at the water surface; at
first, the testa is shed and the seedling floats on the
surface. The very young seedling looks like a
Lemna or a Salvinia prothallium with the flat,
rounded upper surface emerging above the air-
water interface. In these two genera the first and
all later foliage leaves have broad, expanded leaf
blades. Throughout the family the radicle emerges
first at germination but it is short-lived and soon
replaced by adventitious roots.
PHYTOCHEMISTRY. The family follows the gen-
eral chemical pattern of Alismatales in containing
caffeic acid derivatives, hemicellulosis-bound
apiose, flavones and C-gylcosyl flavones; flavonols
and proanthocyanins are rare, and cyanogenic
compounds recorded from Hydrocharis and
Vallisneria (Hegnauer 1963, 1986). In Elodea fla-
vone glucuronides are specific markers at the level
of species (Mues 1983). Cell wall-bound ferulate is
absent from unlignified cell walls.
RELATIONSHIPS WITHIN THE FAMILY. There
have been various and different treatments, and
even suggestions of splitting the family into five
separate families. There are some clearly delim-
ited phenetic groupings. (1) Limnobium, Hydro-
charis, Ottelia and Stratiotes, called here the
Limnobium group. (2) Vallisneria, Maidenia,
Nechamandra - the Vallisneria group, closely
followed by: (3) Apalanthe, Appertiella, Laga-
rosiphon, Hydrilla, Elodea, Egeria - the Elodea
group. The marine genus Enhalus is usually
placed near Vallisneria because it has a similar
pollination mechanism. Thalassia and Halophila,
also marine, show reductions in many morpho-
logical and anatomical features. Most previous
authors have regarded them as separate and inde-
pendent groups. However, in spite of reductions,
they show features that intimate affinities to the
Limnobium group. Blyxa stands more or less
alone; in mariy ways it could be considered the
archetype for the family.*
*A recent work by Les eta!. (1997) using rbcL sequence data
suggests a different arrangement of genera.
242 H ydrocharitaceae
AFFINITIES. The Hydrocharitaceae are certainly
correctly placed in the Alismatidae. They show
clear affinities to the Butomaceae, Alismataceae,
Aponogetonaceae on one side and the Najadaceae
on the other. Examination of the seed coat in the
Alismatidae by Shaffer-Fehre (1991) and an analy-
sis of DNA sequence data from the chloroplast
gene rbcL by Les et al. (1993) indicate that the
Najadaceae have a much closer relationship to
Hydrocharitaceae than to the other families of the
"N ajadales" (Cymodoceaceae, Potamogetonaceae,
Ruppiaceae, Scheuchzeriaceae, Zannichelliaceae,
Zosteraceae). The decision ofShaffer-Fehre (1991)
to place the genus Najas in the Hydrocharitaceae
is not supported by many other morphological
characters. The family Hydrocharitaceae takes a
somewhat basic and central position among the
Alismatales.
DISTRIBUTION AND HABITATS. The Hydro-
charitaceae are cosmopolitan. Their presence in
New Zealand is due to fairly recent introductions.
The centre of generic diversity is tropical
Asia closely followed by tropical Africa. At the
level of species, the centre is tropical Africa. All
species are obligately aquatic, they need free-
standing water for germination. They are found in
a very wide range of aquatic habitats from oceans
to seasonal pools in all but the very coldest
climatic zones.
PARASITES. The genus Hydrilla has been thor-
oughly investigated for insects that may be valu-
able in biological control. The following feed on
Hydrilla: the moth Paraponyx diminutalis which
also feeds on Vallisneria and Potamogeton, and
Acentropus niveus, a leaf-mining fly Hydrellia
pakistanae, a stem-boring weevil, Bagous cf.
limosus, a tuber-boring weevil, Bagous affinis.
This latter species is apparently endemic to the
hibernacula of Hydrilla and has been used in
biological control (Langland 1990). Stratiotes sup-
ports but is not parasitised by an endemic dragon-
fly (Odonata) Aeschna viridis (Miinchberg 1956),
which lays its eggs in the leaf tissue and the larvae
hunt their prey in the rosettes.
PALAEOBOTANY. The genus Stratiotes has an un-
broken fossil record going back to the Middle
Eocene; there are 15 extinct species which are
summarised by Cook and Urmi-Konig (1983a).
The genus Stratiotes is represented over the last 48
million years by an enormous amount of fossil
material. It is the lack of variation which is sur-
prising; the seeds have increased in length about
1 mm each 7 million years. The genus Stratiotites
Heer from the Tertiary of Switzerland is not
assignable to the Hydrocharitaceae; it is perhaps
a member of the Lythraceae. The genus Cerato-
stratiotes H.-J. Gregor from the Early Miocene of
N Austria is an independent genus now extinct; it
is assigned to the Hydrocharitaceae and is in many
ways like Blyxa; D.H. Les (pers. comm.) is of the
opinion that it belongs to the Ceratophyllaceae.
The genus Hydrochartites C. A. Weber from the
Miocene in N Germany and the species Hydro-
mystria expansa (Heer) Hantke described from
the Tertiary of Switzerland are almost certainly
assignable to Hydrocharis as we know it today -
they closely resemble Hydrocharis dubia, a species
found today in Asia. The fragments reported as
Ottelia from the Eocene of France and the
V allisneria from the Miocene of France are of
uncertain identity.
EcoNOMIC IMPORTANCE. Most genera have in-
vaded new regions in historical times and have
sometimes become serious and pernicious weeds.
Elodea canadensis invaded Europe from N
America and became a serious pest at the end of
the last century and the beginning of this one;
since the 1940s it is being replaced by another ~
American species, E. nuttallii. The S American
species E. callitrichoides has also become estab-
lished in Europe but is not considered to be a pest.
Lagarosiphon major, from southern Africa, has
become a very serious pest in New Zealand and a
local nuisance in Europe, particularly in Germany
and Switzerland. Hydrilla verticillata, an Asian
species, has become established in many regions
of the world and has become a very serious pest in
the southern parts of the USA. Egeria densa from
temperate S America has become a local nuisance
inN America, Europe, Australia and New Zealand.
Hydrocharis morsus-ranae from Europe is spread-
ing actively in eastern Canada at the moment and
is blocking canals and side waters. Asiatic species
of Blyxa and Ottelia have established themselves
as weeds in rice fields and irrigation ditches in
Africa, Europe and N America; however, they are
not considered to be serious pests. The genus
Vallisneria from southern Europe is spreading in
northern Europe in heated effluents but has not
yet been reported to be a pest; in parts of India
where it is native it is a serious pest in irrigation
canals. The marine species Halophila stipulacea
entered the Mediterranean Sea via the Suez Canal,
soon after the latter was opened in 1869; it is still
spreading eastwards, but is not considered to be
undesirable.
 Hydrocharitaceae
Most species are of considerable importance as
shelter or food for the local fauna. The leaves and
inflorescences of Ottelia, especially 0. cordata, are
offered for sale in markets for human consump-
tion is SE Asia. Most other species are used locally
as green manure or as fodder for livestock.
KEY TO THE GENERA
1. Leaves clearly differentiated into petiole and blade when
mature
- Leaves never differentiated into petiole and blade
2. Stipules absent; branched roots absent
- Stipules present (lateral or median); branched roots
usually present
3. Leaves much more than 3crn long; sterns contracted and
corrnlike; perianth clearly biseriate with sepals and petal.s;
freshwater 1. Ottella
- Leaves rarely exceeding 3 ern long; sterns elongate and
branched; perianth uniseriate or absent; marine
17. Halophila
4. Petals large and showy, clearly exceeding the sepals in
length· inner stamens adnate in pairs. Old World
' 3. Hydrocharis
- Petals not large and showy, scarcely exceeding the sepals
in length; stamens not united in pairs. New World
4. Limnobium
5. Leaves arranged in rosettes or in bunches on contracted
internodes
- Leaves arranged in regular and repeating patterns along
elongated sterns
6. Leaves rigid, narrowly triangular to linear, with conspicu-
ous, hard spines on the margins; entire plant rising to the
water surface to flower and then sinking and perennating
under water; spathe of 2 free bracts 2. Stratiotes
- Leaves flaccid, without hard spines on the margins (mar-
gins sometimes minutely serrate or if spiny then spines
soft); entire plant not rising to the water surface to flower;
spathe of 2, united bracts . .
7. Plants rhizomatous (horizontal axes wholly buned m the
substrate, relatively thick, fleshy and with short intern-
odes); marine
- Plants not rhizomatous (horizontal axes if present not
wholly buried in the substrate, thin or, if thick and _fleshy,
then short, rarely more than 5 ern long); rnanne or
freshwater
8. Rhizomes monomorphic, bearing foliage leaves at the
apex; flowers rising to the water surface for anthesis; leaf
veins in 2 series, the adaxial series inverted 15. Enhalus
- Rhizomes dimorphic, with extended, horizontal, scale
bearing long shoots and erect, foliage bearing short shoots;
flowers wholly submerged; leaf veins in a single series
16. Thalassia
9. Marine; each internode of the horizontal axis developing 2,
equally sized, scalelike leaves, one embracing the axis the
other embracing a lateral but often undeveloped sho?t;
pollination under or at the water surf~ce 17. Halop~lla
- Freshwater; horizontal axes absent or If present then With-
out 2, equally sized, scalelike leaves at each internode;
pollination at or above the water _surface . .
10. Leaves spirally arranged, not obviOusly distichous
- Leaves distichous (clearly in 2 rows even if spirally wound)
11. Petals large and showy, much wider than the sepals; styles
bifid 1. Ottelia
243
- Petals rudimentary or relatively insignificant, scarcely
wider than the sepals; styles simple 5. Blyxa
12. Leaves confined to basal rosettes; sterns dimorphic (with
horizontal runners and erect contracted sterns); female
flowers borne on long pedicels 14. Vallisneria
- Leaves cauline or sometimes bunched on elongated sterns;
sterns monomorphic; female flowers sessile, borne on long
hypanthia 12. Ne~hamandra
13. At least the middle and upper leaves arranged m whorls of
3 or more 14
- All foliage leaves arranged spirally or in pairs 18
14. Squarnulae intravaginales fringed with fingerlike, orange-
2
brown hairs; styles simple and very much shorter than the
5
sepals; petals in male flowers linear 9. Hydrilla
3
- Squarnulae intravaginales with entire margins; styl~s
branched or if simple then longer than the sepals; petals m
4
the male flowers rudimentary or not linear
15. Leaves crowned by 2 spine cells, each larger than the mar-
ginal spine cells; male flowers liberated from the mother
plant, each with 3, erect, papillose starninodes
11. Lagarosiphon
- Leaves not crowned by enlarged spine cells, all spine cells
about the same size; male flowers liberated or remaining
attached to the mother plant, without starninodes 16
16. Petals inconspicuous, rarely exceeding the sepals or
absent; flowers without nectaries 8. Elodea
- Petals showy, at least twice as large as the sepals; flowers
17
with nectaries
17. Flowers bisexual; stamens 3; prophylls free and lateral or
somewhat oblique; leaves without idioblasts; leaf margins
6
with fibre cells 6. Apalanthe
- Flowers unisexual (plants dioecious); stamens 9; prophylls
13
united at base, median; leaves with colourless idioblasts;
7. Egeria
leaf margins without fibre cells
18. Leaves threadlike (Australia) 13. Maidenia
- Leaves flattened, more or less ribbonlike 19
19. Leaves crowned by 2 spine cells, each larger than the
20
marginal spine cells
Leaves not crowned by enlarged spine cells, all spine cells
more or less the same size 21
7 20. Male flowers with 6 fertile stamens and lacking
starninodes; female flowers with a uniseriate perianth
10. App~rtiella
(3 sepals)
8
- Male flowers with 3 fertile stamens and 3 erect starnmodes;
female flowers with a biseriate perianth (3 sepals and 3
11. Lagarosiphon
petals)
21. Leaf bases not clasping the stern; female flowers borne on
9
long, flexible peduncles; hypanthia absent 14. Valli~neria
Leaf bases clasping the stern; female flowers sessile or
22
subsessile; hypanthia elongate and flexible
22. Midrib ofleaf prominent; male flowers attached to mother
plant; styles entire and more or less cylindrical
5. Blyxa
- Midrib of leaf not prominent; male flowers liberated from
mother plant, free-floating; styles flattened and irregularly
12. Nechamandra
lobed
Genera of Hydrocharitaceae
10
11 1. Ottelia Persoon Fig. 62A
12 Ottelia Persoon, Syn. Pl. 1: 400 (1805); Cook, Syrnoens &
Urrni-Konig, Aquat. Bot. 18: 263-274 (1983); Cook & Urrni-
Konig, Aquat. Bot. 20: 131-177 (1983), rev.
244 Hydrocharitaceae
Polygamous (monoecious, dioecious or trio-
ecious), freshwater annuals or perennials. Roots
unbranched. Stems contracted but sometimes
creeping. Leaves distichous or spirodistichous,
without stipules, usually radical; juvenile leaves
linear, submerged; adult leaves usually differenti-
ated into petiole and blade, with or without spines;
their blades submerged or floating, from linear to
orbicular or cordate. Flowers numerous or soli-
tary. Petals large and showy or reduced in cleisto-
gamous flowers. Stamens 3-15, in female flowers
all staminodial but in male and bisexual flowers
some usually staminodial. Carpels 3-20 or more;
fruit a fleshy capsule; seeds numerous released in
a pulpy mass. 2n = 22, 28, 38, 42, 44, 48, 50, 52, 55,
60, 64, 66, 68, 72, 110, 132. Twenty one spp., tropi-
cal and subtropical Africa, Asia, Australasia and S
America, 0. alismoides (L.) Persoon naturalised in
Europe and N America.
2. Stratiotes L.
Stratiotes L., Sp. Pl.: 535 (1753); Cook & Urmi-Ktinig, Aquat.
Bot. 16: 213-249 (1983), rev.
Dioecious, freshwater perennial, submerged in
winter, rising to the surface to flower. Roots
simple, rarely attached to the substrate. Stems
contracted, complexly branched; branches run-
nerlike; bearing terminal buds which may func-
tion as hibernacula. Leaves spirotristichous, in a
rosette, rigid, sessile, inear or narrowly triangular,
with spinous-serrate margins. Male inflorescence
3- to 6-flowered, female 1- or rarely 2-flowered.
Petals showy. Stamens in male flowers up to 41,
the inner 5-17 fertile; female flowers with 15-30
staminodes. Ovary of 3-6 carpels; fruit a berrylike
capsule. 2n = ±20 - ±64. Only one sp., S. aloides
L., Europe and Central Asia.
3. Hydrocharis L. Fig. 62B,C
Hydrocharis L., Sp. Pl.: 1036 (1753); Cook & Li.itind, Aquat.
Bot. 14: 177-204, (1982), rev.
Monoecious, freshwater perennial. Roots bran-
ched. Stems contracted and bearing leaves or
elongate and leafless. Leaves scalelike or differen-
tiated into stipules, petiole and blade; stipules
either 2 lateral or 1 median; blades floating or
rarely emergent. Flowers unisexual, the females
borne singly, the males in complex cymes. Petals
3, showy, ±, orbicular. Hypanthium absent. Sta-
mens in 4-6 whorls of 3. Ovary of 6 carpels; fruit a
berry or berrylike capsule. 2n = 14, 16, 20, 22, 28.
Three spp., temperate and subtropical Eurasia,
and tropical Africa, H. morsus-ranae L.
naturalised in N America.
4. Limnobium Rich. Fig. 62D,E
Limnobium Rich., Mem. Cl. Sci. Math. Phys. Inst. Imp. France
12 (2): 66, t. 8 (dated 1812, published 1814); Cook & Urmi-
Ktinig, Aquat. Bot. 17: 1-27 (1983), rev.
Monoecious, freshwater perennials. Roots
branched. Stems contracted and bearing leaves or
elongate and leafless. Leaves scalelike or differen-
tiated into stipule, petiole and blade; stipule
median; blade floating or sometimes emergent.
Flowers unisexual, pedicelled, the females borne
singly, the males in complex cymes. Petals 3,
scarcely exceeding the sepals, rudimentary or
absent in some female flowers. Hypanthium ab-
sent. Stamens in 1-6 whorls of 3. Ovary of 3-9
carpels; fruit a berrylike capsule. 2n = 26-30. Two
spp., temperate and tropical America.
5. Blyxa Thouars ex Rich Fig. 62F,G
Blyxa Thouars ex Rich., Mem. Cl. Sci. Math. Phys. Inst. Imp.
France 12 (2): 19-23, 77 (dated 1812, published 1814); Cook
& Li.itind, Aquat. Bot. 15: 1-52 (1983), rev.
Monoecious or dioecious, freshwater annuals or
perennials. Roots unbranched. Stems either elon-
gate or contracted. Leaves submerged, spirally
arranged, flat, ribbonlike, not differentiated into
stipule, petiole and blade, midrib usually pro-
minent, margin usually minutely toothed. Inflo-
rescence with 1-22 or more flowers. Petals
band-shaped, capillary or rudimentary. Stamens
3, 6 or 9, all fertile, in female flowers staminodes
minute. Ovary of 3 carpels, distally elongated into
a hypanthium in bisexual and female flowers; fruit
an elongated, irregularly dehiscent capsule; seeds
smooth or spiny. 2n = 16, 18, 32, 42, 48, 72. Nine
spp., tropical Old World, naturalised inN America
and Europe.
6. Apalanthe Planchon
Apalanthe Planchon, Ann. Mag. Nat. Hist. II, 1: 87 (1848);
Cook, Aquat. Bot. 21: 157-164 (1985), rev.
Monoecious, freshwater perennial or annual.
Roots unbranched. Stems elongate, branched
regularly but at distant intervals. Leaves totally
submerged, scalelike and opposite or foliate spiral
 Hydrocharitaceae
or in whorls of 3-7 or more, alternate near the
base of a stem or branch, sessile, linear to nar-
rowly triangular, regularly spaced along the stem
but contracted towards the apex, minutely
serrulate, apex acute, terminating in a single
spine; squamulae intravaginales paired (or more
at branches). Flowers solitary in leaf axils; spathe
of 2 united bracts. Flowers bisexual, solitary in
spathes, held above the water surface on rigid hy-
panthia; sepals 3, reflexed; petals 3, spreading,
showy, obovate to widely obovate; stamens 3; fila-
ments filiform; staminodes absent. Carpels 3; fruit
ellipsoidal, irregularly dehiscent; seeds up to 7,
fusiform, covered with unicellular hairs. Only one
sp., A. granatensis (Humb. and Bonpl.) Planchon,
tropical S America.
7. Egeria Planchon Fig. 60A-C
Egeria Planchon, Ann. Sci. Nat. (Paris), III, 11: 79 (1849); Cook
& Urmi-Konig, Aquat. Bot. 19: 73-96 (1984), rev.
Dioecious, freshwater perennial or E. najas Plan-
chon occasionally annual. Roots unbranched.
Stems elongate, irregularly branched. Leaves sub-
merged, scalelike and opposite or foliate in whorls
of 3 or more, sessile, linear to widely lanceolate;
squamulae intravaginales paired (or more at
branches). Inflorescences axillary. Male flowers 2-
4, held above the water surface on rigid pedicels;
sepals 3; petals 3, large and showy; stamens 9 or
rarely more; filaments much longer than the an-
thers, staminodes absent; pistillodium 3-lobed.
Female flowers solitary, like males but with sta-
mens replaced by 3 linear staminodes, with an
elongate hypanthium bearing the flower above the
water surface. Carpels 3; fruit ellipsoidal, irregu-
larly dehiscent; seeds ellipsoidal. 2n = 48. Two
spp., subtropical and temperate S America, E.
densa Planchon almost cosmopolitan m warm
regions by naturalisation.
8. Elodea Michx. Fig. 60D-F
Elodea Michx., Fl. Bor. Am. 1: 20 (1803); Cook & Urmi-Konig,
Aquat. Bot. 21: 111-156, (1985), rev.
Dioecious or gynodioecious, freshwater perennial
or occasionally annual. Roots unbranched. Stems
horizontal and stoloniferous below, erect and
spreading above; bulbil-like hibernacula develop-
ing terminally or axillary on erect stems or their
branches. Leaves submerged, scalelike and oppo-
site or foliate in whorls of 3-8, sessile, linear to
245
obovate; squamulae intravaginales paired (or
more at branches). Inflorescences axillary. Male
flowers solitary, floating on the water surface,
attached by a thread-like pedicel or abscising as
buds; sepals 3; petals 3, rudimentary or absent;
stamens 9 or sometimes less, staminodes absent.
Female flowers solitary, subsessile but with a
threadlike hypanthium carrying the flower to the
surface; perianth of 3 sepals and 3 elongate-ovate
petals spreading and floating, staminodes 3, clav-
ate. Carpels 3; fruit ellipsoidal, indehiscent; seeds
cylindric to fusiform. 2n = 12?, 16?, 24,42-56. Five
spp., temperate America, naturalised in many
parts of the Old World and Australasia (E.
canadensis Michx. and E. nuttallii (Planchon) St.
John have become weeds in Europe, E Asia and
Australasia, E. callitrichoides (Rich.) Caspary is
naturalised in Europe).
9. Hydrilla Rich. Fig. 60G-I
Hydrilla Rich., Mem. Cl. Sci. Math. Phys. Inst. Imp. Fr. 12 (2):
9, 61, 73, t. 2 A-K (dated 1812, published 1814); Cook &
Liiond, Aquat. Bot. 13: 485-504 (1982), rev.
Gynodioecious, in fresh or brackish water, peren-
nial or occasionally annual. Roots unbranched.
Stems horizontal and stoloniferous below, erect
and spreading above; bulbil-like hibernacula
developing underground terminally on stolons,
or terminally or axillary on erect stems or their
branches. Leaves submerged, scalelike and oppo-
site or foliate in whorls of 3-12, sessile, linear to
lanceolate or rarely widely ovate; squamulae
intravaginales paired (or more at branches). Inflo-
rescences axillary. Male flowers solitary, small,
abscising as buds and opening explosively on the
water surface; sepals 3, reflexed; petals 3, reflexed;
stamens 3, staminodes absent. Female flowers
solitary, subsessile but with a long threadlike
hypanthium carried to the surface; perianth of 3
sepals and 3 elongate-ovate petals, spreading and
floating; staminodes 3, minute. Carpels 3; fruit
cylindrical, indehiscent; seeds rarely more than 5,
fusiform and smooth. 2n = 16, 24, 32. Only one
sp., H. verticillata (L. fil.) Royle, Old World,
naturalised in America, it has become a very
serious pest in southern USA.
10. Appertiella C.D.K. Cook & L. Triest
Appertie/la C.D.K. Cook & L. Triest in Symoens, Hooper,
Compere, P. (eds), Studies on Aquatic Vascular Plants, Soc.
Roy. Bot. Belgique (Brussels): 75-79 (1982); Appert, Bot.
Helv. 106: 57-61 (distrib., bioi.).
246 Hydrocharitaceae
Dioecious, freshwater annual. Roots unbranched.
Stems elongate, sparsely and irregularly branched.
Leaves submerged, spirally arranged, contracted
towards the apex, sessile, narrowly triangular, up
to 10 em long and 2.5 mm wide. Inflorescences ax-
illary, the male pedunculate and globose contain-
ing 100 or more flowers, the female cylindrical
containing 1 sessile flower. Male flowers very
small, abscising as buds; sepals 3 reflexed at an the-
sis; stamens 6 all fertile and held parallel to the
water surface. Female flowers sessile but with a
long threadlike hypanthium carried to the surface;
sepals 3, spreading and floating; petals and
staminodes absent. Carpels 3; fruit a cylindrical
capsule; seeds ± 5, cylindrical. Only one sp., A.
hexandra C.D.K. Cook & L. Triest, Madagascar.
11. Lagarosiphon Harvey Fig. 60J-L
Lagarosiphon Harvey, Hook. J. Bot. 4: 230 (1841); Symoens &
Triest, Bull. Jard. Bot. Nat!. Belg. 53: 441-488 (1983), rev.
Dioecious or rarely monoecious, freshwater pe-
rennials. Roots unbranched. Stems elongate,
branched regularly, horizontal below and erect
above. Leaves submerged, spiral, subopposite or
in whorls of 3 or more, scalelike and more or less
triangular or foliate and linear{-lanceolate). Inflo-
rescences axillary, the male pedunculate and
globose containing numerous flowers, the female
cylindrical containing 1 sessile flower. Male flow-
ers small, abscising as buds; perianth of 3 sepals
and 3 petals, reflexed at anthesis; stamens 6, 3
fertile and held parallel to the water surface and 3
papillose staminodes held vertically and function-
ing as sails. Female flowers sessile but with long
threadlike hypanthia carrying the flowers to the
surface; perianth of3 sepals and 3 petals spreading
and floating; staminodes 3, linear. Plants with
bisexual flowers have recently been collected in
Madagascar. Carpels 3; fruit a cylindrical capsule;
seeds 5-30, ellipsoidal. 2n = 22. Nine spp., Africa,
Madagascar; L. major (Ridley) Moss is naturalised
in Europe and New Zealand, it has become a very
serious pest in New Zealand.
12 Nechamandra Planchon Fig. 60M-P
Nechamandra Planchon, Ann. Sci. Nat. (Paris), III, 11: 78
(1849); Cook & Liiond, Aquat. Bot. 13: 505-513 (1982), rev.
Dioecious, freshwater annual or perhaps also pe-
rennial. Roots unbranched. Stems elongate, inter-
nodes long or contracted at apparently irregular
intervals. Leaves distichous, flat, band-shaped,
1.5-6 em long. Inflorescences axillary, often in
clusters. Male flowers small, numerous, abscising
as buds; sepals 3, reflexed at anthesis; petal1 rudi-
mentary or absent; staminode 1 or absent; sta-
mens 2, borne parallel to the water surface. Female
flowers solitary, sessile, with long threadlike hy-
panthia, floating on the surface at maturity; sepals
3; petals rudimentary or absent; staminodes
rudimentary or absent. Ovary of 3 carpels; fruit
breaking irregularly; seeds numerous. 2n = 16, 99,
100, 101. Only one sp., N. alternifolia (Roxb. ex
Wight) Thwaites, India and SE Asia, naturalised
in Sudan.
13. Maidenia Rendle Fig. 61A-C
Maidenia Rendle, J. Bot. (Lond.) 54: 313 (1916).
Monoecious or perhaps also dioecious, freshwater
annual or perhaps also perennial. Roots un-
branched. Stems elongated above, sometimes de-
veloping stolons or runners below. Leaves totally
submerged, spirodistichous, threadlike, 2-3 em
long. Inflorescences axillary, often in clusters
along the stem. Male flowers small, numerous,
abscising as buds; sepals unequal, becoming re-
flexed at anthesis; petals absent; staminode 1 or
absent; stamen 1; anther 3-locular (probably de-
rived from 3 united stamens), each locule with up
to 8 pollen grains. Female flowers solitary, long
pedicelled, floating on the surface at maturity;
sepals 3; petals absent; staminodes 3. Ovary of 3
carpels; fruit breaking irregularly; seeds numer-
ous, ellipsoidal, up to 2 mm long. Only one sp., M.
rubra Rendle, NW Australia.
14. V allisneria L. Fig. 61D-G
Vallisneria L., Sp. Pl.: 1015 (1753); Lowden, Aquat. Bot. 13:
269-289 (1992), rev.; Jacobs & Frank, Telopea 7: 111-118
(1997), rev. Austral. spp.
Dioecious, freshwater annuals or perennials.
Roots unbranched. Stems mostly developing sto-
lons or runners. Foliage leaves submerged, disti-
chous or spirodistichous, narrowly strap-shaped,
up to 2m or more long but usually much less.
Inflorescences axillary. Male flowers small, nu-
merous, abscising as buds and opening on water
surface; sepals unequal, reflexed at anthesis; petal
rudimentary or absent; staminode 1 or absent;
stamens 2 or 1. Female flowers solitary, long-
pedicelled, floating on surface at maturity; sepals 3
or 2; petals 3 or 2, minute; staminodes 3 or 2,
alternating with the petals. Ovary of 3 or 2 carpels;
 Hydrocharitaceae
fruit breaking irregularly; seeds numerous, ellip-
soidal, up to 2 mm long. 2n = 20, 30, 40. Six spp.,
tropical to warm regions of the world.
15. Enhalus Rich. Fig. 61H
Enhalus Rich., Mem. Cl. Sci. Math. Phys. Inst. Imp. France 12
(2): 64, 71, 74, (dated 1812, published 1814); Hartog, Verh.
K. Nederl. Akad. Wet. Afd. Natuurkd. 59: 1-275 (1970), rev.
Dioecious, marine perennial. Roots unbranched.
Stem a simple or monopodially branched rhizome
clothed in fibrous remains of leaf sheaths. Leaves
submerged, distichous, usually 2-6 at the apex of
the rhizome, strap-shaped, up to 1m or more long,
up to 1.5 em wide, the veins in 2 series. Male
flowers small, numerous, abscising as buds; peri-
anth 2-seriate; reflexed at anthesis; stamens 3. Fe-
male flowers solitary, long pedicelled, floating on
the surface at maturity; sepals recurved, petals
spreading. Carpels 6; fruit a large and fleshy cap-
sule, 7 em long, 4 em diam.; seeds few, angular.
Only one sp., E. acoroides (L. fil.) Rich. ex Steudel,
coasts of the Indian and W Pacific Oceans.
16. Thalassia Banks ex Konig Fig. 611-L
Thalassia Banks ex Konig, Ann. Bot. 2: 96 (1805); Hartog,
Verh. K. Nederl. Akad. Wet. Afd. Natuurkd. 59: 1-275
(1970), rev.
Dioecious, marine perennials. Roots unbranched.
Horizontal scale-bearing rhizomatous axes (long
shoots) buried in the substrate giving rise to erect
but contracted leaf bearing axes (short shoots).
Leaves submerged, 2-6, distichous, strap-shaped
or somewhat falcate, up to 70 em long and 1.5 em
wide. Inflorescence submerged of 1-4 flowers. Pe-
rianth 1-seriate of 3 segments. Stamens 3-13, all
fertile. Ovary elongated into a hypanthium; fruit a
globose and fleshy capsule. Two spp., coasts of
Caribbean and Indo-Pacific Ocean.
17. Halophila Thouars Fig. 61M,N
Halophila Thouars, Gen. Nov. Madag. 2: 2 (1806); Hartog,
Verh. K. Nederl. Akad. Wet. Afd. Natuurkd. 59: 1-275
(1970), rev.
Monoecious or dioecious, diminutive, marine pe-
rennial or annual. Roots unbranched. Stems regu-
larly branched, creeping and stoloniferous with
subopposite scales or erect and bearing foliage
leaves. Foliage leaves submerged, distichous,
sessile or petiolate, linear to ovate, rarely exceed-
247
ing 3 em long. Flowers solitary. Male flowers pedi-
cellate with a uniseriate perianth of 3 small seg-
ments and 3 stamens; female flowers sessile with a
vestigial perianth. Fruit a fleshy capsule with per-
sistent styles; seeds several, globose. About ten
rather poorly defined spp., tropical to warm seas
of the world.
Selected Bibliography
Ancibor, E. 1979. Systematic anatomy of vegetative organs of
the Hydrocharitaceae. Bot. J. Linn. Soc. 78: 237-266 (plus
microfiche).
Appert, 0. 1996. Zur Verbreitung and Biologie der Gattung
Appertiella (Hydrocharitaceae). Bot. Helvetica. 106: 57-
71.
Brunaud, A. 1976. Ramification chez les Hydrocharitaceae. I.
Ontogenie du systeme des pousses. Rev. Gen. Bot. 83: 397-
413.
Brunaud, A. 1977. Ramification chez les Hydrocharitaceae. II.
Organisation des rameaux lateraux. Rev. Gen. Bot. 84: 137-
157.
Cook, C.D.K. 1982. Pollination mechanisms in the Hydro-
charitaceae. In: Symoens, J.-J., Hooper, S.S., Compere P.
(eds.) Studies on aquatic vascular plants. Brussels, Royal
Botanical Society of Belgium, pp. 1-15.
Cook, C.D.K. 1985. A revision of the genus Apalanthe
(Hydrocharitaceae). Aquat. Bot. 21: 157-164.
Cook, C.D.K., Liiiind, R. 1982a. A revision of the genus
Hydrilla (Hydrocharitaceae). Aquat. Bot. 13: 485-504.
Cook, C.D.K., Liiiind, R. 1982b. A revision of the genus
Nechamandra (Hydrocharitaceae). Aquat. Bot. 13: 505-513.
Cook, C.D.K., Liiiind, R. 1982c. A revision of the genus
Hydrocharis (Hydrocharitaceae). Aquat. Bot. 14: 177-204.
Cook, C.D.K., Liiiind, R. 1983. A revision of the genus Blyxa
(Hydrocharitaceae). Aquat. Bot. 15: 1-52.
Cook, C.D.K., Urmi-Kiinig, K. 1983a. A revision of the genus
Stratiotes (Hydrocharitaceae). Aquat. Bot. 16: 213-249.
Cook, C.D.K., Urmi-Kiinig, K. 1983b. A revision of the genus
Ottelia (Hydrocharitaceae). 2. The species of Eurasia,
Australia and America. Aquat. Bot. 20: 131-177.
Cook, C.D.K., Urmi-Kiinig, K. 1983c. A revision of the genus
Limnobium including Hydromystria (Hydrocharitaceae).
Aquat. Bot. 17: 1-27.
Cook, C.D.K., Urmi-Kiinig, K. 1984. A revision of the genus
Egeria (Hydrocharitaceae). Aquat. Bot. 19: 73-96.
Cook, C.D.K., Urmi-Kiinig, K. 1985. A revision of the genus
Elodea (Hydrocharitaceae). Aquat. Bot. 21: 111-156.
Cook, C.D.K., Liiiind, R., Nair, B. 1981. Floral biology of Blyxa
octandra (Roxb.) Planchon ex Thwaites (Hydrocharita-
ceae). Aquat. Bot. 10: 61-68.
Cook, C.D.K., Symoens, J.-J. Urmi-Kiinig, K. 1983. A revision
of the genus Ottelia (Hydrocharitaceae). 1. Generic con-
siderations. Aquat. Bot. 18: 263-274.
Cox, P.A., Knox, B.P. 1989. Two-dimensional pollination in
hydrophilous plants: convergent evolution in the genera
Halodule (Cymodoceaceae), Halophila (Hydrocharitaceae),
Ruppia (Ruppiaceae), and Lepilaena (Zannichelliaceae).
Am. J. Bot. 76: 164-175.
Cox, P.A., Tomlinson, P.B. 1988. Pollination ecology of a
seagrass, Thalassia testudinum (Hydrocharitaceae), in St.
Croix. Am. J. Bot. 75: 958-965.
Dahlgren, R. et a!. 1985. See general references.
248 Hydrocharitaceae

Davis, G.L. 1966. See general references. Vallisneria americana (Hydrocharitaceae). Am. J. Bot. 61:
Hegnauer, R. 1963, 1986. See general references. 846-866.
Kaul, R.B. 1968. Floral morphology and phylogeny in the Wilder, G.J. 1975. Phylogenetic trends in the Alismatidae
Hydrocharitaceae. Phytomorphology 18: 13-35. (Monocotyledoneae). Bot. Gaz. 136: 159-170.
Kaul, R.B. 1970. Evolution and adaptation of inflorescences in Yeo, R.R., Falk, R.H., Thurston, J.R. 1984. The morphology of
the Hydrocharitaceae. Am. J. Bot. 57: 708-715. hydrilla (Hydrilla verticillata (L.f.) Royle). J. Aquat. Plant
Langland, K.A. 1990. Hydrilla: a continuing problem in Manage. 22: 1-17.
Florida waters. Circular No. 884, Institute of food and
agricultural sciences. Gainesville: University of Florida,
pp. 1-21.
Les, D.H. et a!. 1997. See general references.
McConchie, C.A. 1983. Floral development of Maidenia rubra
Rendle (Hydrocharitaceae). Aust. J. Bot. 31: 585-603.
McConchie, C.A., Kadereit, J.W. 1987. Floral structure of
Vallisneria caulesens Bailey & F. Mueller. Aquat. Bot. 29:
101-110.
Montesantos, M. 1913. Morphologische und biologische
Untersuchungen tiber einige Hydrocharitaceen. Flora
(N.F.) 5: 1-32.
Mues, R. 1983. Species specific flavone glucuronides in Elodea
species. Biochem. Syst. Ecol. 11: 261-265.
Miinchberg, P. 1956. Zur Bindung der Libelle Aeschna viridis
Eversm. an die Pflanze Stratiotes aloides L. Nachrichtenbl.
Bayer. Entomol. 5(12): 113-118.
Nakamura, T., Kadono, Y. 1993. Chromosome number and
geographical distribution of monoecious and dioecious
Hydrilla verticillata (L.f.) Royle (Hydrocharitaceae) in
Japan. Acta Phytotax. Geobot. 44: 123-140.
Scribailo, R.W., Posluszny, U. 1983. The reproductive biology
of Hydrocharis morus-ranae. 1. Floral biology. Can. J. Bot.
62: 2779-2787.
Shaffer-Fehre, M. 1991. The position of Najas within the sub-
class Alismatidae (Monocotyledons) in the light of new evi-
dence from seed coat structures in the Hydrocharitoideae
(Hydrocharitales). Bot. J. Linn. Soc. 107: 189-209.
Singh, V. 1966a. Morphological and anatomical studies in
Helobiae. VIII. Vascular anatomy of the flower of
Hydrocharitaceae-Stratioideae and Thalassioideae. Agra
Univ. J. Res. Sci. 15: 43-59.
Singh, V. 1966b. Morphological and anatomical studies in
Helobiae. IX. Vascular anatomy of the flower of
Hydrocharitaceae-Vallisnerioideae and Halophiloideae.
Agra Univ. J. Res. Sci. 15: 84-106.
Svedelius, N. 1932. On the different types of pollination in
Vallisneria spiralis L. and Vallisneria americana Michx.
Sven. Bot. Tidskr. 26: 1-12.
Symoens, J.J., Triest, L. 1983. Monograph of the African genus
Lagarosiphon Harvey (Hydrocharitaceae). Bull. Jard. Bot.
Nat!. Belg. 53: 441-488.
Takahashi, M. 1994. Pollen development in a submerged plant
Ottelia alismoides (L.) Pers. (Hydrocharitaceae). J. Plant
Res. 107: 161-164.
Tillich, H.-J. 1995. Seedlings and systematics in monocotyle-
dons. In: Rudall, P.J., Cribb, P.J., Cutler, D.F., Humphries,
C.J. (eds.) Monocotyledons: systematics and evolution.
London: Royal Botanic Gardens, Kew, pp. 303-352.
Tomlinson, P.B. 1969. On the morphology and anatomy of
turtle grass, Thalassia testudinum (Hydrocharitaceae). III.
Floral morphology and anatomy. Bull. Mar. Sci. (Miami) 19:
286-305.
Tomlinson, P.B. 1982. Helobiae (Alismatidae). In: Metcalfe
C.R. (ed.) Anatomy of monocotyledons, vol. VII. Oxford:
Clarendon Press, pp. 126-197.
Wilder, G.J. 1974a. Symmetry and development of Limnobium
spongia (Hydrocharitaceae). Am. J. Bot. 61: 624-642.
Wilder, G.J. 1974b. Symmetry and development of pistillate
 Join villeaceae 249

Joinvilleaceae chloroplast-containing cells. Transverse veins are

conspicuous but infrequent. Air canals are present
C. BAYER and 0. APPEL in the sheath (Tomlinson 1969).
According to Cheadle and Kosakai (1980), ves-
sels occur in the roots, rhizomes, stems and leaves.
Simple perforations occur predominantly in
roots and rhizomes, while scalariform perforation
plates prevail in stems and leaves. Lateral pits are
numerous and small.
Joinvilleaceae Tom!. & A. C. Sm., Taxon 19: 888 (1970).
REPRODUCTIVE STRUCTURES. The inflorescence
Perennial herbs from condensed, sympodial rhi- of]oinvillea elegans resembles a bracteate panicle,
zomes; aerial shoots erect, unbranched, hollow but is composed of spikes or racemes instead
except for the nodes, terete or rarely compressed. of single flowers. This structure is similar to that
Leaves distichous; sheath open, more or less ligu- of Flagellaria; both share the conspicuous adaxial
late; blade linear-lanceolate with constricted base, swellings on the base of each lateral axis of the
pubescent, plicate, the veins anastomosing. Inflo-
rescence terminal, many-flowered, paniculiform,
bracteate. Flowers (sub-)sessile, hermaphroditic,
actinomorphic, pentacyclic, trimerous. Tepals
free or almost so, dry, bractlike with thickened
bases, concave, persistent. Stamens free or slightly
adnate to the perianth base; anthers basifixed, sag-
ittate, dithecal, introrse, opening by longitudinal
slits. Ovary superior, globose, trilocular, each loc-
ule containing a single, atropous, apical-axile,
pendulous ovule; styluli 3, free or slightly connate
at the base; stigmas feathery, papillate. Fruit
drupaceous, 3-1-seeded, whitish or reddish to
black at maturity. Seeds with copious, starchy
endosperm and minute discoid embryo.
A monogeneric family with 2 spp. distributed
from the Malay Peninsula to Sumatra, Borneo and
many Pacific Islands.

VEGETATIVE STRUCTURES. The aerial shoots may

reach more than 5 min length (Fig. 63). The occur-
rence of short, unicellular prickle hairs and spiny,
multicellular, occasionally branched hairs is
characteristic at the specific and subspecific level
(Newell1969).
The leaf epidermis is composed of rows of long
cells alternating with short cells that contain silica
bodies. This pattern is interrupted by longitudinal
bands of expansion cells which are found in the
plicae and adjacent to the midrib. Stomata occur
on both sides but are less common on the abaxial
face. They have characteristic dumbbell-shaped
guard cells and are of the grass type. The meso-
phyll contains calcium oxalate crystals, tannins
and silica. It includes a 2- to 3-layered chloren-
chyma of more or less isodiametric cells on both Fig. 63A-D. Joinvilleaceae. A-C ]oinvillea plicata. A Leaf and
fruiting branch. B Fruit. C Same, longitudinal section, left seed
sides of the leaf and a central tissue of large showing embryo. D ]. ascendens, upper part of leaf sheath
colourless cells. Each bundle is enclosed by a showing hyaline stipule-like appendages. (Dahlgren et a!.
fibrous sheath and laterally flanked by a layer of 1985)
250 Joinvilleaceae
inflorescence, which may serve to regulate the
position of the branches. Similar structures occur
in grasses.
At least the inner tepals usually have three veins
(Newell 1969). The thickened tepal bases some-
what remind of the grass Iodides and might re-
semble hypothetical primitive forms of the latter
(Endress 1995). The styluli are linear; they spread
at anthesis and may persist in fruit. The stigmas
are plumose with dispersed, interrupted recep-
tive sites on multiseriate branches, a feature that is
otherwise restricted to Poaceae and some
Restionaceae (Endress 1995).
Septal nectaries are lacking, the flowers are
probably anemophilous. The seeds are dispersed
mainly by birds (Degener and Degener 1973).
EMBRYOLOGY. The tapetum is of the secretory
type (Kircher 1986). Manning and Linder (1990)
and Campbell and Kellogg (1986) reported unique
endothecial thickenings (girdle type), similar to
those of Ecdeiocolea. Microsporogenesis is suc-
cessive. Pollen is shed at the 3-celled stage. The
micropyle is formed by both integuments each
consisting of 2 cell layers. The megagametophyte
is free of starch. Three antipodals with each 2
nuclei are present. Endosperm formation is
Nuclear and starch grains are compound in ma-
ture endosperm (Campbell and Kellogg 1986;
Kellogg and Linder 1995; Linder and Kellogg
1995).
POLLEN MORPHOLOGY. Pollen morphology was
studied by Erdtman (1944, 1952), Chanda (1965,
1966), Chanda and Rowley (1967) and Linder and
Ferguson (1985). The grains are spheroidal, ulcer-
ate and microperforate monads. They closely
resemble those of other Poales, especially
Flagellaria, differing from the latter mainly in
their larger size (about 35 f!m in diameter).
"Graminoid" apertures, covered by a thin endex-
ine layer that extends to some distance under the
margin, are common to Joinvilleaceae, Flagel-
lariaceae, some Restionaceae, Poaceae, Ecdeiocol-
eaceae and Anarthriaceae. In contrast to the latter
three families, an operculum seems to be lack-
ing in the pollen of Joinvilleaceae (Linder and
Ferguson 1985).
KARYOLOGY. Newell (1969) reported several
counts of n = 18 and one of 2n = 36 for ]oinvillea
ascendens and f. elegans (as]. plicata).
PHYTOCHEMISTRY. Harris and Hartley (1980)
observed a characteristic UV-fluorescence of
cell walls, indicating the presence of bound p-
coumaric, ferulic and diferulic acids.
AFFINITIES. Together with Hanguana, ]oinvillea
was originally included in the Flagellariaceae. The
exclusion of Hanguana (see there) has been gener-
ally accepted, but some authors preferred to retain
the more similar genera ]oinvillea and Flagella ria
in one family (Baranova 1975). Tomlinson and
Smith (1970) proposed to treat ]oinvillea as a
separate family that differs from Flagellaria in
characters such as growth form, pubescence, pli-
cate, dorsiventral leaves with open sheaths and
without tendrils, and some anatomical features.
The Joinvilleaceae are generally included in the
Poales. ]oinvillea, Poaceae, some Restionaceae,
and Flagellaria share atropous ovules, which may
be modified in the Poaceae and Restionaceae.
Joinvilleaceae, Poaceae and Restionaceae are
united by their unique complex stigmas (Endress
1995), and pollen morphology supports this alli-
ance. Further evidence was provided by phyloge-
netic analyses of morphological characters and
chloroplast DNA restriction sites and sequences.
According to these studies, ]oinvillea is most
likely to represent the sister group of Poaceae
(Kellogg and Birchler 1993; Chase et al. 1995,
1995a; Davis 1995; Hahn et al. 1995; Kellogg and
Linder 1995; Linder and Kellogg 1995; Stevenson
and Loconte 1995). Both families share alternating
short and long cells in the leaf epidermis and the
6-kb and 28-kb inversion in the chloroplast ge-
nome (Doyle et al. 1992; Linder and Rudall 1993;
Linder and Kellogg 1995).
DISTRIBUTION AND HABITATS. Since there are no
records from eastern Indonesia, New Guinea and
Australia, Newell (1969) regarded the distribution
of ]oinvillea on the Malay Penisula, Sumatra,
Borneo, and on many Pacific Islands as discon-
tinuous. The species occur on non-swampy soils
in moderately humid to dry light forests, forest
margins, thickets and mountain ridges up to more
than 1900m above sea level (Backer 1951; Newell
1969). According to Linder (1987), they probably
germinate exclusively on mineral soils and repre-
sent early successional plants.
Only one genus:
]oinvillea Gaudich. Fig. 63
]oinvillea Gaudich., Voy. La Bonite, Bot, Atlas: pl. 39-40, figs.
7-26 {1841); Newell, J. Arnold Arbor. 50: 527-555 {1969)
rev.; St. John, Phytologia 40: 369-374 {1978), nomenclat.
 Joinvilleaceae
Characters as for family.
Two spp., ]. ascendens Gaudich. ex Brongn. et
Gris and ]. elegans Gaudich.
Selected Bibliography
Backer, C.A. 1951. Flagellariaceae. In: Van Steenis, C.G.G.J.
(ed.) Flora Malesiana I, 4. Djakarta: NoordhoffKolffN. V.,
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Baranova, M.A. 1975. Stomatographic study of the Flagel-
lariaceae. Bot. Zh. (Moscow & Leningrad) 60: 1690-1697.
Brongniart, M.M., Gris, A. 1861. Note sur le genre]oinvillea de
Gaudichaud et sur Ia famille des flagellariees. Bull. Soc. Bot.
Fr. 8: 264-269.
Campbell, C.S., Kellogg, E.A. 1986. Sister group relationships
of the Poaceae. In: Sonderstrom, T.R., Hilu, K.W.,
Campbell, C.S., Barkworth, M.E. (eds.) Grass system~tics
and evolution. International grass symposmm.
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Chanda, S. 1965. On the pollen morphology of the Flagel-
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Calcutta 8: 53-SS.
Chanda, S. 1966. On the pollen morphology of the Cen-
trolepidaceae, Restionaceae and Flagellariaceae, with
special reference to taxonomy. Grana 6: 355-415.
Chanda, S., Rowley, S. 1967. Apertural types in pollen of the
Restionaceae and Flagellariaceae. Grana 7: 16-36.
Chase, M.W. eta!. 1995, 1995a. See general references.
Cheadle, V.I., Kosakai, H. 1980. Occurrence and specialization
of vessels in Commelinales. Phytomorphology 30: 98-117.
Christophersen, E.1931. Notes on]oinvillea. Bernice P. Bishop
Mus. Bull. 9: 2-7.
Dahlgren, R.M.T. eta!. 1985. See general references.
Davis, J.I. 1995. A phylogenetic structure for the monocotyle-
dons, as inferred from chloroplast DNA restriction site
variation, and a comparison of measures of clade support.
Syst. Bot. 20: 503-527.
Degener, 0., Degener, I. 1973. Joinvilleaceae. In: Degener, 0.,
Degener, I. (eds.) Flora Hawaiiensis 7.
Doyle, J.J., Davis, J.I., Soreng, R.J., Garvin, D., Anderson, M.J.
1992. Chloroplast DNA inversions and the origin of the
grass family (Poaceae). Proc. Nat!. Acad. Sci. U.S.A. 89:
7722-7726.
Endress, P.K. 1995. Major evolutionary traits of monocot flow-
ers. In: Rudall, P.J., Cribb, P.J., Cutler, D.F., Humphries, C.J.
(eds.) Monocotyledons: systematics and evolution. Royal
Botanic Gardens, Kew, pp. 43-79.
Erdtman, G. 1944. Pollen morphology and plant taxonomy. II.
Sven. Bot. Tidskr. 38: 163-168.
Erdtman, G. 1952. See general references.
Hahn, W.J., Givnish, T.J., Sytsma, K.J. 1995. Evolution of
the monocot chloroplast inverted repeat: I. Evolution and
phylogenetic implications of the ORF 2280 deletion. In:
Rudall, P.J., Cribb, P.J., Cutler, D.F., Humphries, C.J. (eds.)
Monocotyledons: systematics and evolution. Royal Botanic
Gardens, Kew, pp. 579-587.
Harris, P.J., Hartley, R.D. 1980. See general references.
Kellogg, E.A., Birchler, J.A. 1993. Linking phrlogen~ and ge-
netics: Zea mays as a tool for phylogenetic studies. Syst.
Bioi. 4: 415-439.
Kellogg, E.A., Linder, H.P. 1995. Phylogeny ~f Poales. In:
Rudall, P.J., Cribb, P.J., Cutler, D.F., Humphnes, C.J. (eds.)
Monocotyledons: systematics and evolution. Royal Botanic
Gardens, Kew, pp. 511-542.
251
Kircher, P. 1986. Untersuchungen zur Bliiten- und Inflores-
zenzmorphologie, Embryologie und Systematik der
Restionaceen im Vergleich mit Gramineen und verwandten
Familien. Diss. Bot. 94.
Lee, D.W., Yap Kim Pin, Lew Foo Yew 1975. Serological evi-
dence on the distinctness of the monocotyledonous families
Flagellariaceae, Hanguanaceae and Joinvilleaceae. Bot. J.
Linn. Soc. 70: 77-81.
Linder, H.P. 1987. The evolutionary history of Poales/
Restionales - a hypothesis. Kew Bull. 4: 297-318.
Linder, H.P., Ferguson, I.K. 1985. On the pollen morphology
and phylogeny of the Restionales and Po ales. Grana 24: 65-
76.
Linder, H.P., Kellogg, E.A. 1995. Phylogenetic patterns in tiie
Commelinid clade. In: Rudall, P.J., Cribb, P.J., Cutler, D.F.,
Humphries, C.J. (eds.) Monocotyledons: systematics and
evolution. Royal Botanic Gardens, Kew, pp. 473-496.
Linder, H.P., Rudall, P.J. 1993. The megagametophyte in
Anarthria (Anartiiriaceae, Poales) and its implications for
the phylogeny of the Poales. Am. J. Bot. 80: 1455-1464.
Manning, J.C., Linder, H.P. 1990. Cladistic analysis of patterns
of endothecial thickenings in the Poales/Restionales. Am. J.
Bot. 77: 196-210.
Newell, T.K. 1969. A study of the genus ]oinvillea (Flagel-
lariaceae). J. Arnold Arbor. 50: 527-555.
Newell, T.K., Stone, B.C. 1967. Flagellaria (Chortodes) plicata
Hooker fil. is a ]oinvillea. Taxon 16: 192-194.
Smith, A.C. 1979. Joinvilleaceae. In: Smith, A.C. (ed.) Flora
Vitensis Nova. A new flora of Fiji 1. Honolulu: SB Printers,
pp. 289-290.
Smithson, E. 1956. The comparative anatomy of tiie Flagel-
lariaceae. Kew Bull. 11: 491-501.
Stevenson, D.W., Loconte, H. 1995. Cladistic analysis of
monocot families. In: Rudall, P.J., Cribb, P.J., Cutler, D.F.,
Humphries, C.J. (eds.) Monocotyledons: systematics and
evolution. Royal Botanic Gardens, Kew, pp. 543-578.
Stone, B.C. 1981. Nomenclature of ]oinvillea (Joinvilleaceae).
Gard. Bull. Straits Settlem. III, 34: 223-225.
Tomlinson, P.B. 1969. Commelinales-Zingiberales. In:
Metcalfe, C.R. (ed.) Anatomy of the monocotyledons, Vol.
3. Oxford: Clarendon Press.
Tomlinson, P.B., Smith, A. C. 1970. Joinvilleaceae, a new family
of monocotyledons. Taxon 19: 887-889.
252 Juncaceae
Juncaceae
H. BALSLEV
A.L. Jussieu, Gen. Pl.: 43 {1789), "Junci".
Perennial and usually rhizomatous herbs, more
rarely annuals (some spp. of ]uncus and Luzula),
and very rarely with a stout woody trunk
(Prionium), glabrous or sometimes with hairy leaf
margins (Luzula). Rhizome creeping, ascending
or erect, leafless or with scales. Flowering stems
erect, ascending or rarely procumbent, terete or
laterally compressed, smooth or longitudinally
ridged. Leaves spiral, tristichous or rarely disti-
chous (Distichia, some spp. of ]uncus), basal or
on the lower part of the stem, linear or filiform,
with an open and sometimes auriculate or
closed sheath, the blade dorsiventral and flat or
canaliculate or terete to ± compressed, when ter-
ete hollow, and then sometimes with partitions;
blade sometimes reduced. Inflorescences terminal
or pseudolateral, compound, of open cymose
panicles, or often congested and head- or spike-
like, or flowers sometimes solitary and terminal
(Rostkovia, Marsippospermum) or lateral (Dis-
tichia, Oxychloe and Patosia). Flowers perfect
or unisexual-dioecious or rarely monoecious,
actinomorphic, hypogynous; tepals usually incon-
spicuous, rarely up to 40 mm long (Marsippo-
spermum), 6 (rarely 4) in 2 whorls of 3 (rarely 2),
glumaceous or herbaceous, equal to subequal,
free. Stamens 6 (rarely 4) in 2 whorls of 3 (rarely 2)
alternating with the tepals, or inner whorl some-
times lacking; filaments filiform or flat and wid-
ened at the base; anthers 2-thecate, 4-sporangiate,
oblong to linear, basifixed, obtuse to mucronate,
dehiscing by longitudinal lateral slits; pollen
grains ulcerate, in tetrads. Gynoecium of 3 (rarely
2) united carpels; ovary superior, 3-locular (rarely
2-locular) or sometimes 1-locular and then in-
completely septate; style 3-branched, rarely 2-
branched, or rarely with 3 (2) separate styluli;
stylar branches (stylodia) filiform, sometimes ta-
pering, with adaxial stigmatic papillae, sometimes
twisted. Ovules 3 to many, central, basal or ± pa-
rietal, anatropous. Fruit an orbicular to oblong or
ellipsoid, 3-lobed, round or trigonous, loculicidal
capsule. Seeds 3 (Luzula) to many, with copious
starchy endosperm and a small, broadly cylindric
embryo; outer seed coat hyaline, whitish or light
brown, sometimes with tail-like appendages, often
reticulate and frequently shed before ripening; in-
ner seed coat brown to castaneous or yellow.
A family of worldwide distribution with 8 genera
and ca. 350 species.
VEGETATIVE MORPHOLOGY. All members of the
Juncaceae are herbs, except Prionium which isba-
sally woody and has a trunk up to 1m in length
(Fig. 64). Distichia, Oxychloe, Patosia and some
species of Luzula form flat or bulging cushions up
to several metres wide. The remaining genera are
solitary or caespitose herbs or form mats by
means of densely branching rhizomes. Some are
aquatics.
The roots are adventitious along the rhizome or,
in the cushion-forming genera, emerge among the
leaves and develop in the interior of the cushion
often not reaching the substrate.
The stem is modified in various ways. In the
cushion-forming genera it bifurcates regularly
and dies back at the base; the internodes are then
reduced to 2-3 mm and the stem is completely
covered by the leaf sheaths. In Rostkovia and
Marsippospermum and the perennial species of
]uncus and Luzula the stem is differentiated into a
subterranean long-lived rhizome that dies back at
the base, lateral shoots from the rhizomes, leafy
short shoots and an above-ground ephemeral
flowering stem. Some species of ]uncus branch by
intravaginal shoot formation. In Prionium the rhi-
zome continues into a conspicuous trunk that
may reach 1m or more in height and 5-10 em in
diameter (Fig. 64A).
Foliage leaves are differentiated into a sheath
and blade. The sheath is open or closed (Luzula,
Prionium, ]uncus lomatophyllus, f. engleri), and
often terminates in two membranous or scarious
auricles, which sometimes extend onto the adaxial
side of the sheath to form a ligule. The shape and
texture of the auricles is taxonomically important
in ]uncus. The blade is lanceolate, linear or fili-
form, rarely short and ± conical, as in Distichia.
The margins are hairy in Luzula and serrate in
Prionium and ]uncus trifidus. The leaves of the
rhizome are modified to small membranous flaps
or hard sclerified, scales. Sheathing cataphylls are
often present at the base of the flowering stem.
VEGETATIVE ANATOMY. (Review: Cutler 1969).
Juncaceae lack crystal oxalate raphides. Silica has
been reported only occasionally and is at least
normally absent. The sieve tube plastids contain
cuneate protein crystals but no other inclusions
 Juncaceae
Fig. 64A-M. Juncaceae. A-F Prionium serratum. A Plant with
stout woody trunk and much branched panicle. B Inflores-
cence branch. C Flower. D Ovary with three stylodia. E Many-
seeded 3-locular ovary in cross-section. F Loculicidal capsule.
G-:I Marsippospermum grandiflorum. G Plant with creeping
rhizome and 1-flowered stems. H Pistil. I Loculicidal capsule.
J-M Distichia muscoides. J Plant with forking branches, disti-
chous leaves and subapical lateral staminate flowers. K
Staminate flower. L Fruit originating between six tepals and
supported by two floral bracts, note stipitate prolongation at
base of fruit and lateral insertion of flower near shoot apex. M
1-locular, many-seeded ovary in cross-section. (Takhtajan
1982, with exception ofL which is from Balslev 1996)
253
Fig. 65A-J. Juncaceae. A-E Luzula spicata. A Habit showing
spikelike inflorescence. B Closed leaf sheath. C Flower sub-
tended by ciliate floral bract. D Loculicidal capsule enclosed
by tepals and subtended by ciliate bracts. E Seed. F, G ]uncus
balticus. F Habit showing creeping rhizome, sheathing basal
cataphylls, absence of blade-bearing leaves, and pseudolateral
paniculate inflorescence subtended by stemlike bract. G
Capsule. H- J f. triglumis. H Habit, showing caespitose growth.
I Capsule, note lack of floral bracts below flower. J Seed, the
outer seed coat forming extensions. (Takhtajan 1982)
(Behnke 1981). Vessels are present in roots, stems
and leaves. In Prionium and in Rostkovia,
Oxychloe and the other Andean genera, the vessels
have predominantly scalariform perforation
254 Juncaceae
plates, while in ]uncus and Luzula vessels with
simple perforation plates are present in all organs
(Cheadle and Kosakai 1973).
The root has a 1-layered epidermis with the cells
arranged in longitudinal rows. From normal epi-
dermal cells smaller ones are cut off to form rows
of alternating small and large cells. The small cells
produce the root hairs. The cortex has several lay-
ers of radially arranged parenchyma cells, which
may break down to form airspaces alternating
with lamellae. The endodermis is one cell layer
thick with wall thickenings U-shaped in tran-
section. The central core has 1 to several vessels,
each associated with a group of phloem cells.
The anatomy of the rhizome is similar to that of
the root. An epidermis of a single cell layer covers
a multi-layered parenchymatous cortex, which
may have radially arranged lamellae alternating
with air spaces. A single-layered endodermis with
thickened inner and anticlinal walls separates the
cortex from the central core, in which there are
several vascular bundles scattered in parenchyma-
tous tissue.
The trunk of Prionium, homologous to the rhi-
zome of other genera, has a thin-walled epidermis,
a layer of suberised cells, a cortex of loosely
packed lobate parenchyma cells with dispersed
cortical bundles and a central cylinder with scat-
tered amphivasal bundles. Zimmermann and
Tomlinson (1968) note that this vascular pattern is
similar to that of palms.
The flowering stem in Juncaceae apparently
lacks an endodermis. The vascular bundles are
usually arranged in one or a few peripheral con-
centric rings. The central pith is hollow or consists
of aerenchyma of star-shaped cells.
The leaf epidermis is usually 1-layered with lon-
gitudinal cell rows, but several-layered in ]uncus
imbricatus and related species. The stomata are
paracytic and in xeromorphic species they are of
the Grass type. The shape of the leaf in section
varies greatly and is taxonomically important. In
the broad leaves of Prionium the vascular bundles
are equidistant from each leaf surface. Each
bundle is connected to the surface by parenchyma
girders separated by blocks of chlorenchyma with
air canals. Rostkovia has a well-developed hypo-
dermis of thick-walled fibres that are absent along
2 longitudinal bands, to which the stomata are
restricted. Luzula has flat, unspecialised leaves. In
]uncus several progressive evolutionary series can
be distinguished in the leaf anatomy. Some species
have flat unspecialised leaves like Luzula. In
others, the adaxial epidermis consists oflarge hya-
line cells and is occasionally many-layered where
the leaves are channelled. The involution may be-
come complete so that the leaf is terete and thus
the adaxial surface not recognisable. The terete
leaves evolved independently in 3 different groups
within ]uncus. In 2 of these the leaves of some
species are laterally compressed.
INFLORESCENCE STRUCTURE. The inflorescences
are highly variable and taxonomically important
(Buchenau 1865). Prionium, Luzula, and ]uncus
all have many-flowered inflorescences with per-
fect or gynodioecious flowers U. roemerianus).
Each inflorescence branch originates in the axil of
a bract and has a 2-keeled bifid prophyll on its
adaxial side. Often, though not in most ]uncus
species, 2 or more small bracts are inserted imme-
diately below the flower which they clasp (Fig.
64L). Single-flowered branchlets are developed
successively in a way resembling di- or monocha-
sia. Some ]uncus species have "racemose"
capitulae. Differences in internode length and
congestion of the inflorescences characterise
genera and subgenera. Prionium has a much-
branched panicle. Luzula may either have a
much-branched panicle (L. sylvatica and related
species), may be conspicuously sympodial (L.
elegans) or have a 1-3 times-branched anthela
(the proximal branches overtopping the distal
ones, L. pilosa and related species), or the flowers
may be congested into clusters arranged in pan-
iculate or spikelike inflorescences (L. spicata and
related species).
Distichia, Patosia and some species of Oxychloe
are dioecious with flowers originating in the axils
of normal leaves near the shoot apex. Each flower
is clasped by 1 to 4 membranous bracts. Rostkovia
and Marsippospermum have a solitary, perfect,
terminal flower. The Rostkovia flower has 2 bracts,
1 of which is leaflike and much longer than the
flower. The Marsippospermum flower has 1-2
small, membranous veinless bracts.
Determinate sympodial panicles occur also in
those species of ]uncus, in which the flowers have
2 small bracts. In ]. tenuis and related species the
branches of the inflorescence resemble sickle-
shaped drepania, while in ]. effusus and related
species they often resemble zigzag rhipidia, and
the lowermost inflorescence bract resembles a
continuation of the flowering stem and pushes the
inflorescence into a lateral position.
In the species of ]uncus, where the individual
flowers lack bracts, the flowers are often congested
into capituale, which according to Novara (1976)
have a racemose structure; these capitulae are ar-
ranged in cymes (anthelae), where the proximal
 Juncaceae 255

branches are longer than and overtop the distal and a hypostase may be present. (Development of
branches. the testa see Fruit and Seed, below.)
The subepidermal archesporia! cell cuts off a
FLOWER STRUCTURE. Most Juncaceae have trim- primary parietal cell, which gives rise to 1 or 2
erous flowers, but in some minute species of parietal layers. Sometimes 2 or 3 archesporia! cells
the ]uncus triformis complex they are dimerous are present, but only 1 is functional. After meiosis
(Ertter 1986). the chalaza! megaspore of a linear or T-shaped
The tepals are free and usually <7mm long (but tetrad develops into an embryo sac of the
in Marsippospermum up to 40mm), concave Polygonum type (Distichia, ]uncus, Luzula). The
or flat, usually lanceolate, obtuse, acute or acumi- antipodals are small. The polar nuclei fuse before
nate, green, dull-coloured, reddish brown, fertilisation and the secondary nucleus moves to
castanceous, almost black or rarely yellowish or the chalaza! end of the embryo.
white, often with membranous margins. The outer Endosperm development is helobial (Distichia,
tepals are often slightly longer, thicker and more ]uncus, Luzula). The small chalaza! endosperm
rigid than the inner. chamber becomes few-nucleate (to ca. 12 nuclei),
The 6 (or 3) stamens are free and alternate with sometimes at least few-cellular, or may remain
the tepals. The filaments are filiform or widened at uninucleate (Shah 1963). The chalaza! chamber ul-
the base and white or hyaline. The anthers are 4- timately degenerates, while the large micropylar
sporangiate, 4- or secondarily 2-locular by break- chamber becomes multicellular after several free
down of the inner wall, oblong to linear, basifixed, nuclear divisions and forms the storage tissue of
yellow to whitish, and dehiscent by longitudinal the seed. During endosperm development the
slits. In Distichia, Oxychloe, Patosia, Rostkovia main part of the nucellus is absorbed.
and Marsippospermum the connectives are api- The embryogeny is of the Onagrad type, Luzula
cally mucronate and exceed the thecae. (]uncus) variation. This variation, which also oc-
The gynoecium consists of 3 (rarely 2) united curs in Cyperaceae, Bromeliaceae and some other
carpels opposite the outer tepals. The ovary is su- families, is characterised by an early formation of
perior, globular to oblong, and 3- or 1-locular. The the dermatogen initials through periclinal divi-
common style is 0.5-10 mm long and continues sions of the quadrant cells of the proembryo. The
into 3 linear stigmatic branches with linear to ta- suspensor is few-celled and weakly developed.
pering stigmatic lobes. Most genera have many
ovules on an axile placenta or on placentas on POLLEN.MORPHOLOGY. The pollen tetrads, as far
parietal intruding septae. In Luzula the unilocular as known, are surrounded by a common exine, ca.
ovary has 3 basal ovules only. 1 [lm thick, smooth or minutely granulate (Fig.
66). Apertures are obscure, and recognisable only
EMBRYOLOGY. The family is apparently embryo- as diffuse areas with slightly thinner exine.
logically uniform, but only ]uncus and Luzula The pollen tetrads therefore have been described
have been investigated in detail (e.g. Wulff 1939; as inaperturate, monosulcate, monoporate or
Shah 1963; see also Davis 1966). monoporate-ulcer( oid)ate. Meyer and
The anther wall formation is of the monocotyle- Yaroshevskaya {1976) describe the apertures as "a
donous type. The epidermis is followed by the reduced distal furrow". Tetrad size varies from
endothecium, 1 middle layer and a 1-layered 24-70 [lm. Oxychloe, Patosia, Rostkovia and
secretory tapetum, which consists of uninucleate Marsippospermum have the largest tetrads, ]uncus
cells. and Luzula are intermediate, and Prionium has
Microsporogenesis is of the simultaneous type; the smallest. The thin exine is related to the pre-
sometimes after meiosis I an ephemeral cell plate dominantly anemophilous pollination in the fam-
has been observed (Wulff 1939: reduced succes- ily. The tetrahedral tetrads link Juncaceae to
sive type). The tetrads are usually tetrahedral. The Thurniaceae, and the juncaceous tetrad has been
generative cell is cut off towards the proximal (in- suggested as a precursor for the cyperaceous tet-
ner) pole of each microspore. The pollen grains rad in which 3 of the 4 pollen grains disintegrate.
are 3-celled when shed (]uncus, Luzula, Oxychloe)
and remain in tetrads. KARYOLOGY. A diffuse centromere spread over
The ovules are anatropous, bitegmic and the length of chromosome occurs at least in
(weakly) crassinucellate. The inner integument is Luzula. The spindle fibres attach at several points
2-layered and the outer 2-layered (]uncus) or 3-4- along the chromosomes. Consequently, segments
layered (Luzula, Distichia). A funicular obturator broken off from the chromosomes move to the
256 Juncaceae

FRUIT AND SEED. The fruit is a capsule of variable

shape, globular or more often ovoid to obovoid,
apically truncate, obtuse, acute, acuminate, or
beaked, terete to trigonous or 3-keeled. The num-
ber of locules in the capsule usually agrees with
that in the ovule except in ]uncus repens, ].
uruguensis and species of the ]. drummondii
group, which have free central placentation
through disintegration of the septa in the course
of capsule development. Other species are only
partly 3-locular, the locules being apically
confluent.
Dehiscence is normally loculicidal, but in
Distichia, Oxychloe and Patosia less well defined
and sometimes circumscissile (Balslev 1996). The
valves may inflex, reflex completely, separate only
apically, or separate in the middle and remain
Fig. 66. Juncacaeae. Luzula pilosa, pollen tetrad, SEM X 1700. connate at the apex. In Luzula the capsule
(Palynological Laboratory Stockholm) contains 3 seeds, in the other genera usually 15 to
120.
The seeds are ovoid, obovoid, ellipsoid, oblong
poles and remain functional in the genome or fusiform, often oblique, usually 0.3-2.5 mm
(Nordenskiold 1964). Therefore chromosome long, and 0.1-1 mm wide, but, when fusiform,
fragmentation is obviously important in the evo- sometimes up to 4 mm long. When crowded in the
lution in Luzula. capsule they are sometimes angular. The raphe is
Chromosome numbers reported for 67 of the usually visible as a fold along 1 side of the seed
75 species of Luzula range from 2n = 6 to 2n = 84; coat. The ends are rounded or taper into apiculate
2n = 12, 24, 36, and 48 being the most common or tail-like extensions. In ]uncus the main part of
numbers. Most numbers are multiples of 6, which the seed coat consists of the inner layers of the
is suggested to be the basic number. The number outer and inner integuments. In fusiform seeds
2n = 6 in L. elegans is probably derived in this the outer integument loosely envelops the seed
specialised species. and is elongated in the micropylar region. In
In ]uncus chromosome numbers are reported Luzula the inner layer of the inner integument is
for ca. 65 of the 260 species and range from 2n = thickened while the outer integument forms a thin
18 to 2n = 130. Predominant numbers are 2n = 40 membrane. Sometimes, as in species of ]uncus and
and 80. However, a few reports of 2n = 20 or 18 in Luzula elegans, the outer integument becomes
suggest that x = 10 is the base number for the mucilaginous (Netolitzky 1926).
genus. 2n = 16 is reported for Oxy chloe. The sculpturing of the testa varies from smooth
to reticulate and is characteristic for species or
PoLLINATION. Anemophily is the predominant groups of species (Engelmann 1866, 1868; Ertter
mode of pollination. Adaptations include small 1986; Balslev 1996).
flowers often aggregated in many-flowered inflo- The seeds of many species of Luz ula have a
rescences, prominent stigmatic surfaces, abun- fleshy apical ( = chalaza} e.g. L. pilosa) or basal
dant pollen production, and smooth pollen (= micropylar, e.g. L. multiflora) appendage or
tetrads. Exserted anthers are rare but present in caruncle.
some species of ]uncus. Temporal separation of The larger part of the seed is occupied by a
pollen release and stigmatic receptivity (often starchy endosperm. As in grasses, the peripheral
protogyny) is common. However, autogamy is fre- endosperm layers remain free from starch, but
quent, and cleistogamy occurs in ]uncus bufonius contain aleuron and oil (Shah 1963). The embryo
and ]uncus capitatus. consists mostly of the single cotyledon and is
Entomophily is known in some central Euro- small, straight, cylindrical to conical, and located
pean species of Luzula with bright, white flower medially near the micropyle.
heads and in some ]uncus species in the
Himalayas. Insects may also visit flowers of]uncus DISPERSAL AND REPRODUCTIVE BIOLOGY. Veg-
species probably for collecting pollen. etative propagation by means of large rhizome
 Juncaceae 257

systems is common in many species, especially in AFFINITIES. The Juncaceae are perhaps most
]uncus. Vegetative propagation connected with closely related to the three Thurnia species
water dispersal is known in, e.g. ]uncus bulbosus (Thurniaceae) of northern S America. Both have
and f. pelocarpus. Dispersal is otherwise by seeds. simultaneous microsporogenesis, tetrad pollen
The mucilaginous seeds of some species are and a similar floral structure, but differ in the vas-
adapted to epizoochory and their small size cular anatomy of the leaf, which in Thurniaceae
and ability to adhere to animals explain the has inverted bundles and silica bodies. In most
wide and discontinuous distributions of many recent systems of classification the Juncaceae are
species. The fusiform seeds of some ]uncus species placed with the Thurniaceae in a separate order,
may be anemochorous or (and) hydrochorous. Juncales. There is much evidence favouring a close
Carunculate seeds of Luzula are dispersed relationship between Juncaceae and Cyperaceae.
by ants. The swelling of the testa upon moisten- This is also confirmed by the morphologically
ing in some species appears to cause capsule based cladistic study by Simpson (1995) and the
dehiscence. molecular analysis of Duvall et al. (1993).
The previous association of the Juncaceae with
PHYTOCHEMISTRY. Calcium oxalate (both in the Liliales s.l., based on its liliaceous floral struc-
form of crystals and raphides) is absent and silica ture, is contradicted by a great number of features,
bodies are lacking or at least very rare; Lanning such as stomatal structure, starchy endosperm,
(1972) found silica in the leaves of ]uncus interior anemophily and epicuticular wax structure.
and f. bufonius.
Apart from common phenolics, anthocyanins DISTRIBUTION AND HABITATS. The family is COS-
based on luteolinidin, luteolin 5-methyl ether, mopolitan but in the tropics restricted to the high-
phytotoxic phenolic juncosol, and two coumarins lands (Balslev 1996). ]uncus and Luzula are
(7,8-dihydroxycoumarin, daphnetin and its 8- distributed as the family but have most species in
methyl ether) have been recorded (Williams and the northern temperate zone, though most sub-
Harborne 1975). The presence ofluteolin 5-methyl genera of ]uncus occur in the southern hemi-
ether and daphnetin (in addition to the absence of sphere. Prionium is endemic to the Cape Province
tricin and C-glycosyl flavones) was believed to dis- of S Africa. Distichia, Oxychloe and Patosia are
tinguish the Juncaceae from the Poaceae and the entirely Andean. Rostkovia and Marsipposper-
Cyperaceae when discovered, but these com- mum are trans-Pacific, occurring in New Zealand
pounds have now been reported in the Cyperaceae and Patagonia, Rostkovia also on Tristan da
as well. Cyanogenetic glycosides seem to be re- Cunha and in Ecuador. The greatest concentration
stricted to the septate-leaved species of ]uncus of genera is in the Andes, where all genera except
(Hegnauer 1986). Prionium occur. Marked disjunctions in the
family include the trans-Pacific distribution of
SUBDIVISION AND RELATIONSHIPS WITHIN THE Rostkovia and Marsippospermum. The family
FAMILY. The genera of Juncaceae comprise four probably originated in S America after the
natural groups. (1) Prionium stands clearly apart breakup of Gondwana.
in many characters though not in floral structure Most species are terrestrial or semiaquatic, oc-
and tetrad pollen. It deviates especially in its hard cupying open grass lands or swamps. Some spe-
persistent trunk and leaf anatomy. (2) The cies of ]uncus are anthropochorous and weedy,
cushion-forming genera Distichia, Oxychloe and some apparently restricted to eutrophic grass
Patosia form a natural group. They all have single lands. Several species of ]uncus are halophytic
lateral flowers and are dioecious except for one (Snogerup 1993). The cushion-forming genera
species of Oxychloe. They are all restricted to the Distichia, Oxychloe, and Patosia are well adapted
Andes. (2) Rostkovia and Marsippospermum are to harsh diurnal freezing and thawing; they are
characterised by single terminal flowers and ta- conspicuous elements of wetlands in the high
pering stigmatic branches. They also have mucro- Andes, sometimes reaching the altitudinal limit of
nate anthers and in this resemble the previous the vegetation. Prionium grows in permanent
group. These two genera have a trans-Pacific dis- streams and rivers and is important in preventing
tribution. (4) ]uncus and Luzula are herbaceous erosion during seasonal flooding.
but generally not cushion-forming, they have
many flowered inflorescences and emucronate PARASITES. Many species of ]uncus and Luzula
anthers. Their distribution is mainly north- have been reported as hosts for parasitic fungi of
temperate. the genera Cintracia, Claviceps, Puccinia, Tolypos-
258 Juncaceae
parium, Urocystis and Ustilago. These data suggest
a relationship between the Juncaceae and
Cyperaceae (Savile 1979).
PALAEOBOTANY. Juncaceae are poorly repre-
sented in the fossil record. Tetrads of pollen were
reported from the Neogene of Russia and the
megafossil ]uncus crassulus is known from the
Miocene ofN America. Leaves, stems, and fruits of
]uncus were reported from the Tertiary and well-
preserved inflorescences of Luzula from the
Upper Oligocene of Germany.
ECONOMIC IMPORTANCE. Few important eco-
nomic uses ofJuncaceae are known. Many species
of ]uncus and Luzula are components in pastures
throughout the world. ]uncus effusus is used for
making mats in Central America and Japan, and
so is]. rigid us in theN ear East. In the treeless puna
of Peru, blocks of the cushions of Distichia
muscoides are cut for fuel.
KEY TO THE GENERA
1. Plants with distinct woody trunk (S Africa) 1. Prionium
- Plants herbaceous 2 etal placentation; stigmatic lobes tapering; capsule
2. Flowers solitary; anthers mucronate 3 orbicular. Two spp., subantarctic islands from
- Flowers few to numerous, in inflorescences; anthers not
mucronate 7 and S American mainland in Patagonia and
3. Flower terminal on the flowering stem; plants not forming
cushions 4
- Flower lateral in axil of foliage leaf; plants forming cushions
5
4. Flower subtended by 1-2 membranous bracts shorter than
tepals; capsule longer than wide 2. Marsippospermum
Flower subtended by 2 bracts, one much longer than tepals;
capsule orbicular 3. Rostkovia
5. Leaves distichous; capsule stipitate when ripe 4. Distichia
- Leaves spiral; capsule sessile 6 shoots forking, 5-15cm long, covered with short,
6. Leaf apex acute but not hard and prickly 5. Patosia
- Leaf apex acute and prickly 6. Oxychloe
7. Leaf sheaths closed; leaf margins ::<:: ciliate; capsule 3-seeded
7. Luzula
- Leaf sheaths open or rarely closed; leaves glabrous; capsule
15- to 120-seeded 8. ]uncus
Genera of the Juncaceae
1. Prionium E. Meyer Fig. 64A-F
Prionium E. Meyer, Linnaea 7: 131 (1832).
Basally woody, with a trunk (rarely) to >1m long
and 5-IOcm in diam., continuing into a subterra-
nean rhizome; leaves in a terminal rosette, linear,
tapering, flat or canaliculate, serrate; inflorescence
a terminal erect panicle to 1m tall; anthers
emucronate; styluli free; ovary 3-locular. Only one
sp., P. serratum (L.f.) Drege, rivers and streams,
S Africa.
2. Marsippospermum Desv. Fig. 64G-I
Marsippospermum Desv., J. Bot. (Desv.) 1: 328 (1809).
Herbs to 60 em tall; rhizome creeping, scale-
bearing, sympodial; flowering stems erect, terete;
leaves erect, terete; flowers solitary, terminal;
bracts 1-2, membranous; anthers mucronate;
ovary 1-locular, with parietal placentation; stig-
matic lobes tapering; capsule elongate, ligneous.
Three spp., New Zealand and Patagonia.
3. Rostkovia Desv.
Rostkovia Desv., J. Bot. (Desv.) 1: 324 (1809).
Herbs to 30cm tall; rhizome creeping or ascend-
ing; flowering stem erect, longitudinally ridged;
leaves linear; flowers solitary, terminal, supported
by 2 bracts, the lower foliaceous, the upper bracte-
ate; anthers mucronate; ovary !-locular with pari-
New Zealand to S Georgia and Tristan da Cunha,
Ecuador.
4. Distichia Nees & Meyen Fig. 64K-M
Distichia Nees & Meyen in Meyen, Observ. bot.: 128 (1843).
Cushion-forming herbs to several metres in diam.;
distichous leaves; dioecious; flowers lateral, in
axils of foliage leaves; staminate flowers long-
pedicellate, projecting above the cushion, anthers
mucronate; pistillate flowers in axils of subapical
leaves, with only the stigmatic branches extending
above the cushion; ovary !-locular with parietal
placentation; capsule elevated on a stipe. Three
spp., high altitudes in the Andes from Venezuela
to N Argentina.
5. Patosia Buchenau
Patosia Buchenau, Bot. Jahrb. Syst. 12: 63 (1890).
Cushion-forming herbs to several metres in diam.;
shoots forking, 5-15 em long, covered with short,
spiral leaves; dioecious or rarely with a few her-
maphrodite flowers; flowers lateral in axils of
foliage leaves; staminate flowers long-pedicellate,
 Juncaceae
projecting above the cushion, anthers mucronate;
pistillate flowers in axils of subapical leaves, with
only the long stigmatic branches extending above
the cushion; ovary 3-locular; capsule sessile. Only
one sp., P. clandestina (Philippi) Buchenau in the
Andes from Bolivia to Argentina.
6. Oxychloe Philippi
Oxychloe Philippi, Florula Atacamensis: 52 (1860).
Cushion-forming prickly herbs to several metres
in diam., less dense than the two previous genera;
shoots forking, 5-30 em long, covered with spiral
leaves with hard, prickly apices; usually dioecious;
one sp. with hermaphrodite flowers; flowers lat-
eral, in the axils of foliage leaves, staminate flow-
ers with mucronate anthers, pistillate flowers
concealed in leaves; ovary 3-locular; capsule
sessile. Six spp., high Andes from S Peru to C
Argentina.
7. Luzula DC. Fig. 65A-E
Luzula DC., Fl. fran<;. 3. ed. 1: 198 (1805), ibid. 3: 158.
Perennial, rarely annual (subgen. Marlenia) rhi-
zomatous herbs; leaves spiral, with closed sheath,
and flat, ciliate, acute, acuminate, aristate, or
callose-tipped blade; inflorescence many-
flowered, paniculate, spicate or anthelate; flowers
often ± congested into headlike clusters; bracts
1-2, membranous; flowers hermaphrodite; an-
thers emucronate, ovary 1-locular, with 3 basal
ovules; seeds sometimes carunculate. About 75
spp., a cosmopolitan genus traditionally divided
into the three subgen. Luzula, Anthelaea and
Pterodes (but see Novikov 1990b); one sp. of
Anthelaea is often treated as subgen. Marlenia
(Ebinger 1963).
8. ]uncus L. Fig. 65G-J
]uncus L., Sp. Pl. 1: 325 (1753).
Perennial and rhizomatous or occasionally annual
herbs; leaves spiral; sheath open or rarely closed,
blade linear, flat, canaliculate, terete, or laterally
compressed; inflorescence few- or many-flowered,
separate or congested into headlike clusters;
flowers hermaphrodite, with or without clasping
bracts; anthers emucronate; ovary 3- or !~locular;
capsule 15- to 120-seeded. About 250-275 spp.,
cosmopolitan. Buchenau (1875, 1890, 1906) di-
vided /uncus into 11 subgenera based on inflo-
259
rescence and leaf characters. Novikov (1990a) re-
duced the number of subgenera to two and largely
treated Buchenau's subgenera as sections.
To be excluded:
Voladeria Benoist
Voladeria Benoist, Bull. Soc. Bot. Fr. 84: 637 (1937).
This genus was referred to Juncaceae when de-
scribed. The type is lost, but from the description,
which mentions 6 perianth segments and a 1-
seeded ovary, it appears to belong to Oreobolus in
the Cyperaceae, which is quite common at the type
locality of Voladeria in Paramo del Angel in
Ecuador.
Selected Bibliography
Balslev, H. 1996. Juncaceae. Flora Neotropica Monograph no.
68. New York: The New York Botanical Garden.
Behnke, H.-D. 1981. See general references.
Buchenau, F. 1865. Der Bliithenstand der Juncaceen. Jahrb.
Wiss. Bot. 4: 385-440.
Buchenau, F. 1875. Monographie der Juncaceen vom Cap.
Abh. Naturwiss. Ver. Bremen 4: 393-512.
Buchenau, F. 1890. Monographia Juncacearum. Bot. Jahrb.
Syst. 12: 1-495.
Buchenau, F. 1906. Juncaceae. In: Engler, A. (ed.) Pflan-
zenreich IV, 36: 1-284.
Cheadle, V.I., Kosakai, H. 1973. Vessels in Juncales. I.
Juncaceae and Thurniaceae. Phytomorphology 23: 80-87.
Cutler, D.F. 1969. Juncales. In: Metcalfe, C.R. (ed.) Anatomy of
the monocotyledons 4. Oxford: Clarendon Press, pp. 1-358.
Davis, G.L. 1966. See general references.
Duvall, M.R. et al. 1993. See general references.
Ebinger, J.E. 1963. A new subgenus in Luzula (Juncaceae).
Brittonia 15: 169-174.
Engelmann, G. 1866, 1868. A revision of the North American
species of the genus ]uncus, with a description of new and
imperfectly known species. Trans. Acad. Sci. St. Louis 2:
424-499.
Ertter, B. 1986. The ]uncus triformis complex. Mem. N.Y. Bot.
Gard. 39: 1-90.
Hegnauer, R. 1986. See general references.
Lanning, F.C. 1972. Ash and silica in ]uncus. Bull. Torrey Bot.
Club 99: 196-198.
Meyer, N.R., Yaroshevskaya, A.S. 1976. The phylogenetic sig-
nificance of the development of pollen grain walls in
Liliaceae, Juncaceae and Cyperaceae. In: Ferguson, I.K.,
Muller, J. (eds.) The evolutionary significance of the exine.
Linnaean Society Symposium Series No. 1, pp. 91-100.
Netolitzky, F. 1926. Anatomie der Angiospermen-Samen. In:
Handbuch der Pflanzenanatomie, Vol. 10. Berlin: Born-
traeger.
Nordenskitild, H. 1964. The effect of X-irradiation on diploid
and polyploid Luzula. Hereditas 51: 344-374.
Novara, L.J. 1976. Contribuci6n al conocimiento de las
inflorescencias de ]uncus y su significaci6n taxon6mica.
Kurtziana 9: 41-61.
260 Juncaceae/J uncaginaceae
Novikov, V.S. 1990a. Synopsis of the genus ]uncus L.
(Juncaceae). Bjull. Moskovsk Obsc. Isp. Prir., Odt. Bioi. 95
(5): 111-125.
Novikov, V.S. 1990b. The synopsis of the genus Luzula DC.
system (Juncaceae). Bjull. Moskovsk. Obsc. Isp. Prir., Odt.
Bioi. 95 (6): 63-70.
Savile, D.B.O. 1979. Fungi as aids in higher plant classification.
Bot. Rev. 45: 377-503.
Shah, C.K. 1963. The life history of ]uncus bufonius. ]. Indian
Bot. Soc. 42: 238-251.
Simpson, D. 1995. Relationships within Cyperales. In: Rudall,
P.J., Cribb, P.J., Cutler, D.F., Humphries, C.J. (eds.) Mono-
cotyledons: systematics and evolution, Vol. 2. London:
Royal Botanic Gardens, Kew, pp. 497-509.
Snogerup, S. 1993. A revision of ]uncus subgen. ]uncus
(Juncaceae). Willdenowia 23: 23-73.
Takhtajan, A.L. 1982. See general references.
Williams, C.A., Harborne, J.B. 1975. Luteolin and Daphnetin
derivatives in the Juncaceae and their systematic signifi-
cance. Biochem. Syst. Ecol. 8: 181-190.
Wulff, H.D. 1939. Die Pollenentwicklung der Juncaceen nebst
einer Auswertung der embryologischen Befunde hinsi-
chtlich einer Verwandtschaft zwischen den Juncaceen und
Cyperaceen. Jahrb. Wiss. Bot. 87: 533-556.
Zimmermann, M.H., Tomlinson, P.B. 1968. Vascular con-
struction and development in the aerial stem of Prionium
(Juncaceae). Am.]. Bot. 55: 1100-1109.
Juncaginaceae
R.R. HAYNES, D.H. LES,
and LB. HoLM-NIELSEN
Juncaginaceae Rich., Demonstr. Bot.: IX (1808), nom. cons.
Annual or perennial, rhizomatous, glabrous herbs,
emergent or with floating leaves, in fresh or brack-
ish waters. Roots fibrous or tuberous. Rhizomes
slender to stout. Leaves rosulate, linear, !-several-
veined, dorsiventrally flattened, terete, or uni-
facial, with open, sheathing base; sheath adnate
to the blade, often remaining as fibrous cluster
after the leaves have decayed, with 2 auricles;
intravaginal scales present, membranous. Inflores-
cences ebracteate, terminal spikes or spikelike
racemes, scapose or not; flowers all hermaphro-
dite, or pistillate at the base, hermaphrodite in the
middle, and staminate (or lacking) distally on the
inflorescence, trimerous, dimerous, or mono-
merous. Perfect and pistillate flowers hypogynous.
Perianth mostly present, rarely absent; segments
6 or 1, in 1 or 2 whorls, when in 1 whorl, then
with 1 segment adnate to anther, when in 2 whorls,
then each whorl with 3 free segments; stamens
antetepalous, 0, 1, 4, or 6, sessile or subsessile,
when 4 or 6 in 2 whorls of2 or 3; anthers 2-thecate,
dehiscing by longitudinal slits. Carpels 0, 1, 3, or 6,
when 3 or 6, then coherent or weakly connate
but separating when mature, or rarely apo-
carpous; locules of same number as carpels;
stigma mostly sessile or on a short style, but
the basal pistillate mostly sessile or on a short
style, but the basal pistillate flowers of Lilaea
with an elongate, filiform style; ovules 1-few
per locule, erect, anatropous with basal placenta-
tion, in Maundia atropous and attached apically.
Fruits achenes, rarely with hooks or horns
at apex. Seeds solitary, exalbuminous; embryo
straight.
A family of subcosmopolitan distribution with
4 genera and about 12 species.
VEGETATIVE MORPHOLOGY. The Juncaginaceae
are rooted herbs growing in salt or freshwater
marshes or submersed in temperate freshwater
lakes (Fig. 67). The roots are filiform, unbranched
or branched. In Triglochin, they occasionally form
root tubers distally, which arise from the base of
the stem.
 Juncaginaceae
Fig. 67A-J. Juncaginaceae. A-C Triglochin palustre. A Habit.
B Flower. C Mature fruit. D-F T. calcitrapa. D Habit. E Flower.
F Immature fruit. G-J Tetroncium magellanicum. G Staminate
plant. H Staminate flower. I Pistillate plant. J Pistillate flower.
(Takhtajan 1982)
The plants are typically tufted with linear leaves
arising from an erect, sympodially branched,
cormous axis. Some species are stoloniferous,
with axillary, scale-bearing stolons. The leaves
have a distichous or spirodistichous arrangement
on the rhizome. They are sheathing at the
base. The blades are linear and terete to dor-
261
siventrally or laterally flattened. Intravaginal
squamules are present. Ligules are developed in
Triglochin.
VEGETATIVE ANATOMY. The Juncaginaceae have
paracytic or tetracytic stomata; they lack hairs
(apart from the numerous axillary squamules),
and the mesophyll is lacunose. The main vascular
bundles of the leaf usually have a sclerenchyma-
tous sheath (the outermost sheathing layer often
with U-shaped wall thickenings), but mechanical
tissue is not otherwise developed. Vessels (ques-
tionable in Lilaea ) are restricted to the roots and
are provided with lengthy scalariform perforation
plates. Lilaea is unique in possessing inconspicu-
ous hypodermal laticifers. Crystals occur some-
times in Triglochin, and condensed tannins are
found occasionally in, e.g., solitary mesophyll cells
of Lilaea. Starch is abundant in the stem and root
cortex of Lilaea and Triglochin and also in the leaf
sheath and leaf blade of Triglochin (Tomlinson
1982).
INFLORESCENCE AND FLORAL STRUCTURE. The
inflorescence ofJuncaginaceae is a scapose termi-
nal spike. The flowers are generally ebracteate, but
bracteate in the spikes of Lilaea. The flowers of
Triglochin are perfect, whereas those of Lilaea are
pistillate, hermaphrodite, or staminate, following
in this sequence on the inflorescence axis from
proximal to distal; Tetroncium is dioecious (Fig.
67.) In Lilaea, solitary pistillate flowers are also
borne basally on the scape. The flowers of
Triglochin have 2 whorls each of 3 tepals. There
are up to 6 stamens with tetrasporangiate anthers
which are sessile or on short filaments. The an-
thers dehisce by longitudinal slits. The up to 6
carpels in multicarpellary flowers are more or less
free or united basally and ventrally to a persistent
extension of the floral axis. For further informa-
tion see Tomlinson (1982).
EMBRYOLOGY. The anther wall formation is not
known. The endothecial thickenings are of the spi-
ral type. The tapetal cells are uninucleate and
plasmodial. Microsporogenesis is of the Succes-
sive type, and the homotypic division may aresult
in the formation of isobilateral, linear, T-shaped
or irregular tetrads, such as in, e.g., Butomopsis
(Limnocharitaceae). The pollen grains are dis-
persed as monads in the 2- or 3-celled stage
(Gardner 1976). The ovules are crassinucellate,
with a parietal cell cut off from the archesporia!
cell. The embryo sac is of the Polygonum type, and
endosperm formation is Nuclear.
262 Juncaginaceae
POLLEN MORPHOLOGY. The pollen grains are
dispersed singly. They are globose and inaper-
turate.
KARYOLOGY. Reports on chromosome numbers
are available for several species and include 2n =
12, 18, 24, 30, 32, 36, 48, 60, 96, 120. Base numbers
of x = 6, 8, or 9 have been proposed; it is likely that
the base number of the group is x = 6, and that
polyploidy and aneuploidy are responsible for the
variation (Larsen 1966).
PoLLINATION. The pollination in the Juncagina-
ceae is incompletely known but mainly by wind,
although water pollination may occur as well.
FRUIT. The fruit of Juncaginaceae is basically an
achene, but fruit condition varies. Triglochin has
6 carpels, 3 of which are sterile. These are filled
with parenchymatous tissue; the fruit develops
into a schizocarplike structure, in which the 3 fer-
tile carpels loosen from the others acropetally, to
form 1-seeded achenes (Fig. 67C). In Maundia
and Tetroncium the carpels likewise loosen from
each other and form separate achenes. The
monomerous pistil in Lilaea develops into a trian-
gular, 3-winged achene.
The testa is not sculptured. The embryo is
straight and has a massive cotyledon, a lateral
stem tip, and terminal radicle; it lacks a suspensor.
DISPERSAL. Although no published data exist, we
believe that birds are important in moving the
species over short distances from one water sur-
face to another.
PHYTOCHEMISTRY. The Juncaginaceae, like the
Scheuchzeriaceae, contain the cyanogenic gluco-
side triglochinin. Crystals of oxalate occur in the
tissues.
CLASS! FICA TION AND AFFINITIES. The genus
Lilaea is often treated as a monotypic family
(Lilaeaceae) that is closely related to Jun-
caginaceae (e.g., Hutchinson 1959; Tomlinson
1982). However, embryological, cytological and
palynological evidence indicate a very close rela-
tionship between Lilaea and other genera of
Juncaginaceae (Agrawal 1952; Larsen 1966;
Gardner 1976). Despite the presence of laticifers
(absent in other Juncaginaceae), various authors
(e.g., Cronquist 1981; Dahlgren et al. 1985) have
viewed the features of Lilaea as similar enough
to warrant the inclusion of the genus in the
Juncaginaceae. The presence of laticifers and
some uncertainty regarding the interpretation of
floral morphology in Lilaea and Juncaginaceae
inspired Tomlinson (1982) to retain the family
Lilaeaceae pending further study.
A close relationship between Juncaginaceae and
Scheuchzeriaceae has long been assumed, and
these families are often merged. A compelling ob-
servation is that both families contain the cyano-
genic glucoside triglochinin, which is not known
to occur in other flowering plant families (a simi-
lar derivative is known from Ranunculaceae). In
contrast, the vast number of anatomical and mor-
phological features that differed between Scheu-
chzeriaceae and Juncaginaceae indicate a more
remote relationship (Tomlinson 1982).
Juncaginaceae are traditionally placed (along
with Scheuchzeriaceae) among families in the
order Zosterales ("Najadales"), but have some-
times been considered as closely related to
Alismataceae (Alismatales).
Molecular phylogenetic studies (Les and Haynes
1995; D.H. Les, unpubl.) corroborate the close
relationship of Lilaea ("Lilaeaceae") and Tri-
glochin (Juncaginaceae), but resolve Scheuch-
zeria as relatively distant from these taxa. This
topology agrees well with the relative levels of
divergence in morphological and anatomical
features observed among these families.
Molecular cladograms depict Juncaginaceae
as basal to a clade consisting of Zosteraceae and
Potamogetonaceae. Scheuchzeriaceae are basal to
a clade comprising this group and the remaining
marine families of the Zosterales (Cymodocea-
ceae, Posidoniaceae, Ruppiaceae). Although the
loss of triglochinin from all marine Zosterales and
the Potamogetonaceae is indicated by these phylo-
genetic relationships, losses of secondary metabo-
lites are common in the Potamogetonaceae and
other submersed aquatic plants (Les and Sheridan
1990). All recent evidence confirms the placement
of Juncaginaceae among the Zosterales rather
than the Alismatales.
DISTRIBUTION AND HABITATS. The Juncagina-
ceae are nearly cosmopolitan in cold and tem-
perate regions of all vegetational zones. Lilaea,
Tetroncium, and Maundia are all basically South-
ern Hemispherical, although Lilaea reaches as far
north as southern Canada. Tetroncium is found
only in Patagonia and surrounding islands, and
Maundia is restricted to Australia. Triglochin is
distributed over most regions of the world, and,
like the family, is subcosmopolitan. It is likely that
the family has its origin in Gondwana (Raven and
Axelrod 1974).
 Juncaginaceae
All Juncaginaceae are bound to aquatic habitats.
They grow in fresh or salt marshes, on sea shores
or submersed in lakes and rivers. Whereas
Triglochin, Tetroncium, and Maundia are marsh
plants, Lilaea is an amphiphyte, living part of the
life cycle submersed in freshwater lakes, pools, or
small streams.
EcoNOMIC IMPORTANCE. The leaves of Triglo-
chin maritimum are edible and so is the rhizome
of T. procerum. Some species of Triglochin have
a high content of HCN and cause occasional
problems for livestock that graze in marshes.
KEY TO THE GENERA
1. Flowers both distally in scapose, spikelike racemes, these
perfect, pistillate and staminate, and basally on the scape in
leaf axils, these pistillate and 1-carpellate 4. Lilaea
- All flowers distally in scapose spikelike racemes; carpels 3,
4, or 6
2. Flowers 3-merous; ovules erect 1. Triglochin
- Flowers 2-merous; ovules erect or pendulous
3. Flowers unisexual; ovules basal, erect. Southern S America
2. Tetroncium
- Flowers perfect; ovules apical or pendulous. NE Australia
3.Maundia
1. Triglochin L. Fig. 67A-F
Triglochin L., Sp. Pl.: 338 (1753).
Emersed perennials, occasionally tuberous; stem
short, leaves few, terete, erect. Racemes spikelike;
perianth of 3 + 3 similar, conchiform, separate
segments; stamens mostly 2 + 2 or 3 + 3,
subsessile; carpels 3 or 6, coherent or weakly con-
nate, separating when mature, often embedded in
the angled floral receptacle; styles absent; the stig-
mas plumose or papillose. Fruits angled nutlets.
About 12 spp., cosmopolitan.
2. Tetroncium Willd. Fig. 67G-K
Tetroncium Willd., Ges. Naturf. Freunde Berlin Mag.
Neuesten Entdeck. 2: 17 (1808); Correa, Fl. Patagon. 2: 33-
37 (1969).
Emersed perennials with delicate, fibrous roots;
stem stout, erect, when old sometimes long and
reddish. Leaves few, semiterete, with concave
adaxial surface. Racemes spikelike, scapose. Flow-
ers dioecious, staminate with 4 tepals and 4 sessile
anthers, pistillate with 4 tepals and 4 carpels.
Fruits beaklike; styles persistent on refiexed pe-
duncles. Only one sp., T. magellanicum Willd.,
Tierra del Fuego, Patagonia, and the Falkland
Islands.
263
3. Maundia F. Muell.
Maundia F. Muell., Fragm. 1: 22 (1858).
Stoloniferous, emersed perennial with erect, short
stem. Leaves subequal to scape, semiterete,
sheathing. Spikes scapose, dense, many-flowered.
Flowers perfect, with 2-4 broad-spathulate tepals
and up to 8 stamens in 2 whorls. Carpels 4, weakly
connate, distinct at apex; styles absent; stigmas
linear. Fruits of 4-10 carpels, 4-angled; carpels
joined adaxially, their ventral spongy, their edges
recurved, and their beaks recurved. Only one sp.,
M. triglochinoides F. Muell., NE Australia: S
Queensland, N New South Wales.
4. Lilaea Humb. & Bonpl.
Lilaea Humb. & Bonpl., Pl. Aequinoct. 1: 221 (1808).
2
Emersed annual; roots without tubers; stems
3 short, slender. Leaves terete to fiat. Flowers both
on scapose spike and basal; the former perfect,
pistillate, and staminate, the basal pistillate and
long-styled. Perfect flowers with 1 tepal, the others
naked. Fruit a flattened to angular nutlet. Only one
sp., L. scilloides (Poiret) Hauman, temperate and
tropical montane marshes from southern Canada
to Chile and Argentina.
Selected Bibliography
Agrawal, V.S. 1952. The embryology of Lilaea subulata H. B.
K. with a discussion on its systematic position. Phytomor-
phology 2: 15-29.
Cronquist, A. 1981. See general references.
Dahlgren, R. et al. 1985. See general references.
Gardner, R.O. 1976. Binucleate pollen in Triglochin L. N. Z. J.
Bot. 14: 115-116.
Hutchinson, J. 1959. The families of flowering plants. II.
Monocotyledons. 2nd edn. Oxford: Clarendon Press.
Larsen, K. 1966. Cytotaxonomical note on Lilaea. Bot. Not.
119: 496-497.
Les, D.H., Haynes, R.R. 1995. Systematics of the subclass
Alismatidae: a synthesis of approaches. In: Rudall, P.J.,
Cribb, P.J., Cutler, D.P., Humphries, C.J. (eds.) Monocotyle-
dons: systematics and evolution. Royal Botanic Gardens,
Kew, pp. 353-377.
Les, D.H., Sheridan, D.J. 1990. Biochemical heterophylly and
flavonoid evolution in North American Potamogeton
(Potamogetonaceae). Am. J. Bot. 77: 453-465.
Raven, P.H., Axelrod, D.I. 1974. Angiosperm biogeography
and past continental movements. Ann. Mo. Bot. Gard. 61:
539-673.
Takhtajan, A.L. 1982. See general references.
Tomlinson, P.B. 1982. See general references.
264 Lemnaceae

Lemnaceae In Lemnoideae, the fronds are thin and leaflike

or inflated in some species. They are orbicular,
E. LANDOLT obovate or lanceolate with 1-21 veins. Small pa-
pillae may occur in the upper surface especially
above the node and the tip. In Spirodela the base
of the frond is surrounded by a bifid scale of which
1lobe covers the root primordia on the lower side.
Roots originate at the node on the lower part of
the frond. Spirodela has up to 21 roots, Lemna
Lemnaceae Gray, Nat. Arr. Brit. Pl. 2: 729 (1821), nom. cons. only 1. The roots are surrounded at the base by a
root sheath and enclosed at the tip by a rootcap.
Herbs reduced to smallleaflike poorly differenti- Root hairs are lacking and the roots function
ated fronds, floating on or below the water surface, mainly in stabilising the floating plants. In
mostly reproducing vegetatively. Fronds 1-several Wolffioideae, the fronds are either thin and often
cohering together, thin or thick, 0.5-18mm long, transparent and orbicular to linear or falcate
intensely or transparent green, sometimes red- (Wolffiella, Fig. 69), or thick and globular to cylin-
dish, surrounded at the base by a bifid scale drical or boat-shaped convex (Wolffia). The small-
(Spirodela) or without, consisting of a lower and est species of flowering plants ( Wolffia angusta
an upper epidermis and several cell layers be- and W. globosa) reach a length, width and thick-
tween, with aerenchyma or without ( Wolffia), with ness of 0.8/0.4/l.Omm and 0.8/0.6/0.7mm,
1-16(21) veins (Lemnoideae) or without veins respectively.
(Wolffioideae); with 2 flat lateral reproductive There are three morphological interpretations
pouches at the base of the frond (Lemnoideae) or of the frond as follows: ( 1) The frond corresponds
with 1 terminal pouch or cavity (Wolffioideae). to a leaf (e.g. Hoffmann 1840); (2) the frond cor-
Roots 1-21 (Lemnoideae) or absent (Wolf- responds to a shoot of leaflike shape (e.g.
fioideae), surrounded at the base by a tubular Hegelmaier 1868; Brunaud 1974); (3) the basal
sheath and enclosed at the tip by a cap; root hairs part of the frond represents a shoot, and the distal
lacking. Flowering and fruiting rare in most spe- part represents a phyllome (Engler 1877; Hegel-
cies. Flowers 1-2 per frond, bisexual, either sur- maier 1895; Arber 1919; Meusel 1951; Kandeler
rounded by an utricular to scalelike leaf and 1979). According to the last theory, which is the
emerging from the lateral pouches or originating most convincing and mostly accepted one, the
in a bare cavity on the upper surface. Stamens 2 scalelike leaf of Spirodela corresponds to a
or 1 per flower, 1- or 2-thecate. Ovary 1-locular, prophyll.
bottle-shaped, tapering into a stylar duct with a Many species of the Lemnaceae are able to form
spherical stigma producing a droplet at anthesis. turions in order to survive unfavourable condi-
Ovules anatropous, amphitropous or atropous, tions. These fronds are smaller than normal
bitegmic. Fruits with dry, decaying pericarp. fronds and contain more starch and fewer air
Seeds mostly 1 (up to 5 in Spirodela and Lemna), spaces. They sink to the bottom of the water.
ovoid, retained in pericarp or released, longitudi-
nally ribbed or nearly smooth, endosperm VEGETATIVE ANATOMY. The fronds have 1-10
present. layers of parenchyma between the upper and
The family has four genera and ca. 35 species lower epidermis. This tissue is aerenchymatous
and is distributed worldwide. (except in Wolffia) either throughout the frond or
only around the node. Under certain conditions
VEGETATIVE MORPHOLOGY. The leaflike body gibbous fronds caused by enlarged air spaces are
(frond) of the Lemnaceae is poorly differentiated formed. Veins (only present in Le~noideae) origi-
(Fig. 68). It consists of the rear or basal portion, nate in the node and run through the distal sec-
which is behind the node (vegetative apex) where tion. In the roots, the reduced vascular strands are
the daughter fronds are formed by budding either represented by a central strand of elongated cells.
in flat lateral pouches (Lemnoideae) or in 1 termi- The veins and the strand of elongated cells in roots
nal pouch or cavity (Wolffioideae); a row of in Spirodela and most species of Lemna contain
elongated cells runs from the node to the base some tracheids. In Lemnoideae certain cells con-
continuing as a stipe that connects the parent- and tain oxalate raphides; druses and raphides occur
daughter fronds and the front or distal portion, in Spirodela. Brown pigment cells (myriophylline
which contains veins in the Lemnoideae. idioblasts) are characteristic of Spirodela and
 Lemnaceae
~
0
some of Wolffiella and Wolffia (visible only in
dead fronds as brown dots). Stomata of Lemna
species (except flowering L. trisulca) lack chloro-
plasts and are always open; in Spirodela, Wolffiella
and Wolffia the stomata contain chloroplasts or at
least starch grains. It is known that the stomatal
opening is controlled in S. punctata.
The walls of the epidermal cells are straight
(Spirodela intermedia, Wolffia, Wolffiella) or un-
dulated (Spirodela punctata, Lemna).
INFLORESCENCE AND FLORAL STRUCTURE. The
reproductive part of Lemnaceae is interpreted
either as a single flower, or as an inflorescence
in which all flowers are reduced to either a single
stamen or a single carpel. If the reproductive zone
is treated as an inflorescence, the scalelike leaf
(prophyll) is called spathe. Here the interpretation
as a flower is favoured. In Lemnoideae the flower
265
Fig. 68A-O. Lemnaceae. A-C Spirodela polyrhiza. A Mother
frond with adhering offspring, dorsal view, X4. B Ventral
view, the sheaths of the secondary roots covered by the ventral
scale, X4. C Detail of flowering frond with style and two sta-
mens enveloped by the scalelike leaf, X 12. D-H Lemna
aequinoctialis. D Mother frond with adhering offspring, X8. E
Frond with fruit and daughter frond, dorsal view, X8. F Seed,
X20. G Seed, longitudinal section, X60. H Germinating seed,
showing plumule and first frond, X 10. 1-L L. trisulca. I Sub-
merged plant, X4. JPart of flowering frond, dorsal view, X8. K
Stamen, X 16. L Flower surrounded by the scalelike leaf, X 16.
M Wolffia arrhiza, frond with daughter frond, X 12. N, 0 W.
microscopica. N Flowering plant with daughter frond; X 12. 0
Flower (1 stamen, 1 pistil); X24. (van der Plas 1971 )
consists of 1 membranous scalelike leaf (prophyll)
at the base, 2 4-sporangiate stamens and 1 pistil.
In Wolffioideae the scalelike leaf (prophyll) is
lacking and the single stamen is 2-sporangiate. In
Lemnoideae, the flowering organs originate in the
same pouches in which the daughter fronds are
266 Lemnaceae

lines of the anthers contain pigment cells. Raphide

cells are present in the anther walls of Lem-
noideae. The filaments have at the base a row of
ring tracheids. The pistil of Lemnoideae is in-
serted above the 2 stamens (seen from the upper
frond surface) and between them (seen from the
base of the frond). In Wolffioideae, the pistil is at
the base and proximal to the single stamen. The
pistil is bottle-shaped and tapers into a short style.
The stigma is funnel-shaped, tapering into a style
duct. Pigment cells are present in the stigma of
Spirodela and some species of Wolffioideae. Most
Lemnaceae have only 1 ovule (except Spirodela,
Lemna gibba, and L. disperma with up to 7
ovules). The ovules are anatropous if more than
1 or amphitropous or atropous if solitary.

EMBRYOLOGY. The tapetum is 1-layered and

plasmodial. The ovary consists of a single carpel.
Fig. 69. Lemnaceae. Wolffiella denticulata. (Photo J. Bogner)
The ovule is at first atropous and bitegmic. In
X20 Spirodela and some Lemna species it later be-
comes amphitropous or anatropous. The outer
integument is 2-several-layered, the inner one is
formed. Generally, they appear in pouches instead 2-3-layered. The tegmen layer of the seed coat is
of the second daughter frond. In Wolffioideae the thin but hard and has an apical operculum. Both
flowers originate in a cavity on the upper surface integuments are persistent in ripe seeds. The nu-
of the frond, different from the pouch where the cellus is crassinucellate and in the mature seed
daughter fronds are formed. remains as a small nucellar cap. Embryo-sac for-
There is much controversy about the interpreta- mation is of the normal type in Spirodela (Brooks
tion of the floral structures. Some authors (e.g. 1940; Maheshwari and Maheshwari 1963), of
Hegelmaier 1868; Eichler 1875; Brooks 1940; the Scilla or Allium type in Lemna (Lawalree 1952;
Meuse! 1951; Maheshwari 1958a,b; Daubs 1965; Maheshwari 1956b, Maheshwari and Kapil
Den Hartog and van der Plas 1970; and Kandeler l963a,b) and in Wolffia (Gupta 1935; Maheshwari
1979) assume that Pistia (Araceae) is the closest 1954, 1956a). Embryo development follows the
relative of Lemnaceae and regard the flowering Asterad type. During maturation of the seed,
organs as a reduced inflorescence consisting of the apical cell develops into the cotyledon and
1-2 male flowers and 1 female flower, surrounded the basal cell develops into the hypocotyl with the
in the Lemnoideae by a spathe. In fact, the inter- suspensor and the primordium of the first frond.
pretation as an inflorescence depends totally on In the mature embryo, it is not possible to distin-
the presumed relationship to Araceae. Others guish sharply between cotyledon, hypocotyl and
(Hegelmaier 1871; Lawalree 1945; and Hillman suspensor, and this last organ has been variously
1961) interpret the organs as a flower with 1-2 interpreted. Endosperm formation is Cellular.
stamens and 1 pistil (and 1 prophyll in Lem-
noideae). The inflorescence theory is based on POLLEN MORPHOLOGY. Pollen grains are similar
the supposed derivation of Spirodela from the su- in all species of Lemnaceae. The mature, 3-celled
perficially similar Pistia, which has an inflores- pollen grains are (11) 15-26(52) ~-tm in diameter.
cence. The relationship of Lemnaceae is still They are ulcerate with a circular aperture. The
uncertain, the simpler concept of a flower is here exine is 0.6- 1 11m thick, spinulate with 0.4- 2.5 11m
preferred for the flowering structures of Lem- long spinulae, 1.2-2.5 11m long in the Lemnoideae
naceae. and 0.4-1.0 11m long in the Wolffioideae.
The scalelike leaf (prophyll) in Lemnoideae is a
1-layered membranous utricle either open only at KARYOLOGY. Urbanska-Worytkiewicz (1980)
the top or slit to the base. In this subfamily the 2 counted chromosome numbers of plants from
stamens develop successively, the distal one first. about 1500 localities (30 species). The Lemnaceae
In Spirodela and the Wolffioideae the dehiscent are a very difficult material, "the chromosomes of
 Lemnaceae 267

numerous species being exceedingly small and of- especially ducks, become covered with Lemnaceae
ten tending to stick together". The basic chromo- and transport them to neighbouring waters. Long-
some number for Lemnaceae is x = 10. A diploid distance dispersal is rare as Lemnacean fronds dry
number of 2n = 40 is most common, the euploid out quickly and die. According to Wolek {1981),
numbers 2n = 20, 30, 40, 60, 70, 80 and the aneu- Wolffia arrhiza can survive outside the water only
ploid numbers 2n = 36 and 2n = 42 have been 20-30 min and Spirodela or Lemna about 2 1/2 h
recorded. Similar results regarding the variation (21-26°C, 60-70% air humidity). However, they
were observed by Banerjee {1971) and Beppu and may survive longer under different conditions.
Takimoto {1981). Geber (in lit.) obtained some- Godziemba-Czyk (1970) observed living fronds of
what different results. He could recognise the W. arrhiza after 6h and Ridley (1930) of L. minor
heterochromatin by application of DNA-specific after 12-22h. If the fronds are covered by the
fluorochromes. He counted uniformly 2n = 36 feathers or the hairs of the animals, they may sur-
(5 clones) for Spirodela intermedia, 2n = 40 (5 vive outside the water even longer. Seeds of most
clones), 80 (1 clone) for S. polyrhiza, 2n = 46 (7 species are drought-resistant. They can be trans-
clones) for S. punctata. In the genus Lemna, he ported over distances of several hundred kilome-
observed mostly 2n = 42, 44 and 63. ters by birds. Anthropogenous dispersal together
with aquarium plants is often noted.
PoLLINATION. Pollination biology is not known Vegetative propagation is much more important
in detail. Fruiting is very rare in nature even when than reproduction by seeds. Under optimal condi-
flowering occurs. Species which fruit often (e.g. tions, the number of fronds may double within
L. aequinoctialis, L. perpusilla) are mostly self- 24 h, and one frond can easily develop into a popu-
pollinating, pollen grains being dropped from the lation of over a million fronds within a month.
anthers onto the stigma. Most other species are Seeds are essential only for a few species to survive
protogynous. Presumed pollinators range from dry periods. In some species turions are formed
snails to insects (Ludwig 1909) or wind- and rain under unfavourable conditions, which sink to the
pollination (Maheshwari 1954) or direct contact of bottom. They can stay there for several months
drifting fronds (Lawaln!e 1961) pushed together waiting for better conditions.
by wind or animals. The small number of sticky
pollen grains, and the sucrose nectar produced by PHYTOCHEMISTRY. The literature on phytochem-
the stigma indicate entomogamy (cf. also Blodgett istry of Lemnaceae is so extensive that even a sur-
1923). Indeed, one can observe insects crossing a vey is not possible within this treatment (see
flowering Lemna cover and carrying pollen grains Landolt and Kandeler 1987). The protein content
of Lemnaceae sticking to hairs on their lower side. of the fronds is high (up to 45% dry weight). Most
species have a characteristic pattern of flavonoids
FRUIT AND SEED. Each fruit has a dry pericarp and (McClure and Alston 1966). Proanthocyanidins
contains 1 seed {1-5 in Spirodela intermedia and are present in Spirodela and in some species of
Lemna gibba, 1-2 in S. polyrhiza, S. punctata Lemna. Spirodela and Lemna contain calcium
and L. disperma). The fruits of species with anatro- oxalate up to 4o/o dry weight (Bornkamm 1965),
pous or amphitropous ovules are mostly winged either as raphides and druses in Spirodela, or as
laterally towards the top, but not winged in species raphides only in Lemna.
with atropous ovules (except L. perpusilla). Decay-
ing causes the opening of the fruit and takes place SUBDIVISION AND RELATIONSHIPS WITHIN THE
in some species (e.g. L. aequinoctialis) just after FAMILY. Lemnaceae are clearly divided into two
ripening of the seed, but in most other species subfamilies comprising two genera each. The
much later (up to many weeks). The seeds are Lemnoideae with the genera Spirodela and Lemna
ovoid with a dark, pointed operculum. The chalaza are morphologically more differentiated than the
forms a dark point at the base. The endosperm fills Wolffioideae. They have roots, veins and 2 4-
up part of the ripe seed and, at least in Lemna, is sporangiate stamens per flower. A scalelike leaf
rich in starch (Maheshwari 1956b). (prophyll) is also present at the base of the flower
and at the base of the frond in Spirodela. There
DISPERSAL AND REPRODUCTIVE BIOLOGY. The is a trend to reduction, leading from Spirodela
fronds are very small and easily transported by through Lemna and Wolffiella to Wolffia. Wolffia
wind, water and animals. Mammals such as bea- and W olffiella have pigment cells, plastids in the
vers, rats, muskrats, racoons and wild boar, which guard cells of the stomata, straight walls of the
usually feed on duckweed, as well as water birds, epidermis cells, and flavonols in common with
268 Lemnaceae
Spirodela but not with Lemna. Den Hartog and
van der Plas (1970) have divided the genus Wolf-
fiella into three genera: Wolffiella, Wolffiopsis,
and Pseudowolffia. This separation is not justified
since two intermediate species have recently been
described (Landolt 1980). ·
AFFINITIES. The four genera of Lemnaceae share
many morphological, biochemical and ecological
characteristics and comprise a distinct and rela-
tively uniform family. The reduction of most parts
makes it difficult to estimate relationships to other
families (see above under Vegetative Morphology
and Inflorescence and Floral Structure). Accord-
ing to a widely held view, Lemnaceae are evolved
through Pistia (Araceae) and are members of
Arales. The superficial similarity between Pistia
and Spirodela is indeed striking. Spirodela re-
sembles a reduced Pistia (especially the seedling),
but their shared features are present in other fami-
lies of the monocotyledons. The interpretation of
the reproductive organs as flowers or inflores-
cences depends totally on the presumed relation-
ship. The absence of a basal cell in the embryo
and the absence of a micropylar cap (Maheshwari
1956b) within monocotyledons seem to be typical
of Lemnaceae and Araceae and cellular endo-
sperm is almost exclusive for Lemnaceae and
Araceae. But there are many differences between
Pistia and the Lemnaceae, e.g.: embryo sac of the
Allium type in Lemnaceae, of the Polygonum type
in Pistia; micropyle formed by the inner integu-
ment in Lemnaceae, by the outer integument
in Pistia; pollen grains ulcerate and spinose in
Lemnaceae, inaperturate and plicate in Pistia (but
many Araceae have a spiny exine!); apiose present
in Lemnaceae and absent in Pistia; pollen 3-
nucleate in Lemnaceae, 2-nucleate in Pistia.
Therefore Pistia is clearly not the link between
Araceae and Lemnaceae. Recent molecular work
shows the Lemnaceae rooted in subfam. Aroideae
s.l. of the Araceae (French et al. 1995).
DISTRIBUTION AND HABITATS. The Lemnaceae
are of world-wide distribution. They do not occur
in arctic and desert regions. Of all the continents,
America shows by far the biggest diversity. Three
genera are distributed throughout the world;
Wolffiella is restricted to America and Africa.
Spirodela, the genus representing the most primi-
tive organisation, has its centre of distribution inS
America. Except for Lemna, which has its highest
diversity in N America and E Asia, Lemnaceae
occur mainly in subtropical regions of the south-
ern hemisphere.
All members of the family float on or below the
surface of freshwater. Together with water ferns
(Azolla, Salvinia) and liverworts (Riccia and
Ricciocarpus), they cover the surface of stagnant
waters. They have a relatively high demand for
nutrients in the water. Therefore, they occur only
in meso- to eutrophic waters. Waters with less
than 0.003 mg/1 phosphorous content in the aver-
age of the vegetation period do not contain
Lemnaceae (Liiond 1983). To survive under
unfavourable conditions, some species form
turions (small compact starch-filled buds) which
sink to the bottom of the water.
PALAEOBOTANY. Limnobiophyllum is interpreted
as a link beween Araceae and Lemnaceae (Kvacek
1995); it includes free-floating stoloniferous plants
with 1 or 2 sessile suborbicular to reniform leaves
and simple and branched roots. Two species are
recognised: L. scuta tum from the latest Cretaceous
to Oligocene of western N America and
Palaeocene of E Asia, and L. expansum from the
Miocene of Europe. Recently, specimens of L.
scutatum with attached stamens and monoporate,
spinulate pollen have been described (Stockey
et al. 1997). Records of seeds similar to those of
Lemnaceae date back to the Oligocene and Mi-
ocene, e.g. Lemna tertiaria from Siberia (Dorofeev
1963). Nikitin (1976) describes fossil seeds as
Lemnospermum pistiforme from the Oligocene
and Miocene, which are similar to Spirodela and
Lemna, but larger than Lemna and have about the
size of Spirodela intermedia. Seeds described as
Lemnospermum minimum by Mai and Walther
(1978) probably do not belong to Lemnaceae or
Araceae. Fossil seeds of L. trisulca and L. gibba are
reported from Pliocene and Quaternary deposits
of Russia (Nikitin 1957).
PARASITES. The Lemnaceae have astonishingly
few fungal parasites. The fronds of Spirodela are
attacked by the smut fungus Tracya lemnae.
Spirodela polyrhiza, Lemna gibba, L. minor and
less so L. valdiviana, are susceptible to the infec-
tion of the oomycete Pythium myriotylum. L.
aequinoctialis and S. punctata never show disease
symptoms (Rejmankova et al. 1986). However,
more than 40 insect species and many other ani-
mals feed on duckweeds.
EcoNOMIC IMPORTANCE. Due to the high protein
content and the fast vegetative reproduction
Lemnaceae have some potential economic impor-
tance (cf. Hillman and Culley 1978; Culley et al.
1981). In SE Asia (Burma, Thailand, Laos) Wolffia
 Lemnaceae
globosa is eaten by man (Bhanthumnavin and Me
Garry 1971). In Israel Lemna gibba is estimated as
a vegetable. Lemnaceae are cultivated in many
parts of the world as food for ducks, chicken and
fish (Schulz 1962), and in some regions also for
cattle and pigs. In addition, Lemnaceae are used
for removing nutrients or heavy metals out of
waste water, also for manure, and as sensitive test
plants to demonstrate toxic substances in water.
KEY TO THE SUBFAMILIES AND GENERA
1. Roots present; fronds with veins; daughter fronds and
flowers produced in 2 lateral pouches at the base of the
frond; flowers surrounded by a membranous leaf (spathe
or prophyll), with 2 4-sporangiate stamens; crystal cells
present; seeds with longitudinal ribs (Lemnoideae)
- Roots lacking; fronds without veins; daughter fronds pro-
duced from a single terminal pouch or cavity; flowers pro-
duced in a cavity on the upper surface of the frond, without
a membranous leaf (prophyll); 1 single 2-sporangiate
stamen; crystal cells lacking; seeds nearly smooth
(W olffioideae)
2. Roots 2-21; veins usually 5-16; fronds surrounded at the
base by a small scale; oxalate druses present 1. Spirodela
- Root only 1 per frond; veins 1-5 (rarely 7 in L. gibba); base
of the frond without scale; oxalate druses lacking2. Lemna
3. Fronds fiat, with air spaces, daughter fronds produced in a
terminal fiat pouch; flower originating in a cavity at the side
of the median line of the upper surface 3. Wolffiella
- Fronds globose to ovoid boat-shaped, without air spaces;
daughter fronds produced in a terminal conical cavity of
the mother frond; flower originating on the upper side in a
cavity in or near the median line of the frond 4. Wolffia
I. Subfam. Lemnoideae Engler (1876).
1. Spirodela Schleiden Fig. 68 A-C
Spirodela Schleiden, Linnaea 13: 91 (1839).
Fronds with (3)5-16(21) veins, with several layers
of air spaces, with pigment cells, and with
raphides and druses, sourrounded at the base by a
bifid scale (prophyll) covering the place of attach-
ment of the roots; roots 2-21 per frond (rarely 1).
Pistil with 1 amphitropous or 2-5 anatropous
ovules; seeds 0.7-2.0 mm long, with 10-22longitu-
dinal ribs. Three spp., two widely distributed,
one in S America.
2. Lemna L. Fig. 68D-K
Lemna L., Sp. Pl. 1: 970 (1753).
Fronds with 1-5(7) veins and 1 to several layers of
air spaces and raphides; pigment cells lacking;
fronds not surrounded by a prophyll; 1 root per
frond; pistil with 1 amphitropous or atropous or
269
2-7 anatropous ovules; seeds 0.4-0.9 mm long,
with 8-60 longitudinal ribs. Thirteen spp., mostly
wide distribution.
II. Subfam. Wolffioideae Engler (1889).
3. Wolffiella Hegelm. Fig. 69
Wolffiella Hegelm., Bot. Jahrb. Syst. 21: 303 (1895).
Pseudowolffia den Hartog & van der Plas (1970).
Wolffiopsis den Hartog & van der Plas (1970).
Fronds thin, orbicular, oblong to linear or falcate;
terminal pouche triangular and flat; frond with 1
layer of air spaces; flowers originating in a cavity
2 (2 cavities in W. welwitschii) at the side of the
median line of the upper surface of the frond.
Pistil with 1 atropous ovule. Ten spp., four in
Africa, five in America, and one in Africa and
America.
3
4. Wolffia Horkel ex Schleiden Fig. 68M-P
Wolffia Horkel ex Schleiden, Beitr. Bot. 1: 233 (1844).
Fronds thick, globose, ovoid, cylindrical, conical,
boat-shaped convex; air spaces lacking; flowers
originating in a cavity on or near the median line
of the upper surface. Pistil with 1 atropous ovule.
Eleven spp., three of which on more than one
continent.
Selected Bibliography
For complete bibliography see Landolt (1986) and Landolt
and Kandeler ( 1987).
Arber, A. 1919. On the vegetative morphology of Pistia and the
Lemnaceae. Proc. R. Soc. 91: 96-103.
Banerjee, M. 1971. Chromosome studies in Lemnaceae. Rev.
Roum. Embryo!. Ser. Cytol. 8: 21-27.
Beppu, T., Takimoto, A. 1981. Geographical distribution and
cytological variation of Lemna paucicostata Hegelm. Jpn.
Bot. Mag. (Tokyo) 94: ll-20.
Bhanthumnavin, K., McGarry, G.M. 1971. Wolffia arrhiza as a
possible source of inexpensive protein. Nature 232: 495.
Blodgett, F.H. 1923. The embryo ofLemna. Am. J. Bot. 10:336-
342.
Bornkamm, R. 1965. Die Rolle des Oxalats im Stoffwechsel
hoherer griiner Pflanzen. Untersuchungen an Lemna minor
L. Flora 156: 139-171.
Brooks, J.S. 1940. The cytology and morphology of the
Lemnaceae. Unpubl. PhD Thesis. Ithaca, NY: Cornell
University.
Brunaud, A. 1974. Organisation de Ia pousse vegetative d'une
lemnacee: Spirodela polyrrhiza (L.) Schleid. C.R. Acad. Sci.
Paris D 278: 1183-1186.
Culley, D.D., Rejmankova, E., Kvet, J., Frye, J.B. 1981. Produc-
tion, chemical quality and use of duckweeds (Lemnaceae)
270 Lemnaceae
in aquaculture, waste management, and animal feeds.
J. World Marie. Soc. 12: 27-49.
Daubs, E.H. 1965. A monograph of Lemnaceae. Illinois Bioi.
Monogr. 34. Urbana: The Univ. Ill. Press pp. 1-118.
Den Hartog, C., van der Plas, F. 1970. A synopsis of the
Lemnaceae. Blumea 18: 355-368.
Dorofeev, P.I. 1963. The Tertiary floras of Western Siberia.
Moscow: Akad. Nauk.
Eichler, A.W. 1875. Bliithendiagramme, Vol. 1. Leipzig: W.
Engelmann, pp. 73-80.
Engler, A. 1877. Vergleichende Untersuchungen tiber die
morphologischen Verhaltnisse der Araceae. Nova Acta K.
Leop.-Carol. Dtsch. Acad. Naturf. 39: 135-224.
French, J.C., Chung, M.G., Hur, Y.K. 1995. Chloroplast DNA
phylogeny of the Ariflorae. In: Rudall, P.J., Cribb, P.J., Cut-
ler, D.F., Humphries, C.J. (eds.) Monocotyledons: systemat-
ics and evolution, Vol. 1. Royal Botanic Gardens, Kew, pp.
255-275.
Godziemba-Czyk, J. 1970. Vegetative and resting forms of
Wolffia arrhiza. 2. Anatomy, physical and physiological
properties. Acta Soc. Bot. Pol. 39: 421-443.
Gupta, B.L. 1935. Studies on the development of the pollen
grain and embryo sac of Wolffia arrhiza. Curr. Sci. 4: 104-
105.
Hegelmaier, F. 1868. Die Lemnaceen. Leipzig: W. Engelmann.
Hegelmaier, F. 1871. Ober die Fruktifikationstheile von
Spirodela. Bot. Zeitung 29: 645-666.
Hegelmaier, F. 1895. Systematische Obersicht der Lemnaceen.
Bot. Jahrb. 21: 268-305.
Hillman, W.S. 1961. The Lemnaceae or duckweeds. A review
of the descriptive and experimental literature. Bot. Rev. 27:
221-287.
Hillman, W.S., Culley, D.D. 1978. The uses of duckweeds. Am.
Sci. 66: 442-451.
Hoffmann, J.P. 1840. Beitrage zur naheren Kenntnis von
Lemna arrhiza nebst einigen Bemerkungen tiber L.
polyrrhiza, L. gibba, L. minor und L. trisulca. Wiegm. Arch.
Naturgesch. 6: 138-163.
Kandeler, R. 1979. Lemnaceae. In: Hegi, G. (ed.) Illustr. Fl.
Mitteleur. 2 ( 1) 3rd ed. Berlin: Parry: 335-346.
Kvacek, Z. 1995. Limnobiophyllum Krassilov - a fossil link
between the Araceae and the Lemnaceae. Aquat. Bot. 50:
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Zuer. 71: 566 pp.
Landolt, E., Kandeler, R. 1987. The family of Lemnaceae - a
monographic study. Vol. 2: Phytochemistry; physiology;
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Hochsch. Stift. Rubel Zuer. 95: 638 pp.
Lawaln!e, A. 1945. La position systematique des Lemnaceae et
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Lawalree, A. 1952. L'embryologie des Lemnaceae. Observa-
tions sur Lemna minor L. Cellule 54: 305-326.
Lawalree, A. 1961. La pollinisation de Lemna minor L. Nat.
Belg. 42: 164-165.
Ludwig F. 1909. Lemnaceae. In: Kirchner, 0., Loew, E.,
Schroeter, C. (eds.) Lebensgeschichte der Bliitenpflanzen
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Liiond, A. 1983. Das Wachstum von Wasserlinsen (Lem-
naceae) in Abhangigkeit des Nahrstoffangebots,
insbesondere Phosphor und Stickstoff. Veri:iff. Geobot. Inst.
Eidg. Tech. Hochsch. Stift. Rubel Zuer 80: 1-116.
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Nature 178: 925-926.
Maheshwari, S.C. 1956b. The endosperm and embryo of
Lemna and the systematic position of the Lemnaceae.
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Maheshwari, S.C. 1958a. The Lemnaceae. A contribution to
their biology, morphology and systematics. PhD Thesis,
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tween the aroids and the duckweeds. Nature 181: 1745-
1756.
Maheshwari, S.C., Kapil, R.N. 1963a. Morphological and em-
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914.
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McClure, J.W., Alston, R.E. 1966. A chemotaxonomic study of
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Ekol. Pol. 29: 405-419.
 Limnocharitaceae 271

Limnocharitaceae ultimately forms a new erect plant. The leaves are

spirodistichously arranged in basal rosettes. The
R.R. HAYNES, D.H. LES, bracts of floating inflorescences are often foliose,
and LB. HOLM-NIELSEN but smaller and generally less developed. The long
petiolate leaves have an open sheathing base and
a linear-lanceolate, ovate, or cordate blade (Fig.
70). The petioles are triangular or terete, with con-
spicuous diaphragms. No ligule or squamules are
present. There are few primary veins, these usually
Limnocharitaceae Takhtajan ex Cronquist, Integr. Syst. Class. converging, and a closed reticulum is formed by
Fl. Pl.: 1048 (1981). the secondary and tertiary veins.

Perennial latescent herbs, emergent, submersed, VEGETATIVE ANATOMY (based on Tomlinson

or floating in freshwaters. Roots fibrous, few to 1982). Roots are branched and have a thin-walled
many, from a stout rhizome or stolon. Stems epidermis with trichoblasts that may develop into
fleshy, erect, unbranched. Leaves basal or alter- root hairs. The outer cortex is compacted into an
nate, spirally inserted, petiolate; petioles terete to exodermis of 2 or 3 cell layers, and the remainder
triangular, mostly considerably longer than blade, of the cortex is composed of lacunose paren-
with sheathing base, the sheath without auricles, chyma. An endodermis with Casparian strips
intravaginal scales lacking; leaf blades orbicular to surrounds a single-layered pericycle. The stele is
lanceolate, the margins entire, the apex obtuse to composed of a single metaxylem vessel and few
round-acute, the base cordate to attenuate, vena- protoxylem tracheids, which alternate with single
tion reticulate, with parallel primary veins from sieve tubes. Secretory canals are absent.
base of blade to apex and transverse secondary The rhizome in Limnocharis has a persistent
veins. Inflorescence a scapose, erect to floating, epidermis underlayered by a suberized exodermis.
terminal, involucrate umbel, subtending spathe The well-developed cortex parenchyma is
lacking, with an involucre of few to several mem- lacunose and reticulate without diaphragms and
branous, acuminate bracts. Flowers hypogynous, often stellate. An endodermis similar to that in
perfect, pedicellate; perianth actinomorphic, of 6 the roots is present. The central xylem strands
separate tepals in 2 whorls, the outer 3 sepal-like, are irregular. In the stems of Limnocharis and
persistent, mostly erect and enclosing flower and Hydrocleys, the vascular system is supported by
fruit; the inner 3 petal-like, usually delicate, cadu- about 6 principal vascular bundles in the central
cous. Stamens free, 6-many; anthers 2-locular, parenchymatous core, these bundles without a
basifixed, 4-sporangiate, dehiscing by longitudi- fibrous sheath. Secretory canals are abundant.
nal slits. Carpels 3-many, separate or basally co- The petiole is angular in Limnocharis and terete
herent, 1-locular, each with numerous anatropous in Hydrocleys and Butomopsis. A hypodermis of
or campylotropous ovules; placentation laminar; 1-4 layers of chlorenchyma is present in
styluli short or absent, stigma linear. Fruits folli- Hydrocleys. The aerenchyma consists of a network
cular. Seeds numerous, glandular pubescent or of uniseriate parenchyma plates enclosing large
costate, U-shaped, without endosperm. lacunae traversed by diaphragms of sometimes
Three genera and eight species in the tropics of stellate cells. The core of the vascular system is a
the Old and New World. V-shaped arc of large central vascular bundles.
Secretory canals are abundant.
VEGETATIVE MORPHOLOGY. The roots arise from The floating leaves have hydropoten on the
the rhizome or the nodes of the stem. The abaxial epidermis. The epidermis is thin-walled,
nonchlorophyllous axes are short and erect or but the outer surface is weakly thickened and cuti-
elongated and creeping. The stems, including nized. Stomata are paracytic. The mesophyll is
stolons and scapes supporting floating inflores- dorsiventral with 1(-2) layers of peglike palisade
cences, and petioles have extensive aerenchyma. cells and a very lacunose, spongy parenchyma.
Branching is sympodial and continued from buds The abaxial hypodermis is continuous and com-
developed in the axil of the last leaf of the rosette. pact. Secretory canals are abundant in the leaves
The ultimate bud of the inflorescence of and form a continuous network. An apical pore is
Limnocharis flava always develops into a vegeta- formed abaxially by dissolution of mesophyll cells
tive bud (Wilder 1974), which forms a new at the endings of vascular elements of the major
vegetative shoot after the fruits have fallen, and veins.
272 Limnocharitaceae
Fig. 70A-H. Limnocharitaceae. A-F Limnocharis flava. A
Habit. B Flower. C Fruiting flower with numerous follicles. D
Stamen and staminode. E Follicle. F Seed. G, H Hydrocleys
nymphoides. G Flowering plant. H A carpel with two stamens
and two staminodes. (Takhtajan 1982)
INFLORESENCE STRUCTURE. The inflorescence
is umbel-like and composed of cymose units,
with bracts subtending individual flowers. The
inflorescences often proliferate, especially in
Hydrocleys, where 1 of the flowering axes elon-
gates and develops into another inflorescence
(Fig. 70G).
FLORAL STRUCTURE. The flowers are perfect and
are borne on pedicels longer than the bracts. The
perianth consists of 3 green sepals and 3 white to
bright yellow petals. The petals are twice as large
as the sepals to inconspicuous and smaller than
the sepals. The androecium consists of three to
numerous stamens. An outer circle of up to many
staminodia is found in some species of Hydrocleys
and Limnocharis (Fig. 70D,H). The anthers are
subbasifixed, extrorse or, more generally, latrorse,
as in Limnocharis. Anther dehiscence is by longi-
tudinal slits.
The gynoecium consists of 3 to about 20 carpels.
The carpels of Limnocharis are closely laterally
appressed and scarcely coherent at the base, while
those of Butomopsis and Hydrocleys are fused
below for up to 1/3 of their length. Stigmas are
sessile and capitate in Limnocharis; in Butomopsis
and Hydrocleys, they are horseshoe-shaped sur-
faces at the apex of the beaklike hollow apex of
the carpels.
Floral ontogeny in Hydrocleys reveals a centrifu-
gal stamen inception and a basically trimerous
developmental pattern, in spite of the consider-
able deviation from the monocotyledonous
ground plan of the mature flower (Sattler and
Singh 1973).
EMBRYOLOGY (Davis 1966). Anther wall forma-
tion follows the monocotyledonous type. The en-
dothecium develops fibrous thickenings, and the
middle layer is ephemeral. The tapetum is amoe-
boid and in the early stages consists of uninucleate
cells. Microsporogenesis is of the successive type
and the tetrads are tetrahedral or isobilateral,
rarely linear or T-shaped. The pollen grains are
3-celled at dispersal.
The placentation is laminar. A parietal cell is not
cut off from the primary archesporia! cell, but the
nucellar epidermis divides periclinally to form a
nucellar cap.
The embryo sac is of the Allium type. The en-
dosperm formation is Helobial. Free nuclear
divisions follow for some time at least in the mi-
cropylar chamber, and usually more abundantly
than in the chalaza! chamber. The formation of
the horseshoe-shaped embryo follows the Caryo-
phyllad type. The ripe seeds are without en-
dosperm or perisperm, and the embryo is
cylindrical, curved, linear, and lacks chlorophyll.
PoLLEN MoRPHOLOGY. The pollen grains are dis-
persed singly. They are globose, tri- to panto-
porate (Fig. 71) or inaperturate. In Hydrocleys and
Butomopsis the exine is sculptured by broad-
 Limnocharitaceae 273

DISPERSAL. The seeds are water dispersed, fur-

ther details being unknown. Additionally, bird
dispersal may take place as well.

PHYTOCHEMISTRY. There are no tannins, antho-

cyanins or anthocyanin pseudobases in Limno-
charitaceae. Flavonoids are not reported for the
family.

RELATIONSHIPS WITHIN THE FAMILY. Based on

number of stamens, thickness and ornamentation
of the testa, Pichon (1946) placed Limnocharis
and Hydrocleys in subtribe Limnocharitinae and
Fig. 71. Limnocharitaceae. Hydrocley martii, pantoporate Butomopsis ( Tenagocharis) in subtribe Tenago-
pollen, SEM X 1750. (Palynological Laboratory, Stockholm) charitinae. In contrast, Hooper and Symoens
(1982) stated that Hydrocleys and Butomopsis
show closer similarities to each other than to
Limnocharis, and that even the generic distinct-
based spines of varying densities. The spines are ness of Butomopsis is questionable.
sometimes interspersed with short columnar pro-
cesses. In Limnocharis the exine is granular and AFFINITIES. The relationship of Limnocharita-
pitted with small cavities (Argue 1973; Harley ceae to Alismataceae and Butomaceae has long
1982). been recognized, but authors have differed con-
siderably in their opinion regarding the specific
KARYOLOGY. The chromosome morphology of placement of the family. The shared presence of
Butomopsis and Hydrocleys are strikingly similar, features such as follicular fruits and laminar pla-
according to Rao (1953). He found, however, that centation motivated Hutchinson (1959) to com-
Butomopsis has the base number x = 7 and bine the Limnocharitaceae with Butomus in an
Hydrocleys x = 8. Kenton (1981) observed 2n = 16 enlarged family Butomaceae. The generic name
for H. nymphoides and 2n = 14 for H. modesta. Butomopsis (Limnocharitaceae) further alludes
Kenton went on to suggest that different numbers to the superficial similarity of these families. How-
within the genus could have arisen either by fu- ever, Cronquist (1981) observed many features
sion of 2 chromosomes (x = 8 to x = 7) or by that separate Limnocharitaceae (lactiferous ducts,
fission of 1 chromosome (x = 7 to x = 8). Rao leaves with petioles and expanded blades, cadu-
(1953) determined Limnocharis to have 2n = 20. cous petals, campylotropous ovules, seeds with
He proposed that Butomopsis, Hydrocleys, and curved embryos) from Butomaceae, which have
Limnocharis form a progressive aneuploid series. contrasting characteristics.
Other treatments have merged Butomopsis,
POLLINATION. Pollination in the family is in- Hydrocleys, and Limnocharis with the Alisma-
completely known. However, we have observed taceae, leaving the Butomaceae as monotypic
Limnocharis and Hydrocleys species frequently (Pichon 1946; Dahlgren et al. 1985). The Limno-
visited by flies. charitaceae differ from the Alismataceae by nu-
merous ovules per carpel, laminar placentation,
FRUIT AND SEED. The carpels develop into fol- and dehiscent fruits. The Alismataceae have 1-few
licles opening towards the center of the flower. ovules per carpel with basal placentation and in-
The seeds are horseshoe-shaped and sculptured dehiscent fruits. Cronquist (1981) recognized all
(Fig. 70F). The surface of the testa in Limnocharis three families as separate.
consists of strongly elongated cells upturned at the Phylogenetic analyses using both morphological
ends to produce prominent transverse ridges or data (Haynes and Holm-Nielsen 1992) and mo-
flanges that cross the seed at regular intervals. The lecular data (Les and Haynes 1995; D.H. Les,
testa surface of Butomopsis and Hydrocleys con- unpubl.) resolve a clade with Butomus as basal to
sists of relatively short cells and a smooth surface genera of both Alismataceae and Limnochari-
interrupted by small papillae in Butomopsis and taceae. Although these analyses show Limno-
by hairlike projections with globular ends in charitaceae to be monophyletic, molecular data
Hydrocleys. depict the family as derived from a paraphyletic
274 Limnocharitaceae
Alismataceae (D.H. Les, unpubl.). However, the
paraphyly of Alismataceae remains uncertain
because trees only 3 steps longer resolve both
Alismataceae and Limnocharitaceae as mono-
phyletic families (see discussion for Alis-
mataceae). Given the questionable paraphyly of
Alismataceae, and the several defining character-
istics of the Limnocharitaceae, we have retained
these families as distinct taxonomically.
DISTRIBUTION AND HABITATS. The Limno-
charitaceae are pantropical, Limnocharis and
Hydrocleys occur naturally from Argentina to
Mexico and the Caribbean. Butomopsis occurs in
Africa, Asia, and northern Australia. Hydrocleys
and Limnocharis are naturalized in the USA and in
the Paleotropis.
All species of the Limnocharitaceae occur in
aquatic habitats, especially those with stagnant
or nearly stagnant freshwater. Limnocharis and
Butomopsis grow in muddy substrates as
amphiphytes or rosette-helophytes. Hydrocleys is
a typical hydrophyte with floating leaves and more
or less floating inflorescences. Although Hydro-
cleys has been reported as free-floating it is mostly
rooted in muddy substrates or, if floating, then a
member of floating island associations.
EcoNOMIC IMPORTANCE. Members of the family
are often troublesome weeds in the tropics, par-
ticularly in paddy fields, although not as serious
as Eichhornia. Limnocharis flava is cultivated in
ponds in India and SE Asia, where the leaves are
used as a vegetable and as pig fodder. Limnocharis
and Hydrocleys are now receiving much attention
in the aquarium and ornamental plant trade.
KEY TO THE GENERA
1. Carpels 15 or more in number, semicircular; stigmas sessile
1. Limnocharis
- Carpels 3-9, linear-lanceolate; styles present
2. Leaf blades about as broad as long; petioles terete; inflores-
cences decumbent or floating; Neotropical 3. Hydrocleys
- Leaf blades much longer than broad; petioles triangular;
inflorescences erect; Paleotropical 2. Butomopsis
Genera of Limnocharitaceae
1. Limnocharis Humb. & Bonpl. Fig. 70A-F
Limnocharis Humb. & Bonpl., Pl. Eequinoct. 1: 116 (1808).
Plants emersed. Stems short, rhizomatous, occa-
sionally with stolons; leaves basal, emersed,
petiole triangular, aseptate, blade lanceolate to
oval. Inflorescences up to 10, on aseptate scapes,
occasionally proliferating; the pedicels somewhat
dilated, often winged, inflated trigonous; petals
yellow, ovate to suborbicular, longer than the
sepals; stamens many, the outer often sterile,
filaments flattened; carpels 15-20, laterally com-
pressed, scarcely coherent at base. Follicles semi-
circular, membranous, dorsally furrowed; seeds
transversely multicostate. Two spp., neotropics.
2. Butomopsis Kunth
Butomopsis Kunth, Enum. Pl. 3: 164 (1841).
Tenagocharis Hochst. (1841).
Plants partly submersed, stems terete; leaves
basal, erect, petiole triangular, oblong-lanceolate,
the base cuneate. Inflorescences 1-3, on a long
septate scape, 1- to 2-whorled with cymes of
usually 3-11 flowers; pedicels angular; petals
white, delicate, oblong-obovate, orbicular, slightly
smaller than sepals; stamens 8-12, in 3 whorls;
filaments filiform; carpels 4-9, basally coherent,
tapering into a style, this curved outward. Follicles
dehiscing along the inner margins. Seeds with
testa ornamented with small papillae. Only one
sp., B. latifolia (D. Don) Kunth, tropical Africa, S
and SE Asia and N Australia.
3. Hydrocleys Rich. Fig. 70G,H, 71
Hydrocleys Rich., Mem. Mus. Nat. Hist. 1: 368 (1815); Haynes
& Holm-Nielsen, Fl. Neotr. 56: 1-34 (1992), rev.
Plants submersed; stems short, stolons often
present; leaves basal, either floating and petiolate,
or submersed phyllodia; petioles terete, with a
sheathing base. Inflorescences few to many on
long septate scape, comprising few to numerous
flowers, occasionally proliferating; pedicels terete;
petals yellow to white, delicate, longer than or
2
shorter than the sepals; stamens 6-many, in !-
several series, the outer often sterile, the filaments
linear or lanceolate; carpels 3-8, basally cohering,
attenuate into the style, this curved inward. Fol-
licles dehiscing along the inner margins. Seeds
sparsely to densely glandular pubescent. Five spp.,
tropical America.
Selected Bibliography
Argue, C.L. 1973. The pollen of Limnocharis jlava Buch.,
Hydrocleis nymphoides (Willd.) Buch., and Tenagocharis
latifolia (Don) Buch. (Limnocharitaceae). Grana 13: 108-
112.
 Limnocharitaceae/Lowiaceae
Cronquist, A. 1981. See general references.
Dahlgren, R.T.M. eta!. 1985. See general references.
Davis, G.L. 1966. See general references.
Harley, M.M. 1982. Palynological evidence of a close associa-
tion between Butomopsis Kunth and Hydrocleys L.C. Rich.
(Limnocharitaceae). In: Symoens, J.J., Hooper, S.S. (eds.)
Studies on aquatic vascular plants. Brussels: Royal Botani-
cal Society of Belgium, pp. 61-65.
Haynes, R.R., Holm-Nielsen, L.B. 1985 [1987]. A generic treat-
ment of Alismatidae in the neotropics with special reference
to Brazil. Acta Amazonica 15: 153-193.
Haynes, R.R., Holm-Nielsen, L.B. 1992. The Limnocharitaceae.
Flora Neotropica 56: 1-34.
Hooper, S.S., Symoens, J.J. 1982. Observations on the family
Limnocharitaceae Takhtajan ex Hooper & Symoens. In:
Symoens, J.J., Hooper, S.S. (eds.) Studies on aquatic vascu-
lar plants. Brussels: Royal Botanical Society of Belgium,
pp. 56-60.
Hutchinson, J. 1959. The families of flowering plants. 2 vols.
2nd edn. Oxford: Clarendon Press.
Kenton, A. 1981. A Robertsonian relationship in the chromo-
somes of two species of Hydrocleys (Butomaceae sens. lat.).
Kew Bull. 36: 487-492.
Les, D.H., Haynes, R.R. 1995. Systematics of subclass
Alismatidae: a synthesis of approaches. In: Rudall, P.J.,
Cribb, P.J., Cutler, D.F., Humphries, C.J. (eds.) Monocotyle-
dons: systematics and evolution. Royal Botanic Gardens,
Kew pp. 353-377.
Pichon, M. 1946. Surles alismatacees et les butomacees. Not.
Syst. (Paris) 12: 170-183.
Rao, Y.S. 1953. Karyo-systematic studies in Helobiales. Proc.
Nat!. Int. Sci. India 19: 563-581.
Sattler, R., Singh, V. 1973. Floral development in Hydrocleys
nymphoides. Can. J. Bot. 51: 2455-2458.
Takhtajan, A.L. 1954. Proiskhozhdenie pokrytosemennykh
rastenii. Moskva: Nauka.
Takhtajan, A.L. 1982. See general references.
Tomlinson, P.B. 1982. See general references.
Wagner, R. 1918. Ober den Aufbau der Limnocharis laforestii
Duchass. Sber. Akad. Wiss. Wien, Math.-Nat. Kl. I, 127:
317-327.
Wilder, G.J. 1974. Symmetry and development of Butomus
umbellatus (Butomaceae) and Limnocharis flava (Limno-
charitaceae). Am. J. Bot. 61: 379-394.
275
Lowiaceae
K. LARSEN
Lowiaceae Ridley, Fl. Malay Penins. 4: 291 (1924), nom. cons.
Perennial, glabrous herbs without aerial stem, of-
ten tufted from a horizontal or vertical, sympodial
rhizome bearing scalelike leaves. Leaves disti-
chous, glabrous, eligulate, base sheathing, petiole
well defined, lamina entire, lanceolate, dorsiven-
tral, rolled up from one side to the other in bud;
venation pinnate with parallel secondary veins
emerging from the prominent main vein at nar-
rowly acute angles and regularly connected by
fine, transversal veins (conspicuous in dried
material, but not so in living plants). Inflores-
cences dense or lax, consisting of sympodially con-
nected paracladia each producing a single flower.
Flowers malodourous, bracteate, bisexual, epigy-
nous, with the ovary prolonged into a solid exten-
sion. Sepals (outer tepals) 3, subequal, narrow,
fused basally into a tube. Petals (inner tepals) 3,
very unequal, the 2 lateral small, narrow, the me-
dian one large, forming a variously shaped label-
lum. Stamens 5, free from each other, the adaxial
median reduced; filaments short, adnate to base of
petals; anthers tetrasporangiate, introrse, opening
by long slits, apically terminated by a short con-
nective tip. Gynoecium of 3 fused carpels with sep-
tal nectaries; placentation axile; style elongate,
widening apically; stigmatic lobes 3, large, undi-
vided or flattened, laciniate to fimbriate, wet.
Ovules numerous, anatropous, bitegmic, crassi-
nucellate. Fruit a dry, elongate, loculicudal cap-
sule terminated by a beak formed by the proximal
part of the ovary extension. Seeds numerous, glo-
bose or pyriform, hairy, albuminous, with a lacer-
ate aril and an operculum in front of the radicle.
A monogeneric family with ca. ten species in SE
Asia.
VEGETATIVE ANATOMY. The anatomical obser-
vations available refer to two species (Tomlinson
1969). The roots are relatively thick and air chan-
nels are present in the inner cortex. Vessels with
scalariform perforation are present in the metaxy-
lem of the root, the perforation plates in the ob-
lique end walls have numerous bars. In the outer
cortex of the rhizome fibrous strands are found in
276 Lowiaceae

various species; as an inner limit of the cortex a term ovary extension for this structure was intro-
distinct endodermoid layer is found. The leaves duced by Holttum (1970). The sepals (or outer
are totally glabrous. Few stomata are found on the tepals) are equal; the petals (or inner tepals) are
upper surface, but uniformly and diffusely scat- arranged as in an orchid flower with 2laterallobes
tered on the lower. Isolated fibres and fibrous and a labellum (Fig. 73). The lateral lobes are small
strands are frequent near the epidermis. The chlo- and overlap the stamens; the labellum is variously
renchyma consists of an irregular mixture of small shaped, in some species having a narrower proxi-
and larger cells without a differentiated palisade mal part and a broad distal part, in other species
layer. Air channels are present in the leaf midrib.
Calcium oxalate is common in all parts of the
plants both as crystals and raphides; crystal sand
is abundant in the rhizome. Silica occurs as "hat-
shaped" bodies in stegmata cells along the vascu-
lar bundles in all organs except the roots. Tannin
is common only in the rhizomes. The rhizomes are
also rich in small, angular starch grains, quite dif-
ferent from those found in the Musaceae and the
Strelitziaceae.

INFLORESCENCE AND FLOWER STRUCTURE. The

inflorescences are terminal on short leafy
branches of the stem; they persist for a long time
(Fig. 72). The partial inflorescences are composed
of sympodially connected paracladia, each of
which consists of 5 nodes with 5 bracts in trans-
versal-distichous order. The 2-keeled prophyll is
sterile, the following 2 bracts, called interphylls,
bear buds that eventually may develop into new
branches of the cymoid. The 4th bract subtends
the only flower of the paracladium, and the last
bract is reduced and envelops the open end of the
paracladium axis (Kunze 1985). The single flower
is to be interpreted as the reduced coflorescence.
The ovary has an apical prolongation which is
sometimes called a calyx tube or hypanthium. It is,
however, solid with a central vascular strand con-
necting the base of the style with the ovary; the

Fig. 73A-E. Lowiaceae A-C Orchidantha longiflora. A Habit.

B Androecium and stigma, abaxial view. C Style and stigmatic
Fig. 72. Lowiaceae. Orchidantha borneensis, flowering plant. lobes. D, E 0. maxillarioides. D Habit. E Arillate seed.
Sarawak. (Photo J. Bogner) (Takhtajan 1982)
 Lowiaceae
such as 0. holttumii the whole labellum is rather
broad but distally curved back and forming a
clawlike structure. The labellum often has lamel-
lae in the central part along the vascular strands.
These structures may play a role in pollination.
The 5 stamens are of a simple nature, opening
introrsely, in a close semicircle around the style
and facing the labellum; the position of the miss-
ing stamen is median posterior, as in Musa.
POLLEN MoRPHOLOGY. The pollen is inaper-
turate, psilate, and spheroidal and resembles
much that of Ravena/a (Strelitziaceae).
KARYOLOGY. In three species, 0. holttumii, 0.
maxillarioides and 0. siamensis, the chromosome
number has been determined as 2n = 18 (K.
Larsen 1993, and unpubl.).
PoLLINATION. Flowers of 0. fimbriata, 0. sia-
mensis, and 0. holttumii have a foul smell. The
first two species have flowers which are placed
near the soil and have a large, white labellum. The
succulent central part along the midvein of the
labellum consists of nutritious tissue with protein
cells. W.J. Kress (pers. comm.) observed that tree
shrews feed on the labellum in nighttime. 0.
holttumii has an erect labellum that is dark purple
during an thesis and may attract flies of beetles.
SEED STRUCTURE. The seeds are spheroidal or
pyriform; the testa is smooth ( 0. holtthumii) or
covered with hairs (0. chinensis ); the aril consists
of 3-4 coarse hair-like outgrowths; the raphe is
inconspicuous; endosperm is copious, perisperm
very little, mostly only one cell layer thick and at
the chalazal part several cell layers thick; the em-
bryo is cylindric (Wen et al. 1997; H.-J. Tillich,
person. comm.).
PHYTOCHEMISTRY. Unknown.
AFFINITIES. It is undisputed that the Lowiaceae
belong in the "banana group" of the Zingiberales.
Since the description of the first species of
Orchidantha by N.E. Brown in 1886, the genus has
been referred to the Musaceae, until a separate
family close to the Musaceae was proposed for it
by Ridley in 1924. This viewpoint has repeatedly
received support (e.g. Kunze 1986; Kress 1990),
even if in several respects the Lowiaceae have
an isolated position. Unfortunately, the seed
anatomy is only superficially known; a better
knowledge might help to a deeper understanding
of their affinities.
277
EcoNOMIC IMPORTANCE. The species are of no
economic importance. Only one sp., 0. maxil-
larioides, widely grown in public greenhouses in
Europe.
DISTRIBUTION AND HABITATS. The genus is dis-
tributed from southern China (Kwangtung pro-
vince and Hainan) through Vietnam, Laos and
Thailand to W Malaysia, Indonesia (Borneo) and
Brunei. Frequent quotations in recent literature to
"Pacific Islands" are due to an error. The plants
occur in lowland evergreen forests along streams
and waterfalls, sometimes also in clearings after
logging.
Only one genus:
Orchidantha N.E. Brown Fig. 72, 73
Orchidantha N.E. Brown, Gard. Chron. II, 26: 519 (1886).
Holltum {1970) included in his revision six spp.,
but to date ten spp. have become known.
Selected Bibliography
Holttum, R.E. 1970. The genus Orchidantha (Lowiaceae).
Gard. Bull. (Singapore) 25: 239-246.
Kress, W.J. 1990. The phylogeny and classification of the
Zingiberales. Ann. Mo. Bot. Gard. 77: 698-721.
Kunze, H. 1986. Infiorescenz- und Bliitenmorphologie von
Orchidantha maxillarioides (Rid!.) K. Schum. (Lowiaceae).
Beitr. Bioi. Pfianz. 61: 221-234.
Larsen, K. 1961. New species of Veratrum and Orchidantha
from Thailand and Laos. Bot. Tidsskr. 56: 345-350.
Larsen, K. 1973. A new species of Orchidantha (Lowiaceae)
from Vietnam. Adansonia II, 13: 481-482.
Larsen, K. 1993. A new species of Orchidantha (Lowiaceae)
from Borneo. Nord. J. Bot. 13: 285-288.
Larsen, K. 1996. Lowiaceae. Fl. Males. II, 12: 763-774.
Loesener, Th. 1930. Musaceae. Unterfam. III. Lowioideae. In:
Engler, A., Prantl, K. (eds.) Natiirliche Pfianzenfamilien,
2nd edn., 15a: 541. Leipzig: W. Engelmann.
Long, H., Wen, Y. 1997. Pollen morphology ofLowiaceae from
China. J. Trop. Subtrop. Bot. 5(3): 6-9.
Schumann, K. 1900. Musaceae. Subfam. III. Lowioideae. In:
Engler, A. (ed.) Pfianzenreich IV. 45: 40-42. Leipzig: W.
Engelmann.
Takhtajan, A.L. 1982. See general references.
Tomlinson, P.B. 1969. Commelinales-Zingiberales. In:
Metcalfe, C.R. (ed.) Anatomy of the monocotyledons, Vol.
III. Oxford: Clarendon Press.
Wen, Y., Liao, J., Wu, Q. 1997. Anatomy and histochemistry of
the seed of Orchidantha sinensis T.L. Wu. Guihaia 17: 235-
241.
Wu, T.L. 1964. Lowiaceae, a family new to China. Acta
Phytotaxon. Sin. 9: 335-343, 2 plates.
278
Marantaceae
L. ANDERSSON
Marantaceae Petersen in Engler & Prantl, Nat. Pflanzenfam. II.
6: 33 (1888), nom. cons.
Terrestrial rhizomatous herbs, sometimes liane-
scent and high-climbing, very variable in devel-
opment of aerial shoots. Leaves distichous,
segregated into sheath, petiole proper (often miss-
ing), pulvinus and blade; blade with a strong mid-
rib and thin, parallel and closely set lateral veins,
these ± sigmoid, fusing marginally, intercon-
nected by closely set, parallel transverse veinlets.
Inflorescence terminal or lateral, simple or a ±
complex synflorescence; inflorescence unit (flor-
escence) a usually spiciform or capitate thyrse
with distichous or spiral spathes ("bracts") sub-
tending ± elaborate flower clusters composed of
1-several (rarely numerous) 2-flowered cymules
(1-flowered in Monotagma and Monophrynium),
each with a prophyll on the dorsal side at base,
sometimes also with a scale-like interphyll on
the ventral side and 1 or 2 ± dorsal bracteoles
(additional "lateral bracteoles" in one species
of Calathea). Flowers perfect, epigynous, penta-
cyclic, heterochlamydeous, completely asymmet-
ric (the 2 flowers of a pair being mirror images);
sepals distinct; petals, androecial elements, and
style fused to form a floral tube ("corolla tube")
greatly variable in length. Outer androecial whorl
rarely missing, usually of 1 or 2 staminodes, which
are usually petaloid and showy, but sometimes
acicular, or ± rudimentary; inner androecial
whorl of 3 members, 1 fertile monothecic stamen
(often with a petaloid appendage), 1 hood-shaped
staminode (staminodium cucullatum), and 1 con-
spicuously firm and fleshy staminode (stamino-
dium callosum); the inner staminodes basally
fused to form a firm and fleshy staminal tube
which ± exceeds the level where the corolla lobes
separate from the floral tube. Ovary 3-carpellate
and 3-locular, but 2locules often empty and com-
pressed; fertile locule(s) 1-ovulate with the anatro-
pous ovule inserted at base; septa with completely
sunken septal nectaries. Style fused basally with
the floral tube, in mature untriggered flowers the
style enclosed in the hood of the cucullate
staminode, bent backwards and held under ten-
Marantaceae
sion; when released by a pollinator, snapping
forward and curling up in a circinate or (Thalia)
helical fashion; stigmatic surface restricted to the
inner side of a funnel-shaped depression in the
apex of the style. Fruit usually a loculicidal cap-
sule, more rarely berry-like (Sarcophrynium,
Thaumatococcus), or caryopsislike (Thalia). Seeds
rather large (3-20 mm), without endosperm at
maturity, embryo horseshoe-shaped, embedded
in copious starchy perisperm; perisperm with
a simple or forked, central canal running from
base to the level of the embryo bend, or slightly
further.
A family of 31 genera and ca. 550 species, essen-
tially pantropical, but absent from Australia, with
a few members outside the tropics in S and N
America.
VEGETATIVE MORPHOLOGY. The basic contribu-
tion in this field is still that of Eichler (1884); some
particulars are proffered by Andersson (1977,
1981), Holttum (1951), Kennedy (1978) and
Tomlinson (1961).
The general organisation is that of a sympodial
rhizome with branches terminating in aerial
shoots and renewal branches appearing near the
base of the aerial shoots. Shoots are mostly mono-
morphic but dimorphic shoots (leafy shoots and
inflorescence shoots) are not uncommon. Roots
are usually borne on the rhizome only, but in
numerous species roots are ± regularly formed
at the nodes of aerial stems.
The roots are usually fibrous, but in many spe-
cies of Calathea, in Saranthe leptostachya, and in
Monotagma sp., terminal root tubers have been
recorded (Mats Hagberg, pers. comm.; Tomlinson
1961, pers. observ.).
The shape of the rhizome is quite variable and
its structure is often obviously correlated with
phenology. In perennating species, rhizome
branches are commonly moderately elongated
with thin scalelike leaves, which may be rather
ephemeral, and of a firm, often ± woody texture.
Stoloniferous rhizomes are found in some species
of Maranta (Andersson 1986) and Marantochloa
(Tomlinson 1961). In highly seasonal species,
where the aerial shoots die in the dry season,
rhizomes have specialised, ± fleshy and starch-
storing branches (e.g. Maranta arundinacea).
In Monophyllanthe oligophylla, the rhizome axis
is thin, the rhizomes tending to be stoloniferous,
but the terminal scale leaves are thickened and
starch -storing.
The Marantaceae are remarkable for the great
organisational diversity of the aerial shoot system.
 Marantaceae
All shoot systems are made up, however, of unit
branches with a uniform sequence of parts. The
1st and 2nd internodes are very short. The 1st
node is provided with a 2-keeled prophyll backing
the mother axis, the 2nd with a bladeless sheath,
the interphyll (mesophyll of some authors, rarely
missing). Subsequent internodes are short or ±
elongate and the phyllomes are either sheaths
or "normal" leaves. "Rhythmic contractions"
("several-leaved nodes") are characteristic of the
major axes of some genera or parts thereof (e.g.
Stromanthe, Ischnosiphon). Phyllotaxis is uni-
formly distichous in vegetative shoots. In lateral
shoots, the plane of distichy of the prophyll and
interphyll is paralell to that of the mother axis
(prophyll always backing the mother axis), but is
then often turned 90°.
The simplest growth form is the "rosulate"
model, characteristic of most low-growing spe-
cies: a basal leaf cluster is produced from a short
axis at ground level and the axis is eventually ter-
minated by an inflorescence. In the rare "cannoid"
model, the unbranched axis is elongated, carrying
± evenly distributed leaves separated by distinct
internodes, the axis being closely wrapped up by
the leaf sheaths. In some genera more elaborate
models are found, without leaves at ground level
and with the first distinct aerial internode much
elongated (up to several m). On top of this intern-
ode the leaves are often clustered in a fanlike fash-
ion (Ischnosiphon spp., Stromanthe spp., with
leaves on the main shoot; Donax with leaves on
lateral shoots). In some species of Maranta
subgen. Maranta and Stromanthe, the shoot is ex-
cessively branched to form a crown of leafy
branches on top of the long basal internode
("shrub-shaped" model of Andersson 1977). In
the "bambusoid" model of Ischnosiphon sections
Bambusastrum and Rotundifolii, the major
branches have a long sequence of elongated inter-
nodes, the ultimate branchings being scandent.
Assimilating leaves and inflorescences are borne
in ± brachyblastic clusters laterally on these ma-
jor axes. Also in Hypselodelphys, Trachyphrynium
and Haumania, scandent shoots are found, but
the details of branching have not been described.
A complete marantaceous leaf is segregated into
4 distinct zones: (1) sheath, (2) petiole proper, (3)
pulvinus ("callus", "superior part of petiole") and
(4) blade. The petiole proper is missing in many
species, and in species with elaborate aerial
shoots, it shows an evident decrease in develop-
ment from basal to distal leaves. All divisions of
the mature leaf develop from the Unterblatt and
the Oberblatt remains small, identifiable in young
279
leaves as a small precursory tip, and disintegrates
(Kaplan 1973). Leaf development was described
by Tran (1974).
The leaf blade has a prominent midrib from
which closely set parallel lateral veins diverge at an
acute angle (longitudinal veins of Tomlinson
1961). The lateral veins have a ± pronouncedly
sigmoid course and fuse near the blade margin.
Some of the lateral veins, ± equidistant, are
slightly thicker than the others and have some-
times, for descriptive purposes, been termed
major lateral veins. The lateral veins are intercon-
nected by closely set, ± equidistant veinlets
("transverse veins"), which are perpendicular to
the lateral ones.
In bud, 1/2 of the leaf blade is rolled up around
the other half (supervolute ptyxis), and this side is
always narrower than the other in the mature leaf.
Two symmetry forms may thus exist. The term
homotropic leaves has been used for the case
when all leaves of a shoot are of the same symme-
try form, and the term antitropic leaves for the
case where successive leaves on a shoot are of
different symmetry form.
VEGETATIVE ANATOMY. Root stele polyarc. Hairs
common, unicellular. Epidermis cells with anti-
clinal walls sinuous in surface view. Stomata
paracytic or tetracytic according to Tomlinson
(1974); according to Olatunji (1980), mainly
paracytic, but also tricytic, tetracytic and poly-
cytic, the development being mainly biperi-
geneous. Hypodermis 1-layered below each
surface. Silica cells common in all parts except
roots; cells either thin-walled ("false stegmata") or
irregularly thick-walled ("true stegmata"); silica
bodies mostly "hat-shaped" or druselike. Rhom-
bohedral or rodlike calcium oxalate crystals are
common in the ground tissue of many species, but
raphides are unknown. Tannin common in the
ground tissue, both in large specialised cells and
in unmodified ones. Secretory cells present in the
adaxial hypodermis and ground tissue of the leaf
blade in Hypselodelphys. Mucilage canals present
in Thalia. Aerial stem without clear demarcation
between cortex and central cylinder; stems of spe-
cies with a well-developed aerial shoot system
usually have a distinct peripheral mechanical zone
of numerous fibre bundles, which are sometimes
confluent into a fibrous sheath. Leaf axis with
schizogeneous air lacunae arranged in a single
abaxial arc; an additional lacuna present distally
in sheath and petiole of Thalia and Halopegia;
Haumania has a single wide lacuna in the centre
of the sheath. Vascular bundles arranged in three
280 Marantaceae

main systems: (1) main arc of large bundles alter-

nating with, and adaxial to air lacunae; (2) abax-
ial system of small irregularly arranged bundles,
showing gradual transition from fibre to vascular
bundle in adaxial direction; (3) adaxial incon-
spicuous system, sometimes partly absent. Pulvi-
nus with epidermis of ± isodiametric cells with
non-sinuous walls; inside the chlorenchyma with
a continuous palisadelike layer of colourless cells
at oblique angle to the longitudinal axis, indi- \·::··.
vidual cells often more than ten times as long as
wide, 2-3-nucleate (Tran 1974). Leaf blade with
lateral (longitudinal) veins sheathed on both sides
\'
I

\.··.
with fibres, sheaths connecting with hypodermis
on both sides; transverse veins similar to lateral
ones but much narrower and not connecting with
hypodermis. Colour patterns of leaf blades are
achieved in several ways: (1) variation in size and
number of chloroplasts (up to complete absence
in certain areas); (2) presence of anthocyanin in
the epidermis; (3) predominance of xanthophyll
in the assimilating tissue of certain areas; (4)
variation in size and distribution of air-filled
intercellular spaces.
Non-structural nectaries are present at the
junction of petiole and sheath in some species
of Ctenanthe and Stromanthe (Kirchoff and
Kennedy 1985).

INFLORESCENCE STRUCTURE. According to

Andersson's (1976) interpretation, the inflores-
cence unit is a ± spiciform or capitate thyrse. Such
units, florescences, are often complexed into
synflorescences, which may be highly distinct Fig. 74A-E. Marantaceae. A, B Pleiostachya pruinosa. A
Synftorescence. B Flower. C, D Monotagma secundum. C
superunits (e.g. Monotagma, Fig. 74C; Phrynium), Synflorescence. D Seed. E M. laxum, flower. (Kennedy et al.
or ± diffuse (e.g. Maranta, Marantochloa), up to 1988)
the point where the concept is quite pointless.
Simple inflorescences, i.e. solitary thyrses, are
characteristic of most species of Calathea. The unit thyrse has an indeterminate main axis,
Synflorescences (and simple inflorescences) are or, if the flower pair is interpreted as a cymule,
usually terminal, but lateral in many species with then it is determinate in some species of
elaborate shoot systems (e.g. Ischnosiphon sects. Stromanthe and Maranta (cf. Andersson 1981;
Bambusastrum and Rotundifolii), and also in Kunze 1985). The main axis carries distichous or
Myrosma and Saranthe (Andersson 1981). A typo- spiral spathes ("bracts" of most taxonomic de-
logical interpretation radically different from that scriptions), which subtend determinate flower
of Andersson was given by Kunze (1985). Kunze custers, florescence components. The florescence
observed that flower pairs occur terminally in component is made up of 1- or 2-flowered
"Calathea stromata" (supposedly a species of cymules, each with a dorsal prophyll at the base.
Stromanthe), Ctenanthe and Stromanthe san- The cymule axes may be long (dolichoblastic
guinea. Based on this observation, he interpreted cymules; e.g. Donax, Schumannianthus, Maranto-
the terminal flower pair as the main florescence. chloa, Maranta, Fig. 75; Stromanthe) or nearly
While the occasional flower pairs were taken imperceptible (brachyblastic cymules; e.g.
as the basis of interpretation by Kunze, they Calathea, Ischnosiphon). There is no sharp dis-
were regarded as insignificant aberrations by tinction between these categories. A ventral scale,
Andersson (1981). the interphyll, is present immediately above the
 Marantaceae
Fig. 75A-E. Marantaceae A, B Maranta arundinacea. A
Habit. B Flower (arranged). C-E M. amazonica. C Flower
(arranged). D Fruiting cymule enclosed by its subtending
spathe. E Seed. (Kennedy eta!. 1988)
prophyll in several genera, e.g. Phrynium and
Calathea. In genera with interphylls, it is evident
that secondary, tertiary, etc., cymules arise from
the interphyll axils. Bracteoles are characteristic
of several genera (e.g. Donax, Schumannianthus,
Sarcophrynium, Calathea, Ischnosiphon). They are
very variable in shape and often characteristic at
generic level. In many species of Calathea and
Ischnosiphon they are strap-shaped with a glandu-
lar swelling at the tip, and in the Donax group they
are small and glandular throughout. These glands
have been shown to be extrafloral nectaries
(Kirchoff and Kennedy 1985).
The 2-flowered cymule has been interpreted
in several ways: (1) as a reduced dichasium with
the terminal flower aborted (Eichler 1875;
Muller 1885); (2) as a 2-flowered monochasium
(Andersson 1976); (3) as a 2-flowered spikelet
(Kunze 1985), or (4) the entire florescence compo-
nent is a deformed spikelet with the flowers re-
stricted to 1 side of the axis (Thompson 1933). The
relations between branchings and bracteoles in
dolichoblastic cymules seem to favour the mono-
chasial model, which also conforms with the
branching patterns of the rest of the plant. Some
281
arguments raised in favour of Eichler's and
Thompson's interpretations have proved to be
based on erroneous observations (Fahn 1953;
Andersson 1976). Also, the observation (Kirchoff
1983) that the flower pair arises through biparti-
tion of an apical meristem seems in line with the
monochasium rather than with the dichasium
interpretation. Kunze's interpretation was based
on the observation that in some species a 3rd
meristem occurs on the pedicel of 1 of the flowers.
This meristem was thought by Kunze to represent
the apex of the spikelet axis, but its nature seems
obscure. It may equally well be interpreted as a
floral meristem, and, in fact, 3-flowered "flower
pairs" were reported from one species of
Stromanthe by Muller (1885), who interpreted this
3rd flower as the terminal flower of a dichasium.
Andersson (1976) interpreted Muller's observa-
tion as a 3-flowered cymule.
It seems that a moderately dolichoblastic
cymule like that of Phrynium is the most probable
ancestral type, from which both extremely
dolichoblastic and markedly brachyblastic ones
have been derived. Phyletic trends are obscure in
inflorescence structure, however.
FLORAL STRUCTURE AND ANATOMY. In all mem-
bers thoroughly examined, the ovary is trilocular,
but in many genera 2 of the locules are empty
and completely compressed (tribe Maranteae of
Petersen 1889; Schumann 1902; Loesener 1930;
and others). Transitional conditions with ± sup-
pressed ovules in 2 of the locules are found in
Phrynium parviflorum (Tilak and Pai 1968) and
Halopegia (Andersson 1981). Well-developed sep-
tal nectaries are present in all members examined
and have the shape of narrow slots lined with a
gladular tissue (Pai and Tilak 1965). The slots
penetrate the septa down to the base of the locules
and open out with a small pore at the base of the
style.
The androecium has been variously interpreted,
but both ontogeny (Eichler 1884; Kirchoff 1983;
Kunze 1984) and floral anatomy strongly support
Eichler's classical interpretation. According to
this, the androecium consists of 2 whorls. The
outer whorl is represented by 1 or 2 (rarely none)
staminodes, the inner one by a fertile stamen and
2 staminodes. The outer staminodes are usually
petaloid and showy, but sometimes subulate
(Megaphrynium), ± rudimentary (several spp. of
Calathea and Stromanthe), or entirely absent
(Thaumatococcus, several spp. of Stromanthe,
some spp. of Calathea). The stamen is monothecic
and has often a petaloid lateral appendage inter-
282 Marantaceae
preted as a transformed theca. One of the inner
staminodes, the staminodium cucullatum, is a
hood-shaped structure (Fig. 76D), which holds
the style backwards-curved and under tension
until the pollination mechanism is triggered. It
has a lateral appendage (2 in Thalia, a bilobed 1 in
Sarcophrynium, Megaphrynium, Hypselodelphys,
and Trachyphrynium), which is the actual trigger
of the pollination mechanism. The other inner
staminode, staminodium callosum, is firm and
fleshy, at least at base, and is provided with fleshy
knobs on the inner surface. The style is narrowly
cylindrical with the stigmatic surface confined to
the inner surface of a ± funnel-shaped terminal
depression. On the dorsal side of the style, just
below the stigmatic orifice, is a flattened part, the
stamp, to which pollen is stuck when the anther
opens. The stamp is lined distally by a glandlike
Fig. 76A-F. Marantaceae. Thalia pavonii. A Habit. B Flower
in approximately natural arrangement. C Style. D Cucullate
staminode. E Fruit. F Seed. (Kennedy et al. 1988)
swelling, from which a sticky substance is
produced by disintegration of the epidermis; this
substance aids in attaching the pollen onto the
pollinator.
Floral anatomy, mainly the vascularisation, has
been described for Calathea (four spp., Tilak and
Pai 1970), Phrynium (two spp., Tilak and Pai 1968)
and Schumannianthus virgatus (Tilak and Pai
1966).
EMBRYOLOGY. No information is available on an-
ther formation and microsporogenesis. The least
incomplete account of ovule development and
embryology is that of Mauritzon (1936), who also
discussed the older literature. The embryo-sac de-
velopment of Phrynium capitatum was described
by Venkataswaralu (1937).
The ovules are anatropous until tetrads have
been formed, but continued differential growth of
the chalaza and nucellus eventually leads to an
amphitropic ovule. During this process of growth,
chalaza! tissue seems to "enter" the enlarged
nucellus, forming a simple, or ± dilatated or
branched strand of chalaza! tissue within the nu-
cellus. This strand contains elongated, apparently
conductive, cells. As the seed matures, the nucel-
lus is transformed into a starch-storing perisperm,
and the intrusive chalaza! tissue disintegrates,
leaving the so-called perisperm canal.
The ovules are bitegmic and crassinucellate. In
Phrynium, the micropyle is formed by the 2 in-
teguments together, in the other genera the inner
integument is protruding and alone forms the mi-
cropyle. The inner integument is thin and under-
goes little differentiation. The outer integument is
much differentiated in the micropylar region. At a
stage where the embryo sac is organised, divisions
in the central layers of cells give rise to a ridge-
shaped structure on the inner surface. This ridge
continues to enlarge to form a cylinder of integu-
mentary tissue, the Mikropylarkragen, penetrat-
ing into the micropylar region of the nucellus. At
about the same time, periclinal divisions are initi-
ated above the micropylar collar. By continued
growth and subsequent sclerification, these will
eventually give rise to the operculum. Periclinal
divisions of the outer epidermis near the micro-
pyle give rise to the aril. In Thalia dealbata, where
the mature seeds lack an aril, aril development
stops after a few divisions.
The nucellar epidermis cells become high and
palisadelike at an early stage. Peripherally, they
undergo periclinal divisions to form a multiseriate
epidermis, while in the centre they remain undi-
vided and form a ± sclerified nucellar cap.
 Marantaceae 283

The archespore, usually unicellular, differenti- The structural peculiarities of the marantaceous
ates from a hypodermal cell. The archesporia! cell flower are well explained by the pollination
cuts off one external parietal cell and then func- mechanism. The anther develops precociously. In
tions as macrospore mother cell. The embryo sac bud it is pressed against the stamp of the style
is formed from the chalazal macrospore and (stylar cavity of Kennedy 1978). When still in bud,
embryo-sac development proceeds according to it opens and the sticky pollen is attached to the
the normal type. At late stages, an increase in an- stamp. The developing style is included in
tipode number may occur, or in number of anti- the cucullate staminode, by which it is held
pode nuclei. Endosperm formation seems to be backwards-curved under tension due to differen-
Nuclear, but the initial divisions have not been tial lengthening. At this point the flower opens.
observed. When a pollinator forces its head down between
Embryo development has not been described in the callose and cucullate staminodes, guided by
any detail. It seems from Mauritzon's illustrations the callosities of the former, it touches the ap-
(his Figs. 4, 5) that the first divisions are trans- pendage of the cucullate staminode, pressing it to
verse, giving rise to a linear quartet of cells. The the side. This releases the style, which snaps for-
micropylar one remains undivided, or divides ward and downward. The course is such that the
once longitudinally, and has haustoria! function. apex of the style enters the proboscidial fossa of
The remaining three cells give rise to the embryo. the visitor, scraping existing pollen into the
In the mature seed no endosperm is present and funnel-shaped stigmatic cavity. Immediately
the fairly large U-shaped embryo is embedded in afterwards, new pollen is deposited from the
copious starchy perisperm. stamp. Deposition is aided by the sticky substance
formed by the glandlike tissue distally lining the
PoLLEN MoRPHOLOGY. Detailed descriptions of stamp.
pollen morphology in the Marantaceae are lack- Some species of Calathea are pseudocleisto-
ing. As seen under the light microscope, pollen gamous. The flowers do not open spontane-
grains are large, spherical and completely sym- ously, but are forcibly opened by the pollinator
metric at maturity. Sporopollenin is (nearly) ab- (Kennedy 1978).
sent from the sporoderm and the thick (ca. 15 !J,m) The species studied by Kennedy are mostly
pollen wall seems to be made up of a pectic, 2- monophilic, while the bees are polytrophic. The
layered "intine". The thick outer layer is radially factors determining the specificity of the pollina-
striate. It seems therefore that the pollen wall is tors are partly ecologic (habitat preferences of
basically similar to that of Canna (described in plants and pollinators), partly "morphologic fit",
detail by Kress and Stone 1982), except that the i.e. the relationship between lengths of floral tube
surface is psilate. and proboscis.
From the artificial pollinations made by
KARYOLOGY. Numerous counts have been re- Kennedy (1978), it seems that most species are
ported (see Bisson et al. 1968; Mahanty 1970), but self-compatible, but one case of suspected self-
mostly for material of uncertain identity. Many incompatibility was also reported. Seed set is gen-
reports, possibly the majority, are obviously based erally very low in Marantaceae, often as low as 1
on erroneous counts. The chromosomes are very seed per 100 flowers, or less. This may relate to the
small ( < 1-1.5 !J.m), sticky and difficult to stain. A hazards of a complex pollination mechanism,
reexamination is badly needed. which are obviously compensated for by large
numbers of flowers, large seeds with copious
POLLINATION AND REPRODUCTIVE SYSTEMS. perisperm, and by the longevity of the mature
Data are available only for American members of plant.
the family (Kennedy 1978). Here, euglossine bees Truly cleistogamous flowers (as opposed to the
of the genera Euglossa, Eulaema, Euplusia, and pseudocleistogamous ones referred to above) are
Exaerete have been reported to visit representa- known to occur in two species of Calathea sect.
tives of most New World genera. Pollination by Microcephalum (Kennedy et al. 1988) and in one
species of Euglossa and Eulaema has been studed species of Maranta (L. Andersson, unpubl.). Auto-
in detail. One aberrant species of Calathea, C. gamy in chasmogamous flowers is also suspected
timothei, is supposed, on circumstantial evidence, on the basis of unexpectedly high fruit set in most
to be bird-pollinated (Kennedy 1977). No data ex- species of Maranta subgen. Maranta (Andersson
ist for species of the Old World, where euglossines 1986), in Thalia geniculata, and several species of
are absent. Ctenanthe and Stromanthe. In several species of
284 Marantaceae
Maranta and one of Ctenanthe, these suspicions
are further strengthened by a high fruit set under
greenhouse conditions in temperate regions,
where effective pollinators are absent (L.
Andersson, unpubl.). The mechanism behind this
has not been described.
FRUIT AND SEED. The basic type of fruit in the
family is a loculicidal capsule with a moderately
thick parenchymatous wall. Such capsules are
found in several genera, e.g. Calathea, Stroman the
and Maranta. In several genera the pericarp is
very thin with the outer layers ± sclerified and
the inner layers degenerated (e.g. Monotagma,
Ischnosiphon, Calathea spp.). By further reduction
of the pericarp, an indehiscent, caryopsislike fruit
is formed with a thin papery or membranous peri-
carp (e.g. Thalia).
Sarcophrynium has berrylike indehiscent fruits
with a fleshy pericarp. In Thaumatococcus, also
with indehiscent fruit, the pericarp is leathery and
the seeds are surrounded by a mucilaginous pulp.
The seeds are fairly large (3-20mm long) and
subglobose to pyramidal or deformed-ellipsoid.
The seed coat is smooth to knobbly but never
quite regularly sculptured. The mature testa con-
sists mainly of a silicified endotesta, derived from
the inner layers of the outer integument, and a
tanniniferous exotesta derived from the middle
layer of the outer integument. The outer layers of
the outer integument and the inner integument
are compressed or ± totally obliterated in the
mature seed (Grootjen 1983). There seems to be a
considerable variation in testa anatomy (Desler
1994), but this has not been systematically
explored.
The micropyle persists in the mature seed as a
germinal aperture, closed by a lid formed from the
inner epidermis of the outer integument. In spe-
cies with dehiscent fruits, the seeds are always aril-
late (see Embryology, above), the aril being much
reduced in tardily dehiscent fruits and absent in
indehiscent fruits or, perhaps, transformed into a
mucilaginous pulp in Thaumatococcus.
Endosperm is absent from the mature seed and
the vermicular horseshoe-shaped embryo is em-
bedded in a starch-storing perisperm. [The em-
bryo was reported by Desler (1994) to be straight
in Donax, Monotagma and Phacelophrynium.]
Starch is present in the form of excentric, single or
aggregated grains that fill the cells ± completely
(Desler 1994).
From the point of funicular attachment, the seed
is penetrated by a canal, which is formed through
the disintegration of an apparently conductive tis-
sue formed by an "intrusion" of chalazal tissue
into the nucellus (Grootjen 1983). This perisperm
canal may be simple throughout (Ischnosiphon,
Monotagma), laterally dilatated in various ways
( Calathea, Haumania), distally branched (Donax,
Hypselodelphys, Maranta, Myrosma, a.o.), or
branched throughout (Thalia).
Seeds germinate only after a prolonged period
of dormancy (1-several months, or even a year),
and often only after removal of the aril. Desicca-
tion seems to kill the seeds of all species of wet
evergreen forest, but may be tolerated by some
species of more seasonal habitats (as indicated by
Kennedy's data).
DISPERSAL. Most species with arillate seeds are
probably myrmecochorous. Horvitz and Beattie
(1980) report that seeds of Calathea microcephala
and C. ovandensis are collected by ponerine
ants and carried to the nest where the arils are
eaten and the seeds discarded. Twenty one species
of ants were observed to be attracted by Calathea
seeds. Large numbers of ants have also been seen
coming to stored seed packages of Calathea
altissima, Ischnosiphon hirsutus, and I. cerotus
(pers. observ.). Horvitz and Beattie report that
germination is much improved by removal of the
aril, which is also the case with seeds of several
other species of Calathea, Monotagma and
Ischnosiphon (pers. observ.). When macerated
under water, the aril releases an oily substance
(Andersson 1977), which may be the actual attrac-
tant. Arils of Ischnosiphon arouma have an intense
spicy fragrance, not observed in other species.
Some species of Calathea, such as C. altissima
and C. inocephala, have bright blue seeds that re-
main well exposed in the opened capsule for a long
time. These may be bird-dispersed, although this
has not actually been observed. Fruits with fleshy
pericarp or pulp are obviously also dispersed by
birds or other vertebrates, but again factual obser-
vations are lacking. The caryopsislike fruits of
Thalia may be water-dispersed.
PHYTOCHEMISTRY (Hegnauer 1963, 1986). The
family is notable for the accumulation of silica (in
form of stegmata), calcium oxalate (as rhomboid,
prismatic and similarly shaped crystals, but not
as rap hides) and starch. Condensed tannins
are based on procyanidin, the floral pigments
(in Thaumatococcus danielli) on delphinidin,
cyanidin and pelargonidin. Proteins of high mo-
lecular weight, which are contained in the aril, the
thaumatins, have been found to be 1600 times
sweeter than sucrose. Cell wall-bound ferulate has
 Marantaceae
been recorded for Maranta bicolor by Harris and
Hartley (1980).
SUBDIVISION AND RELATIONSHIPS WITHIN THE
FAMILY. Petersen (1888) and most subsequent
authors (e.g. Schumann 1902; Loesener 1930) have
subdivided the family into two tribes, Maranteae
(1 fertile ovary locule) and Phrynieae (3 fertile
locules). This classification separates very similar
genera, and variation in virtually all other charac-
ters shows no or little correlation. The classifica-
tion was criticized by Andersson (1981), who
proposed an informal arrangement of the Ameri-
can genera into five groups. A sixth group was
proposed by Andersson in Dahlgren et al. (1985)
for the Palaeotropical genera related to Phrynium.
A recent limited analysis by Mats Hagberg (pers.
comm.) indicates that Monotagma is related to
Calathea and Ischnosiphon, while Koernickanthe
is closely related to Maranta subgen. Koernickea,
thus indicating that Andersson's (1981) Mono-
tagma group is artificial. For many palaeotropical
genera, data are still lacking or incomplete on
many critical points, and there is presently no ba-
sis for a formal classification accounting for the
entire family. In the following treatment, genera
are grouped according to overall similarity and on
the basis of available data, however imperfect
these may be.
AFFINITIES. The position of the family in the
order Zingiberales has never been doubted. It is
unusually well supported by evidence from
vegetative and reproductive morphology, as well
as anatomy and embryology. It is also supported
by analyses of rbcL sequences (Chase et al. 1993;
Duvall et al. 1993). Cladistic analyses of morpho-
logical and anatomical data (Kress 1990) unam-
biguously support the hypothesis that the
Cannaceae form the sister group of the Maranta-
ceae and that these, together with the Zingi-
beraceae, form a monophyletic group within the
order. The relationships among the rest of the
families of the order are still ambiguous, but most
of Kress' (1990) analyses indicate that the other
families form a paraphyletic grade relative to the
herbaceous families.
The Marantaceae are well characterized by a
number of unique synapomorphies: pulvinus,
shoots with a characteristic sequence of organs, a
highly specialised pollination mechanism with a
far-reaching modification of the inner androecial
whorl and the style. Synapomorphies shared with
the Cannaceae and Zingiberaceae sensu lato in-
clude 1 arc of air canals in the leaf axis, absence of
285
raphide sacs, druse-shaped silica bodies, 1 fertile
stamen, inner median stamen fertile, outer me-
dian stamen not fertile, outer lateral stamens
staminodial, inner whorl lateral stamens stamina-
dial, and perisperm present. Synapomorphies
shared with the Cannaceae include asymmetric
flowers, free sepals, petals fused at base, free
staminodes, bisporangiate anthers, modified style
and nuclear endosperm (Kress 1990).
DISTRIBUTION AND HABITATS. The Marantaceae
are nearly exclusively tropical, albeit a few species
slightly exceed the tropics in N and S America.
The majority of species are neotropical, i.e. about
450, some 300 of which are contributed by the
genus Calathea alone. Four major ranges may be
discerned in the Americas, one comprising the
Pacific lowlands and western Andean slopes, and
Central America north to Guatemala and S
Mexico; one comprising the eastern Andean
slopes and western Amazon Basin from Bolivia to
central Colombia; one including the Guianas and
the eastern Amazon Basin; and one comprising
the coastal rain forest of SE Brazil.
The African flora is poor is species, some 30-35
in total, but comparatively rich in genera (10-12).
This reflects great diversity in growth habit,
inflorescence and fruit structure. The African dis-
tribution extends from Sierra Leone to Sudan,
Uganda and Zimbabwe, but is heavily centred in
the west.
The Asian flora is partly poorly understood. No
modern treatment exists covering any substantial
part of the range. It is presently believed to include
some 50 species in eight genera, but this is prob-
ably an underestimate, at least for the species
number. The Asian range extends from Sri Lanka
and Assam to Formosa and the Philippines, but is
pronouncedly centred in the Indomalesian area.
Most species are confined to tropical rainforest
habitats at altitudes below 1000m, where they
show a considerable amount of ecologic diversifi-
cation. They are mainly "jungle weeds" in the
sense that they require some degree of disturbance
in order to flower and fruit. Rather few are "true"
understorey species flowering and fruiting in deep
shade (e.g. Monophyllanthe oligophylla, some
Calathea spp.). Some species, usually small to
medium, germinate and persist in a vegetative
state in the full shade of a closed canopy, being
ready to flower as soon as opportunity arises.
Others, usually larger species, are unable to grow
for long in deep shade, but complete their life
cycle in larger light gaps of longer duration, such
as are caused by landslides, extensive storm-
286 Marantaceae

felling etc. Relatively few species are able to persist 5. Lower branch of cucullate staminode appendage several
in permanent openings such as stream margins, times the length of the upper one; caulescent plants 6
- Branches of cucullate staminode appendage (sub)equal;
etc. Species of the two latter categories often show rosulate plants 7
good ability to invade secondary habitats, while 6. Fruit indehiscent; seeds exarillate 13. Hypselodelphys
the species of small light gaps mostly perish where - Fruit dehiscent; seeds arillate 14. Trachyphrynium
forests are cut over. 7. Bracteoles 1 per cymule; outer staminodes stiff, subulate;
Some few species occur in semideciduous and fruit dehiscent 15. Megaphrynium
- Bracteoles 2 per cymule; outer staminodes petaloid; fruit
deciduous forests (e.g. Maranta arundinacea). indehiscent 16. Sarcophrynium
They show a highly seasonal pattern of growth, 8. Cymules 1-flowered 9
often dying back to the rhizome in the dry season. - Cymules 2-flowered 10
The genera Thalia and Halopegia (H. azurea, at 9. Ovary with 3 fertile locules 3. Monophrynium
- Ovary with I fertile locule 10. Monotagma
least) are confined to open marshes. A larger num-
10. Ovary with 3 ovulate locules, but 1 or 2 of the ovules
ber of species (in Calathea, Ischnosiphon, Mono- sometimes poorly developed 11
tagma) have their primary habitats in swamp - Ovary with a single ovulate locule, the 2 others completely
forest and riverine inundation forest (igap6 and compressed and empty 23
varzea). 11. Outer staminodes absent; fruit ca. 3 em in diam., winged,
with a thick leathery exocarp and a mucilaginous pulp
29. Thaumatococcus
EcoNOMIC IMPORTANCE. The only important - Outer staminodes present (except in a few species of
crop plant of the family is Maranta arundinacea, Calathea), petaloid; fruit smaller, not winged, and of dif-
cultivated for the high-quality starch of the rhi- ferent texture 12
zomes. Several species of Calathea (e.g. C. 12. Cymule axes ± crescent-shaped in cross-section; 1 sepal
much smaller than the 2 others 27. Halopegia
lancifolia, C. makoyana) and Maranta (e.g. M. hi- - Cymule axes elliptic or circular to subrectangular in
color, M. leuconeura) are economically important cross-section; sepals subequal 13
ornamentals in common trade. A much larger 13. Spathes large and suborbicular, showy white, persistent
number of species of Calathea, Maranta, Saran the 31. Haumania
and Ctenanthe occur occasionally in horticultural - Spathes ± elliptic, greenish, purplish, or brownish, often
caducous 14
trade, but have little significance economically, 14. Outer staminodes 2 per flower 15
since they are impossible to cultivate indoors - Outer staminodes solitary 21
without special arrangements. It seems, however, 15. Caulescent plants with leaves clustered distally on long,
that the importance and diversity of Marantaceae canelike stems; cucullate staminode unappendaged
2. Ataenidia
as ornamentals in tropical and subtropical horti-
- Rosulate plants with most leaves at ground level, or leaves
culture is rapidly increasing. solitary from the rhizome; cucullate staminode
In aboriginal economy, a fairly large number of appendaged 16
species are used. Many species of Calathea and 16. Corolla tube at least twice the length of the lobes 17
Maranta with tuberous roots are locally culti- - Corolla tube much shorter, usually equalling the lobes or
shorter 18
vated, and inflorescences of several Calathea spp. 17. Sepals about 1/2 the length of corolla tube 30. Cominsia
are cooked and eaten by the natives in Central - Sepals 1/3 the length of corolla tube or, usually, much less
America (Kennedy 1978). Split stems of tall- 5. Stachyphrynium
growing species of Ischnosiphon are used for 18. Cucullate staminode appendage at the base of the lobe;
basket-weaving and the leaf blades of many larger cymules usually conspicuously dolichoblastic with both
peduncular and pedicellar axes well developed 19
species are used for wrapping up food items. - Cucullate staminode appendage from distal half of the
lobe; cymules usually subbrachyblastic to brachyblastic,
either peduncle or pedicel poorly developed 20
KEY TO THE GENERA 19. More or less robust plants, usually 1-3m tall, leaves and
inflorescences on the same shoot 17. Marantochloa
1. Bracteoles present, small and glandular 2 - Small plants with leaves and inflorescences on separate
- Bracteoles absent, or scale-, sheath-, or ribbon-shaped 8 shoots 18. Afrocalathea
2. Appendage of cucullate staminode simple 3 20. Spathes spirally arranged; inflorescence often a much
- Appendage of cucullate staminode 2-branched 5 congested, ± capitate synflorescence 1. Phrynium
3. Corolla tube long and narrow; ovary densely tuberculate; - Spathes distichous; inflorescence simple or a lax
outer staminodes solitary 6. Calathea synflorescence of fasciculate florescences
- Corolla tube short; ovary smooth; outer staminodes paired 4. Phacelophrynium
4 21. Bracteoles absent; corolla tube usually equalling the lobes,
4. Fruit indehiscent; stamina! tube obsolete 11. Donax or shorter 4. Phacelophrynium
- Fruit dehiscent; stamina! tube long and conspicuous - Bracteoles usually present, scale-, sheath- or ribbon-
12. Schumannianthus shaped; corolla tube usually longer than lobes 22
 Marantaceae 287

22. Florescence coarsely spiciform to capitate, often several Genera of the Marantaceae
em in diam. 6. Calathea
- Florescence narrowly spiciform with tightly rolled spathes,
ca. 1 em in diam. or less 7. Sanblasia I. Phrynium group
23. Cucullate staminode with double appendage; style with a
very long process from the lower rim of the stigmatic Cymules brachyblastic to slightly dolichoblastic,
orifice; perisperm canal branched from base 28. Thalia bracteoles absent, interphylls usually present;
- Cucullate staminode with simple, sometimes obsolete corolla tube (short to) moderately long (to long),
appendage; stigmatic orifice without long process; peri-
sperm canal simple, or branched distally 24 (1)-2 outer staminodes.
24. Bracteoles present 25
- Bracteoles absent 28
25. Corolla tube 10-30 times longer than wide 26
1. Phrynium Willd.
- Corolla tube twice as long as wide, or shorter 27 Phrynium Willd., Sp. Pl. 1: 17 (1797), nom. cons.; emend. K.
26. Florescences laterally flattened, the spathes conduplicate Schum., Pflanzenreich IV. 48: 28, 52-58 (1902); Milne-
also in bud; bracteoles keeled 8. Pleiostachya Redh., Kew Bull. 1952: 167-168; Holttum, Gard. Bull.
- Florescences ± terete, the spathes tightly rolled, at least in Singapore 13: 279-294 (1951).
bud; bracteoles not keeled 9. Ishnosiphon
27. Corolla tube as long as wide, or longer; callose staminode
distally petaloid 25. Ctenanthe Rosulate herbs. Inflorescence a usually congested
- Corolla tube shorter than wide; callose staminode firm and and richly branched synflorescence, florescences
fleshy throughout 26. Stroman the with spiral spathes; cymules 2-flowered, +
28. Leaf sheath flanks beginning to wither as soon as the blade brachyblastic to slightly dolichoblastic, usually
unfolds, leaving a reticulum of fibres ("fenestrate
sheaths") 24. Hylaeanthe with interphylls, bracteoles absent. Sepals half the
- Leaf sheaths not fenestrate 29 length of corolla tube or (usually much) longer;
29. Small plants, each aerial shoot with a single basal leaf corolla tube half the length oflobes, rarely longer;
20. Monophyllanthe outer staminodes 2, petaloid; cucullate staminode
- Basal leaves 2 or more together, or larger plants with leaves
on ± richly branched shoot systems 30
with appendage near apex; ovary with 3 ovulate
30. Leaves homotropic, all blades of the same shoot being of locules. Fruit dehiscent; seeds arillate. About 20
the same symmetry form 31 spp., India and Sri Lanka eastwards to New
- Leaves antitropic, the broadest side of the blade alter- Guinea.
nating between right and left in successive leaves of an axis
35
31. Corolla tube more than twice the length of the lobes 32 2. Ataenidia Gagnep.
- Corolla tube less than twice the length of the lobes 33
32. Cymules brachyblastic with axial system ca. 2mm long or Ataenidia Gagnep., Bull. Soc. Bot. Fr. 55: XLI-XLII (1908);
less 21. Koernickanthe Milne-Redh., Kew Bull. 1952: 167 (1952).
- Cymules ± dolichoblastic with axial system 5 mm long or
(usually much) more 19. Maranta Caulescent herbs. Inflorescence a much-branched,
33. Florescence strongly monosymmetrical; spathes white or
pale green when fresh, straw-coloured when dry
congested synflorescence; spathes persistent;
22.Myrosma cymule + brachyblastic, interphyll present,
- Florescences usually radially symmetrical or bisymmetric; bracteoles absent. Corolla tube ± equalling the
spathes green or sometimes purplish tinged 34 lobes; outer staminodes 2, slightly unequal; cucul-
34. Florescence component usually of several cymules; lateral late staminode without appendage; ovary with 3
synflorescences absent; corolla tube as long as wide, or
longer 19. Maranta
ovulate locules. Fruit with papyraceous pericarp;
- Florescence component of a single cymule; lateral seeds arillate. Only one sp., A. conferta (Benth.)
synflorescences usually present; corolla tube as long as Milne-Redh., W Africa to Central African
wide, or shorter 23. Saranthe Republic, Sudan and Uganda.
35. Corolla tube as long as wide, or longer; callose staminode
distally petaloid; corolla and staminodes exceeding the
sepals 36 3. Monophrynium K. Schum.
- Corolla tube shorter than wide; callose staminode firm and
fleshy throughout; corolla and inner staminodes usually Monophrynium K. Schum., Pflanzenreich IV. 48: 68 (1902).
not exceeding the sepals 26. Stroman the
36. Florescence of several or numerous, densely imbricated Rosulate herbs. Inflorescence a ± richly branched
spathes; axial system of cymule less than 1 em long
25. Ctenanthe
synflorescence, florescences narrowly spiciform,
- Florescence of 1 or 2 slightly overlapping spathes, cymule spathes persistent. Florescence component of 1-
peduncle ca. 2 em, or more 19. Maranta several !-flowered cymules; cymule subbrachy-
blastic, with or without interphyll, bracteoles
absent. Corolla tube ± equalling the lobes, or
288 Marantaceae
somewhat shorter; outer staminodes 2, petaloid;
cucullate staminode with a small, simple append-
age; ovary with 3 fertile locules. Fruit and seed
unknown. A few spp., endemic to the Philippines.
4. Phacelophrynium K. Schum.
Phacelophrynium K. Schum., Pflanzenreich IV. 48: 120-123
(1902); Holttum, Gard. Bull. Singapore 13: 273-279 (1951).
Rosulate plants. Inflorescence a moderately
branched synflorescence; florescence bisym-
metric; spathes persistent. Cymule subbrachy-
blastic to brachyblastic; interphyll usually present;
bracteoles absent. Sepals equalling the corolla
tube or longer; corolla tube equalling lobes or
shorter; outer staminodes 1, petaloid; ovary with 3
ovulate locules. Fruit dehiscent, seeds arillate.
About six spp., Thailand, Malacca Peninsula,
Malesian Islands and the Philippines.
5. Stachyphrynium K. Schum.
Stachyphrynium K. Schum., Pflanzenreich IV. 48: 45-49
(1902); Holttum, Gard. Bull. Singapore 13: 275-279 (1951).
Rosulate plants. Inflorescence simple or a sparsely
branched synflorescence, florescences ± coarsely
spiciform, spathes distichous, persistent. Cymule
brachyblastic, interphyll present or absent,
bracteoles absent. Sepals 1/3 the length of corolla
tube or (usually) much shorter; corolla "tube"
solid, twice the length of lobes or longer; outer
staminodes 2, petaloid; ovary with 3 ovulate
locules. Fruit dehiscent, seeds arillate. About ten
spp., India, Sri Lanka and Indonesia. The distinct-
ness of the genus was doubted by Holttum, but the
very long-tubed flowers with short, triangular,
membranous sepals seem to be a good distinctive
character.
II. Calathea group
Cymules brachyblastic, interphylls usually
(Calathea) or occasionally present, bracteoles
usually present; corolla tube long or very long, one
outer staminode.
6. Calathea G. F. W. Meyer
Calathea G. F. W. Meyer, Primit. Fl. Essequebonensis: 6-7
(1818); Andersson, Nord. J. Bot. 1: 237-238 (1981);
Kennedy, Univ. Calif. Pub!. Bot. 71 (1978); Kennedy, Fl.
Ecuad. 32: 14-136 (1988).
Thymocarpus Nicolson, Steyerm. & Sivadasan (1981).
Rosulate herbs. Inflorescence simple or a sparsely
to moderately branched synflorescence; flore-
scences usually coarsely spiciform to capitate,
spathes distichous or spiral. Cymules brachy-
blastic or, exceptionally, subbrachyblastic or
dolichoblastic, interphyll usually present, brac-
teoles usually present, 1-4 per cymule, glandular,
scale- or sheathlike, or ribbon-shaped and
claviculate. Corolla tube 5-25 times longer than
wide, rarely longer; outer staminode 1, rarely 0,
petaloid; ovary with 3 ovulate locules. Fruit dehis-
cent, seeds arillate. About 300 spp., all over the
humid American tropics.
7. Sanblasia L. Andersson
Sanblasia L. Andersson, Nord. J. Bot. 4: 21 (1984).
Rosulate plants. inflorescence a sparsely branched
synflorescence, florescences narrowly spiciform
with rolled spathes. Cymule brachyblastic,
interphyll present, bracteoles 2 per cymule,
sheathlike. Corolla tube ca. 20 times longer than
wide; outer staminode 1, petaloid; ovary with
3 fertile locules. Only one sp., S. dressleri L.
Andersson, Serrania de San Blas, Panama.
Sanblasia is nearly intermediate between Calathea
and Ischnosiphon.
8. Pleiostachya K. Schum. Fig. 74A,B
Pleiostachya K. Schum., Pflanzenreich IV. 48: 164 (1902);
Andersson, Nord. J. Bot. 1: 238 (1981).
Rosulate herbs. Inflorescence a moderately
branched synflorescence; florescences rather
stoutly spiciform, laterally flattened, with condu-
plicate spathes, cymules brachyblastic, interphyll
sometimes present, bracteoles 2, sheathlike.
Corolla tube ca. 15 times longer than wide; outer
staminode 1, petaloid; ovary uniovulate. Two spp.,
Central America and the Pacific lowland south to S
Ecuador.
9. Ischnosiphon Korn.
Ischnosiphon Korn., Mem. Soc. Imper. Nat. Moscou 11: 346
348 (1859); emend. K. Schum., Pflanzenreich IV. 48: 156-
158 (1902); Andersson, Opera Bot. 43 (1977), rev., Nord. J.
Bot. 4: 25-32 (1984).
Rosulate or caulescent, sometimes very tall plants.
Inflorescence simple, or a ± distinct synflore-
scence; florescences narrowly spiciform, terete
with rolled spathes (before anthesis). Cymules
brachyblastic, interphyll rarely present, bracteoles
 Marantaceae
1 or 2, usually ribbon-shaped and claviculate,
sometimes scalelike, never sheathlike. Flowers as
in Pleiostachya. About 35 spp., throughout the
humid American tropics.
10. Monotagma K. Schum. Fig. 74C-E
Monotagma K. Schum., Pflanzenreich IV. 48: 166 (1902).
Rosulate herbs. Inflorescence a usually richly
branched synflorescence, often congested;
florescences narrowly spiciform with sclerotic,
rolled spathes. Cymules 1-flowered, subbrachy-
blastic, interphyll (usually) and bracteoles absent.
Corolla tube very long and narrow; outer
staminode 1, petaloid; ovary with 1 ovulate locule.
Thirty seven spp. (Mats Hagberg, pers. comm.),
widespread in humid parts of S America.
III. Donax group
Cymules ± dolichoblastic, interphylls absent,
bracteoles present, small and glandular; corolla
tube short (to moderately long), 2 outer sta-
minodes.
11. Donax Lour.
Donax Lour., Fl. Cochinch.: 14 (1790); Holttum, Gard. Bull.
Singapore 13: 266-271 (1951); Rolfe, J. Bot. 45: 242-244
(1907).
Actoplanes K. Schum. (1902).
Tall plants with compound leaf cluster on top of
canelike stem. Inflorescence a synflorescence of
narrowly spiciform florescences. Cymules mark-
edly dolichoblastic, interphyll absent, bracteoles 2
per cymule, small, glandular. Corolla tube equal-
ling the lobes or shorter; stamina! tube incon-
spicuous; outer staminodes 2, petaloid; ovary
with 3 ovulate locules. Fruit indehiscent, seeds
exarillate. Three or four spp., SE Asia from
Malaysia eastwards to the New Hebrides. Donax
differs from Schumannianthus in length of the
stamina! tube and in having indehiscent fruit. The
distinction is not entirely convincing.
12. Schumannianthus Gagnep.
Schumannianthus Gagnep., Bull. Soc. Bot. Fr. 51: 169-177
(1904); Holttum, Gard. Bull. Singapore 13: 271-273 (1951);
Rolfe, J. Bot. 45: 244 ( 1907).
Habit like Donax. Cymules markedly dolicho-
blastic, without interphyll, with 1 or 2 glandular
289
bracteoles. Corolla tube 1/2 the length of the lobes
or less, split to the base; stamina! tube conspi-
cuous; outer staminodes 2, petaloid; ovary with 3
ovulate locules. Fruit dehiscent, seeds arillate.
Two spp., one from Sri Lanka and Malabar, S.
virgatus (Roxb.) Rolfe, and one from Malaysia,
Indochina and the Philippines, S. dichotomus
(Roxb.) Gagnep.
13. Hypselodelphys (K. Schum.) Milne-Redh.
Hypselodelphys (K. Schum.) Milne-Redh., Kew Bull. 1950:
158-161 (1950); Hepper, Fl. Trop. W Africa 3(1): 88-89
(1968).
Trachyphrynium subgen. Hypselodelphys K. Schum. (1902).
Scandent or subscandent, much-branched plants.
Cymules distinctly dolichoblastic, without inter-
phyll, bracteoles 2 per cymule, glandular. Corolla
tube ca. 1/4 of the length of the lobes; outer
staminodes 2, ± unequal, petaloid; cucullate
staminode with 2-branched appendage, the upper
branch much shorter than the lower one; ovary
with 3 ovulate locules. Fruit 3-lobed, densely
muricate, indehiscent; seeds exarillate. Four spp.,
Sierra Leone to the Congo Basin.
14. Trachyphrynium Benth.
Trachyphrynium Benth. in Benth. & Hook. f., Gen. Pl. 3: 651
(1883); emend. Milne-Redhead, Kew Bull. 1950: 157-158
(1950).
Hybophrynium K. Schum. (1892).
Scandent, much-branched plants. Cymules slightly
dolichoblastic, prophyll ephemeral, interphyll
absent, bracteoles 2, glandular. Flowers as in
Hypselodelphys. Fruit dehiscent, seeds arillate.
Only one sp., T. braunianum (K. Schum.) Milne-
Redh., W Africa, the Congo Basin, Sudan and
Uganda. The only important difference from
Hypselodelphys is in fruit structure, and the dis-
tinction is not very convincing.
15. Megaphrynium Milne-Redh.
Megaphrynium Milne-Redh., Kew Bull. 1952: 169-170 (1952).
Rosulate plants. Inflorescence a richly branched,
congested synflorescence; spathes caducous;
florescence component of a single cymule. Cymule
± distinctly dolichoblastic, without interphyll,
bracteole 1 (rarely 2), glandular. Corolla tube ca.
1/2 the length of the lobes; outer staminodes 2 ( 1),
often markedly unequal, subulate and erect;
cucullate staminode with 2-branched appendage,
290 Marantaceae
branches ::±: equal, divergent; ovary with 3 ovulate
locules. Fruit dehiscent. About four spp., W
Africa, the Congo Basin, Sudan and Uganda. Very
close to Sarcophrynium, and possibly better treated
as a subgenus.
16. Sarcophrynium K. Schum.
Sarcophrynium K. Schum., Pfianzenreich IV. 48: 35-40 (1902);
emend. Milne-Redh., Kew Bull. 1952: 168-169 (1952).
Rosulate herbs. Inflorescence a congested syn-
florescence, spathes persistent or tardily cadu-
cous; florescence component of ca. 5 cymules;
cymules evidently dolichoblastic, without inter-
phyll, bracteoles 2, glandular. Corolla tube ca.
1/2 the length of the lobes, or less; outer stami-
nodes 2(1), petaloid; cucullate staminode as in
Megaphrynium; ovary with 3 ovulate locules. Fruit
fleshy, indehiscent; seeds with a mucilaginous,
mantlelike aril. About three spp., W Africa, the
Congo Basin and Uganda.
IV. Maranta group
Cymules ::±: dolichoblastic, interphylls absent,
bracteoles absent or rudimentary; corolla tube
moderately long to long, (usually) 2 outer
staminodes.
17. Marantochloa Brogn. ex Gris
Marantochloa Brogn. ex Gris, Bull. Soc. Bot. Fr. 7: 320-322
(1860).
Shrub-shaped plants, usually 1-3m tall. Cymules
::±: dolichoblastic, often very pronouncedly so,
interphyll absent (or very rarely present),
bracteoles absent (rarely rudimentary). Corolla
tube very short to ::±: equalling the lobes; outer
staminodes 2, petaloid, equal to markedly
unequal; cucullate staminode with a short flat
appendage near middle of lobe; ovary with 3
ovulate locules. Fruit (tardily) dehiscent; seeds
arillate. About 15 spp., W Africa, the Congo Basin,
Sudan, Uganda, Tanzania, Reunion and the
Cornaro Islands.
18. Afrocalathea K. Schum.
Afrocalathea K. Schum., Pfianzenreich IV. 48: 51-52 (1902).
Small acaulescent herbs with leaves and inflore-
scences on separate shoots, inflorescence simple.
Cymule evidently dolichoblastic, interphyll and
bracteoles absent. Corolla tube ::±: equalling the
lobes (ca. 10 times longer than wide); outer
staminodes 2, slightly unequal, petaloid; cucullate
staminode with a short, flat appendage near base
of lobe; ovary with 3 ovulate locules. Fruit and
seed unknown. Only one sp., C. rhizantha K.
Schum., W Africa. The genus has been considered
close to Calathea (Andersson 1981), but inflore-
scence organisation and floral structure agree
better with Marantochloa.
19. Maranta L. Fig. 75
Maranta L., Sp. Pl. 1: 2 (1753); emend. Eichler, Abh. K. Akad.
Wissensch. Berlin 1883: 75-77 (1884); K. Schum.,
Pfianzenreich IV. 48: 123-137 (1902); Andersson, Nord. J.
Bot. 1: 227-228 (1981), ibid. 6: 729-756 (1986), rev.
Leaves homotropic or rarely antitropic; cauline
leaves separated by distinct internodes. Cymules
::±: markedly dolichoblastic, interphyll absent,
bracteoles absent (rarely rudimentary). Corolla
tube 1/2 to twice the length of the lobes; outer
staminodes 2, petaloid; cucullate staminode with a
small, flat appendage near base oflobe; ovary with
1 ovulate (and 2 compressed) locules. Fruit dehis-
cent, seeds arillate, perisperm canal distally
branched. About 25 spp., humid areas of tropical
America.
20. Monophyllanthe K. Schum.
Monophyllanthe K. Schum., Pflanzenreich IV. 48: 165-166
(1902); Andersson, Nord. J. Bot. 1: 228-229 (1981).
Aerial shoots with a single basal leaf each, and a
small, simple inflorescence. Cymule markedly
dolichoblastic, interphyll absent, bracteoles rudi-
mentary, ridge-shaped. Flowers like Maranta, but
1 outer staminode often much reduced or absent.
Fruit dehiscent; seeds arillate, perisperm canal
distally branched. One or two spp., the Guianas
and northern Amazon Basin. Very close to
Maranta, the generic rank being debatable.
21. Koernickanthe L. Andersson
Koernickanthe L. Andersson, Nord. J. Bot. 1: 240-241 (1981).
Rosulate herbs with homotropic leaves. Inflore-
scence a moderately branched synflorescence;
florescences narrowly cylindrical; spathes bright
orange, ::±: papyraceous when dry. Cymules 2-
flowered, subbrachyblastic, interphyll and brac-
teoles absent. Corolla tube very long and narrow,
ca. twice the length of lobes; outer staminodes 2,
 Marantaceae
petaloid, subequal, or 1 of them :±:: reduced or
absent. Only one sp., K. orbiculata (Korn.) L.
Andersson, S Brazil and the eastern Amazon
Basin.
V. Myrosma group.
Cymules subbrachyblastic to dolichoblastic, in-
terphylls absent, bracteoles absent or present;
corolla tube short or very short, usually 2 outer
staminodes; perisperm canal distally branched.
22. Myrosma L. f.
Myrosma L. f., Suppl. Pl.: 8, 80 (1781); Andersson, Nord. J. Bot.
1: 230-231 (1981).
Leaves homotropic. Inflorescence simple or a
sparsely branched synflorescence, terminal, some-
times also lateral; florescence markedly mono-
symmetrical, spathes white or pale green,
persistent. Florescence component of a single
cymule; cymule subbrachyblastic, interphyll and
bracteoles absent. Corolla tube less than 1/2 the
length of the lobes; outer staminodes 2, :±:: equal,
petaloid; callose staminode distally petaloid;
staminal appendage large and showy. Only one
sp., M. cannifolia L. f., widespread in S America,
especially in dry forests.
23. Saranthe (Regel & Korn.) Eichler
Saranthe (Regel & Korn.) Eichler, Abh. K. Akad. Wissensch.
Berlin 1883: 85 (1884); Andersson, Nord. J. Bot. 1: 232-233
(1981), ibid. 5: 61-63 (1985).
Maranta subgen. Saranthe Regel & Korn. (1857).
Ctenophrynium K. Schum. (1902).
Rosulate plants with homotropic leaves. Inflore-
scence a sparsely to richly branched synflore-
scence, synflorescences lateral and mostly also
terminal; florescence bisymmetrical or (usually)
:±:: monosymmetrical, spathes greenish, usually
caducous. Florescence component usually of a
single cymule; cymule subbrachyblastic to evi-
dently dolichoblastic, interphyll and bracteoles
absent. Corolla tube ca. 1/2 the length oflobes, or
less; outer staminodes 2, :±:: equal, petaloid; callose
staminode distally petaloid; staminal appendage
small and obsolete. Five to ten spp., SE Brazil.
24. Hylaeanthe Jonker & Jonker
Hylaeanthe Jonker & Jonker, Acta Bot. Neerl. 4: 175 (1955);
Andersson, Nord. J. Bot. 1: 233-234 (1981).
291
Rosulate herbs with homotropic leaves, leaf
sheath flanks early decaying into a reticulum of
fibres. Inflorescence a sparsely branched syn-
florescence, terminal; florescence :±:: markedly
monosymmetrical; spathes green when young,
often marcescent. Florescence component of 2-3
cymules; cymules evidently dolichoblastic to
(rarely) subbrachyblastic, interphyll and brac-
teoles absent. Corolla tube equalling the lobes or
1/2 longer; outer staminodes 2, :±:: markedly
unequal (one rarely entirely missing), the larger
one, at least, petaloid; callose staminode distally
petaloid. Five or six spp., widespread in the humid
American tropics.
25. Ctenanthe Eichler
Ctenanthe Eichler, Abh. K. Akad. Wissensch. Berlin 1883: 81-
84 (1884); Andersson, Nord. J. Bot. 1: 235 (1981).
Rosulate or caulescent plants with antitropic
leaves; cauline leaves mostly clustered. Inflore-
scence a sparsely to moderately branched, some-
times diffuse synflorescence, terminal; florescence
bisymmetrical to markedly monosymmetrical;
spathes greenish, usually persistent, florescence
component of 2-12 cymules, cymules brachy-
blastic to (usually) subbrachyblastic, interphyll
absent, bracteoles usually present, 1-2, scale- or
sheathlike. Corolla tube 1/2 the length of lobes
to equal; outer staminodes 2, petaloid; callose
staminode distally petaloid. About ten spp., wide-
spread in the humid American tropics, but con-
centrated in SE Brazil.
26. Stromanthe Sonder
Stromanthe Sonder, Neue allgem. deutsche Gart. Blumen-
zeitschr. 5: 225-227 (1849); Andersson, Nord. J. Bot. 1: 235-
236 (1981).
Usuaully caulescent herbs with antitropic, usually
clustered leaves. Inflorescence a richly branched,
but often diffuse, synflorescence; spathes often
brightly coloured, often caducous. Cymules :±:: evi-
dently dolichoblastic, interphyll absent, bracteoles
rarely present, rudimentary or scalelike. Corolla
tube less than 1/2 the length oflobes, mostly obso-
lete; outer staminodes absent or 2, rudimentary
to petaloid; callose staminode firm and fleshy :±::
thoughout. Ten to 15 spp., all over the humid
American tropics, but concentrated in Central
America and SE Brazil.
292 Marantaceae

GENERA OF UNCERTAIN AFFINITY Schumann (1902) subdivided the genus into four
subgenera, which is hardly motivated.
27. Halopegia K. Schum.
29. Thaumatococcus Benth.
Halopegia K. Schum., Pflanzenreich IV. 48: 49-51 (1902).
Thaumatococcus Benth. in Benth. & Hook. f., Gen. Pl. 3: 652
Rosulate marsh plants. Inflorescence simple or a (1883).
sparsely branched synflorescence. Florescence
component of a single cymule; cymule :±:: evidently Rosulate herbs. Inflorescence simple or a very
dolichoblastic, axes conspicuously flattened and sparsely branched synflorescence; spathes cadu-
furrowed, interphyll and bracteoles absent. Sepals cous. Florescence component of a single cymule;
strongly unequal; corolla tube very short to cymule subbrachyblastic, interphyll absent,
obsolete; outer staminode(s) 1, or 2 very unequal, bracteole solitary, glandular. Corolla tube 1/5-
the smaller one sometimes caducous; callose 1/10 the length of lobes; outer staminodes absent;
staminode conspicuously large with narrow peta- ovary with 3 ovulate locules. Fruit large, with thick
loid rim; cucullate staminode with a simple, rather wings, indehiscent, exocarp leathery, endocarp
obsolete appendage near top of the lobe; ovary mucilaginous; seeds with aril of membranous
with 3 ovulate locules, 2 of them poorly developed sheets sticking to testa. Only one sp., T. danielii.
(to completely empty and compressed). Fruit 1- (Benn.) Benth., W Africa and the Congo Basin.
seeded, caryopsislike, indehiscent. About four With a strikingly peculiar fruit, and general
spp., W Africa, the Congo Basin and Indonesia. An anonymity in other characters, affinity is not
isolated genus without obvious affinities within easily guessed. Bracteole shape may indicate
the family. In some features it shows resemblance distant relationship with the Donax group.
to Thalia: organisation of florescence component,
short-tubed flowers with very large callose 30. Cominsia Hemsl.
staminodes, and caryopsislike fruits. Interest-
ingly, 1 of the outer staminodes, and 2 of the ovary Cominsia Hems!., Ann. Bot. 5: 508 (1891).
locules are :±:: reduced in Halopegia, and the two
genera are the only ones in which an additional Inflorescence a :±:: richly branched synflorescence;
adaxial air lacuna has been demonstrated in the spathes caducous. Cymule subbrachyblastic,
leaf axis. interphyll and bracteoles absent. Corolla tube very
long and narrow, 2-3 times the length of lobes;
outer staminodes 2, petaloid; callose staminode
28. Thalia L. Fig. 76 with large petaloid portion; cucullate staminode
with small simple appendage. Fruit a hard-
Thalia L., Sp. Pl. 1: 1193 (1753); Andersson, Nord. J. Bot. 1:
241-243 (1981).
textured, dehiscent capsule; seeds arillate. Only
one sp., C. guppyi Hemsl., in Molucca Islands, New
Rosulate or caulescent marsh plants. Inflorescence Guinea, and Solomon Islands. The very long
an often very richly branched synflorescence; corolla tube is reminiscent of the Calathea group,
spathes caducous. Florescence component of a but cymule structure rather indicates relationship
single cymule; cymule subbrachyblastic, interphyll with the Phrynium group.
and bracteoles absent. Corolla tube obsolete; outer
staminode 1, petaloid, showy; callose staminode 31. Haumania J. Leonard
conspicuously large with narrow petaloid rim; cu-
Haumania J. Leonard, Bull. Jard. Bot. Bruxelles 19: 453-455
cullate staminode with two appendages near base (1949); Milne-Redh., Kew Bull. 1950: 161-163 (1950).
oflobe; style helically twisted when triggered, with Trachyphrynium subgen. Lasiodelphys K. Schum. (1902),
a very long liplike protrusion from the ventral rim partly.
of the stigmatic orifice. Fruit caryopsislike; peri-
sperm canal branched from the base. Five to seven Slender-stemmed, climbing plants. Spathes per-
spp., mainly in seasonally humid parts of S sistent, :±:: orbicular, white. Cymules evidently
America, two spp. also in southern USA. T. dolichoblastic, interphyll and bracteoles absent.
geniculata L. has a widespread population in Corolla tube ca. half the length of lobes; outer
Africa, sometimes regarded as a separate species staminodes 2, petaloid; ovary with 3 ovulate
(cf. Andersson, Nord. J. Bot. 1: 48-56, 1981). locules. Fruit indehiscent, tuberculate; seeds
 Marantaceae
exarillate. Two spp., one W Africa (E Cameroon to
Gabon) and one the Congo Basin. A distant rela-
tionship to Trachyphrynium may be guessed.
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of the Marantaceae II. Vascular anatomy of the genus
Phrynium Willd. Proc. Indian Acad. Sci. 68: 240-249.
Tilak, V.D., Pai, R.M. 1970. Studies in the floral morphology of
the Marantaceae III. Vascular anatomy of some species of
the genus Calathea. Marathwada Univ. J. 9 Sci. 2: 31-41.
Tomlinson, P.B. 1961. Morphological and anatomical charac-
teristics of the Marantaceae. J. Linn. Soc. (Lond.) 58: 55-78.
Tomlinson, P.B. 1974. Development of the stomatal complex
as a taxonomic character in the monocotyledons. Taxon 23:
109-128.
Tran V.N. 1974. Sur le callus des marantacees. Bull. Soc. Bot.
Fr. 121: 97-108.
Venkataswaralu, V. 1937. A note on the development of the
embryo sac in Phrynium capitatum W. J. Indian Bot. Soc.
16: 95-98.
294 Mayacaceae
Mayacaceae
D.W. STEVENSON
Mayacaceae Kunth, Abh. Konig!. Akad. Wiss. Berlin, Phys.
Abh. 1840: 93 (1842), nom. cons.
Aquatic to telmatic perennial, rooted herbs.
Leaves simple, spirally arranged, sessile, without
a basal sheath, lanceolate to linear or nearly
filiform, commonly bidentate apically. Flowers
axillary, solitary on short to long pedicels, emer-
gent, actinomorphic, perfect, hypogynous, trimer-
ous, with strongly differentiated calyx and corolla;
sepals 3, equal, free, green, valvate to subvalvate;
petals 3, equal free, white, imbricate, short-clawed;
stamens 3, alternate with the petals; filaments
slender, glabrous; anthers basifixed, tetraspo-
rangiate to bisporangiate, opening by apical pores
or porelike slits or sometimes by pores at the end
of a tubular apical appendage. Ovary unilocular,
composed of 3 united carpels, placentation pari-
etal; style simple, terminal; stigma short capitate
or slightly trifid. Ovules several to numerous,
biseriate on each of the 3 parietal placentas
atropous, bitegmic, tenuinucellate. Fruit a loculi-
cidal capsule with 3 lines of dehiscence, each
midway between a placenta. Seeds ovoid to glo-
bose, striate, operculate; embryo small, globose,
undifferentiated, apical; endosperm copious,
composed of starch and aleurone. x = 8.
One genus with four to ten spp., one native to
tropical West Africa, the others to tropical and
warm-temperate America.
VEGETATIVE ANATOMY AND MORPHOLOGY.
Plants of Mayaca are rooted aquatics with adven-
titious roots (Fig. 77), containing septate cortical
air canals, produced randomly at the nodes
(Uphof 1924). Root hairs develop from specialized
rhizodermal cells and lateral roots develop oppo-
site the protophloem poles. Roots, stems, and
leaves contain vessels with scalariform perfora-
tions. Sieve-tube plastids are of the P2c type. The
slender stems have a stele composed of only 3 to 5
collateral vascular bundles arranged in a single
ring which is separated by an endodermis with
Casparian strips from an aerenchymatous cortex
composed of septate canals. Leaves are arranged
in 3/8 or 5/13 phyllotactic spiral. The leaf axils
bear short, uniseriate, ephemeral, glandular hairs.
Branching is axillary with only a few axils actually
bearing buds. Ptyxis is adplicate (flat). Each leaf
has a single vascular bundle with a septate air
canal on each side and paracytic stomates
(Tomlinson 1969).
FLORAL MoRPHOLOGY AND EMBRYOLOGY. Al-
though seemingly terminal, flowers are borne in
the axils of nonvascularized sheathing bracts. Flo-
ral initiation occurs near the shoot apex and is
accompanied by suppression of vegetative growth
while the flower and fruit develop to maturity.
Flowers may occur at only a single node or at
several successive nodes. Although the anthers are
tetrasporangiate, some tri- and bisporangiate an-
thers also occur (Horn af Rantzien 1946; Thieret
1975). Often, an individual anther may be tetra-
sporangiate in its apical region and bisporangiate
basally. Anther wall development is of the
monocot type. The endothecium fails to develop
thickenings. An exothecium develops from the
persistent epidermis with the inner tangential
walls developing reticulate thickenings. The tape-
tum is secretory with uninucleate cells. Meiosis is
successive, producing isobilateral and decussate
tetrads. Pollen is sulcate; its sculpturing is finely
reticulate with columellate ultrastructure (Zavada
1983). The embryo sac is of Polygonum type.
Endosperm formation is nuclear. Embryogeny
conforms to the Onagrad type with a well-
developed hypostase. The embryo is opposite the
micropyle with the flat side facing the endosperm.
Both integuments form a micropyle with a broad
exostome and narrow endostome. The cells of the
inner integument become radially elongated to
close the micropyle and form an operculum with-
out a micropylar collar. The cotyledon of the seed-
ling consists of a closed sheath and distal
haustoriallimb.
AFFINITIES. The relationship of the Mayacaceae
has generally been thought to be with the Com-
melinaceae (Hutchinson 1973; Cronquist 1981;
Takhtajan 1997), although both Cronquist and
Takhtajan point out evidence for possible affinity
with the Xyridaceae. The latter is supported
particularly by embryological evidence. Only
Xyridaceae and Mayacaceae have an exothecium,
secretory tapetum, parietal placentation, and a
seed coat and operculum formed primarily from
an inner integument, whereas the Commelinaceae
have an endothecium, usually an amoeboid tape-
tum, and a seed coat and operculum formed pri-
marily from the outer integument. In fact, the

presence of these unique shared embryological
characters led Stevenson (1983) and Johri et al.
(1992) to suggest that Mayacaceae should be
placed nearer to Xyridaceae than to Commelina-
ceae. The results presented in Stevenson and
Loconte (1995) also support this conclusion and
expand the number of characters to include veg-
etative features. These are the absence of rap hides,
calcium oxalate crystals, and silica as well as
nonsheathing leaves, lateral roots arising opposite
the protophloem poles, and the presence of sca-
lariform vessels throughout the plant.
DISTRIBUTION AND HABITATS. The Mayacaceae
exhibit a disjunct distribution pattern with one
species, M. baumii Gurke, occurring in Zaire,
Angola, and Zambia in West Africa. The other
three to nine species are New World and wide-
spread, ranging from theSE United States through
the West Indies and Central America to Paraguay.
Plants of Mayaca are rooted in the substrate and
often submerged (at least seasonally). The seeds
are resistant to desiccation and hence they can
grow along river banks and lake margins that
experience seasonal drying. They generally grow
on depleted sandy soils in the absence of other
vegetation, indicating that they do not do well in
competition.
Only one genus:
Mayaca Aubl.
Mayaca Aubl., Hist. pl. Guiane 1: 42, Tab. 15 (1775).
Mayacaceae 295
Fig. 77A-J. Mayacaceae. A Mayaca longipes, flowering shoot.
B-D M. fluviatilis. B Flower. C Lateral view of stamen. D Seed
in longitudinal section, endosperm stippled, embryo un-
shaded, apically the operculum. E M. se/lowiana, adaxial view
of stamen. F, G M. baumii. F Young stamen. G Dehisced sta-
men, note apical dehiscence. H-J M. vandellii. H Capsule. I
Dehisced capsule with one seed and remains of placentae on
valves. J Seed, the micropyle up. (Takhtajan 1982}
Characters as for the family.
Selected Bibliography
Cronquist, A. 1981. See general references.
Dahlgren, R. et a!. 1985. See general references.
Faden, R. 1985. Mayacaceae. In: Dahlgren, R., Clifford, T., Yeo,
P. (eds.) The families of the monocotyledons. Berlin,
Heidelbery, New York: Springer, pp. 387- 388.
Horn af Rantzien, H. 1946. Notes on the Mayacaceae of the
Regnellian Herbarium in the Riksmuseum Stockholm.
Sven. Bot. Tidskr. 40: 405-424.
Hutchinson, J. 1973. The families of flowering plants. Oxford:
Oxford Univ. Press.
Johri, B., Ambegaokar, K., Srivastra, P. 1992. Comparative
embryology of angiosperms Vol. 2. Berlin, Heidelbey, New
York: Springer.
Lourteig, A. 1952. Mayacaceae. Notul. Syst. (Paris) 14: 234-
248.
Stevenson, D. 1983. Systematic implications of the floral mor-
phology of the Mayacaceae. Am. J. Bot. 70 (5, pt. 2}: 32.
Stevenson, D., Loconte, H. 1995. Cladistic analysis of monocot
families. In: Rudall, P., Cribb, P., Cutler, D., Humphries, C.
(eds.) Monocotyledons: systematics and evolution. Royal
Botanic Gardens Kew, pp. 543-578.
Takhtajan, A.L. 1982. See general references.
Fig. 77 Takhtajan, A. 1997. See general references.
Thieret, J.W. 1975. The Mayacaceae in the southeastern
United States. J. Arnold Arbor. 56: 248-255.
296 11ayacaceae/11usaceae
Tomlinson, P.B. 1969. Commelinales-Zingiberales. In: 11et-
calfe, C.R. (ed.) Anatomy of the monocotyledons. Oxford:
Oxford Univ. Press.
Uphof, J.C.T. 1924. The physiological anatomy of Mayaca
fluviatilis. Ann. Bot. 38: 389-393.
Venturelli, 11., Bouman, F. 1986. Embryology and seed devel-
opment in Mayaca fluviatilis (11ayacaceae). Acta. Bot.
Neerl. 35: 497-516.
Vyshenskaya, T.D. 1985. 11ayacaceae. In: Takhtajan, A. (ed.)
Comparative seed anatomy. 1: 177-179. (in Russian.).
Leningrad: Nauka.
Zavada, 11. 1983. Comparative morphology of monocot pollen
and evolutionary trends of apertures and wall structures.
Bot. Rev. 49: 331-379.
Musaceae
L. ANDERSSON
11usaceae Juss., Gen. Pl.: 61 (1789}, nom. cons.
Mostly tall and robust, suckering or monocarpic
herbs with gigantic leaves; stem massive, cormlike;
leaf sheaths forming pseudostem around scape;
lateral buds absent or leaf-opposed. Leaves alter-
nate, spirally arranged, differentiated into sheath,
petiole and blade; sheath without distinct ligule;
blade entire (but often split due to wind action),
with coarse, compound midrib and closely set,
parallel, slightly sigmoid lateral veins fusing near
margin; lateral veins interconnected by tertiary
cross-veins. Inflorescence a terminal thyrse, bear-
ing cincinnate flower clusters in axils of
spathaceous bracts on an indeterminate main axis.
Flowers ebracteolate, functionally unisexual with
male flowers in distal part of inflorescence and
female ones in proximal part (rarely bisexual
flowers in proximal part), trimerous and 5-cyclic,
heterochlamydeous, zygomorphic; unpaired outer
tepal in anterior (abaxial) position; outer tepals
fused mutually and with 2 inner tepals into 5-
dentate or 5-lobate lamina; adaxial inner tepal free,
small; stamens usually 5, the missing stamen then
mostly staminodial, or occasionally 6; filaments
free from each other, narrow and filiform; anthers
basifixed, elongate, 2-thecate, 4-sporangiate.
Ovary inferior, 3-locular; ovules anatropous, nu-
merous on axile placentae; style 1, filiform, often
dilated distally. Fruit a baccate, mostly indehiscent
capsule; seeds operculate, exarillate, with scanty
perisperm and copious starchy endosperm.
Two genera and about 40 spp.; tropical Africa
and from NE India to N Australia and the
Philippines.
VEGETATIVE MORPHOLOGY. Various aspects of
vegetative morphology have been described by
Fisher (1978), Kaplan (1973) and Tomlinson
(1962).
In Musa, the plants have a sympodial rhizome
with branches terminating in aerial shoots, and
with renewal branches arising near the base of the
aerial shoot (Tomlinson 1962). Lateral branches
arise from leaf-opposed buds (Fisher 1978). Ensete
is strictly monocarpic, with vegetative branching
 Musaceae 297

occurring only as a result of damage. Rhizome with vascular bundles in all parts of the plant
branches are usually short and massive, becoming except roots.
cormlike at soil surface. Leaves are produced from
the "corm" tip. The short rhizome branches typi- INFLORESCENCE. The inflorescence is terminal,
cally give the plants a caespitose habit. Starch is its formation terminating the growth of the entire
commonly stored in the ground parenchyma of aerial shoot. (Fig. 78).
the rhizome (Tomlinson 1959).
The only aerial stem is the inflorescence scape. It
is soft-textured and depends on support from the
tightly enclosing leaf sheaths, which form a
pseudostem.
The leaf is usually differentiated into sheath,
petiole and blade, but a distinct petiole may be
missing. All divisions of the leaf are derived from
the proximal (Unterblatt) part of the primordium,
the distal (Oberblatt) part giving rise only to a
precursory tip, which is usually not indentifiable
in the mature leaf (Kaplan 1973). Ptyxis is
supervolute.
The leaf blade has a massive, compound midrib
from which closely set, parallel secondary veins
diverge. There have a slightly sigmoid course and
fuse near the leaf margin. They are interconnected
by parallel tertiary veins, which are perpendicular
to the secondary ones.

VEGETATIVE ANATOMY. Vegetative antomy was

described in detail by Tomlinson (1959). Stoma
development was described by Tomlinson (1974)
and Olatunji (1980).
Root stele with wide vessels and phloem strands
scattered throughout the stelar ground tissue.
Hairs absent from aerial parts. Rhizomes with the
central cylinder demarcated by an endodermis.
Leaf axis with a main arc of vascular strands,
which are continuous with the major lateral veins
of the lamina, with several arcs of smaller and
shorter strands, some entirely fibrous, on the
abaxial side of the main arc, and with small vascu-
lar and fibre bundles close to the adaxial surface;
with only 1 arc of large air canals. Epidermis of
lamina with anticlinal walls non-sinous; stomata
predominantly paracytic, guard cells symmetrical
in cross-section; hypodermis usually of 2 or more
layers on adaxial side, 1-2-layered abaxially; me-
sophyll with transverse, vascularised or non-
vascularised septa; all veins sheathed by fibres,
sheaths connected to hypodermis on both sides.
Silica cells are associated with vascular bundles in
all parts of the plant except the roots, silica bodies Fig. 78A-J. Musaceae. A-F Musa acuminata. A Inflorescence.
through-shaped. Raphide sacs are present in the B Distal part of inflorescence, the reflexed spathe showing a
ground tissue of all organs. Tannin present in staminate partial inflorescence. C Portion of inflorescence axis
with scars of the spathe and fruits. D Staminate flower. E
morphologically unspecialised cells in all parts of Pistillate flower. F Seed. G M. maclayi, inflorescence. H-J
the plant. Articulate lacticifers containing a muci- Ensete ventricosum. H Inflorescence. I Fruit. J Seed.
laginous substance present in M usa, associated (Takhtajan 1982)
298 Musaceae

The main inflorescence axis is indeterminate. It trichomes is similar to those of arils in other
bears spathaceous bracts (spathes), in the axils of Zingiberales.
which partial inflorescences (hands) arise. The
spathes are spirally arranged and caducous EMBRYOLOGY. The embryology of Musa was ex-
(Musa) or persistent (Ensete). The flowers are amined by Bouharmont (1963) and Simmonds
ebracteolate. The partial inflorescences in the (1960, summarised in Simmonds 1962, 1966).
proximal part of the inflorescence have female (or Some details are also provided by Humphrey
rarely bisexual) flowers, those in the distal part (1896).
have male ones. The ovule is anatropous, with 2 integuments, the
The partial inflorescences have been variously outer with 6-9, and the inner one with 2 cell layers.
interpreted. The classical (and still most com- The micropyle is formed by the outer integument.
monly accepted) interpretation, suggested by The nucellus is crassinucellate. Megasporogenesis
Eichler (1875), is that it is a cincinnus. The whole follows the Polygonum type. No parietal cell
inflorescence is then a thyrse with monchasial par- seems to be formed. A circular thickening, the so-
tial inflorescences. Thompson ( 1933) and Lane called Mikropylarkragen is formed from the outer
(1955), however, considered the partial inflores- integument. The epidermis of the nucellus is
cences to be deformed spikes. The classical view palisadelike at the stage when the embryo sac is
has been supported by Fahn (1953), who con- mature. Endosperm formation is Nuclear.
cluded that Thompson's interpretation was based The first division of the zygote is transverse or
on erroneous observations. oblique in Musa (Bouharmont 1963), the plans of
the second division being perpendicular to that of
FLORAL STRUCTURE AND ANATOMY. The flowers the first one. The young embryo (25 days) is still
have a basic trimerous, pentacyclic organization, undifferentiated and ± globose, a suspensor de-
with 2 whorls of tepals, 2 of stamens, and 1 of veloping at a later stage (ca. 60 days), at which the
carpels. They are oriented with the unpaired proembryo is cylindrical and the meristems differ-
sepal in the anterior position. The perianth entiating into radicula, plumule and cotyledon are
consists of a large, 5-lobed or 5-dentate structure, vaguely defined.
and a small simple, scalelike one. The larger
structure is usually interpreted as having evolved POLLEN MORPHOLOGY. Pollen grains are large or
by fusion of all 3 outer and the 2 paired inner very large (to 140~-tm diam. in Musa), radiosym-
tepals, which is supported by the course of metric and isopolar. There is no structurally
vascular strands (Tilak 1968; Fahn and Kotler defined aperture. The surface is psilate in Musa,
1972; Tilak and Pai 1974) and by observations of but has usually large sporopollenin gemmae
occasional structural aberrants (Varela de Vega or warts in Ensete. The exine is endexinous in
1972). nature, uniformly thin (0.02-0.06~-tm in Musa) or
Musa usually has 5 stamens with a staminode irregularly thickened (0.02-1.1~-tm, in Ensete). In
present or absent. Occasional aberrants have 6 Musa, it is made up of several thin layers, some of
fertile stamens (Varela de Vega 1972). The which may slough off during development. The
staminode, when present, belongs to the inner intine is thick. In Musa the exintine has an outer
whorl. layer that is granular or has elliptic channels and
The gynoecium is formed by 3 carpels. The an inner, channelled layer. The endintine is made
ovary is inferior and trilocular, with numerous up of longitudinally oriented fibrils. The exintine
ovules on axile placentae. Septal nectaries are in Ensete is channelled throughout and the
present in the axile tissue of the distal part (pro- endintine is similar to that of Musa (W.J. Kress,
longation) of the ovary, descending as far down as pers. comm.).
the uppermost part of the locular region (Kirchoff
1992). The nectary duct is 6-armed proximally, KARYOLOGY. Detailed reviews of karyology are
becoming 3-radiate distally, ending with 3 sepa- given by Simmonds (1962, 1966). Wild species of
rate ducts below the style. In Musa, trichomes are Musa have 2n = 14, 18, 20 and 22, different basic
present on the raised parts of the placentae and numbers being characteristic for different sec-
sometimes on the funicle. These trichomes have tions. Polyploidy is important in banana cultivars,
been interpreted as aril homologues (e.g. but is unknown for wild material. The numbers 2n
Friedrich and Strauch 1975). This interpretation = 18 and 20 have been reported for Ensete. Most
was rejected by Kirchoff (1992) on the grounds chromosomes have satellites (Cheesman and
that neither the position, nor the structure of these Larter 1935; Bisson et al. 1968).
 Musaceae 299

POLLINATION AND REPRODUCTIVE SYSTEM. Strelitziaceae. At present (e.g. Tomlinson 1962;

Musa and Ensete are primarily pollinated by Cronquist 1981; Dahlgren et al. 1985), the ten-
macroglossine bats, but pollination by sunbirds dency is to include only Musa and Ensete in the
(Nectariniidae) and tree shrews (Tupaia spp.) Musaceae.
have also been reported (Nur 1976). In a phylogenetic analysis of the Zingiberales
Musa and Ensete are usually monoecious with (Kress 1990) the Musaceae occupy a basal
male flowers in the distal part and female flowers position. They differ from all other families
in the proximal part of the inflorescence. Selfing of the order in possessing laticifers, spiral
must, therefore, be very rare in natural popula- phyllotaxy, unisexual flowers, and in lacking
tions. Parthenocarpy is not infrequent even in arils.
wild plants, however (Simmonds 1962), and self-
compatible hermaphrodite flowers have been DISTRIBUTION. The Musaceae have most extant
observed (Nur 1976). species in Asia, NE Australia and the western Pa-
cific, but three species of Ensete are African. The
FRUIT AND SEED. The fruit is indehiscent, or, centres of diversity are in Assam-Burma and
rarely, dehiscent by decay. The pericarp is leathery Indonesia-Borneo.
and the seeds are surronded by a starchy, sweetish
pulp derived from the placental trichomes PALAEOBOTANY. Spirematospermum, a fossil seed
(Friedrich and Strauch 1975). The evolution of known from the Upper Cretaceous of North
seedless, edible bananas is described in detail by America and frequent in the European Tertiary,
Simmonds (1962, 1966). for long had been related to the Zingiberaceae, but
Seeds of Ensete and Musa were studied by Desler Rodriguez de la Rosa and Cevallos-Ferriz (1994)
(1994). They are generally large (ca. 5-15mm recognised it as musaceous. Manchester and Kress
diam.), exarillate, and have an operculate germi- (1993) reported seeds of Ensete from the Eocenic
nation pore in the micropylar end. A double mi- Clarno flora of Oregon.
cropylar collar and a distinct hypostase are
present in the mature seed. The outer integument ECONOMIC IMPORTANCE. The most important
forms a massive sclerenchyma in the mature testa. economic product of the Musaceae is the edible
The inner integument is 2-layered, with distinct banana, which is, after citrus, the most important
cellular structure. In Musa, the nucellus is fruit in world trade. Bananas are commonly grown
represented by 1 cell layer in which a fatty in monocultures in the tropical lowlands around
substance but no starch is stored. In Ensete, the the world, but are also, together with plantains,
nucellus remnant is strongly compressed. Starch cultivated in backyard gardens or in mixed crop-
is stored in the endosperm in the form of large, ping systems, sometimes as shade canopy in cof-
simple, concentric grains. The embryo is fee plantations. The dessert bananas are sugary
mushroom-shaped. and most often eaten raw. Plantains are generally
more starchy and are cooked, fried or roasted. In
DISPERSAL. The fruits are taken by mammals many tropical countries, especially in Africa and
(bats, monkeys and rodents) and birds. The hard SE Asia, bananas and plantains are the staple food.
seeds are later dropped, regurgitated or passed They are also dried, powdered, chipped, flaked,
through the digestive tract. juiced and pureed as well as used for producing
beer.
SUBDIVISION AND RELATIONSHIPS. The close The male buds of M usa inflorescences are
relationship of Musa and Ensete has never cooked as vegetables in some parts of Asia. The
been doubted; Ensete has, in fact, often been in- swollen basal parts of leaf sheaths of Ensete
cluded in Musa (e.g. Schumann 1900; Winkler ventricosum are used as a source of starch in east-
1930). Traditionally (e.g. Schumann 1900; Winkler ern Africa. Musa leaves are widely used to wrap
1930), they have been included in a broadly cir- food items when cooking.
cumscribed family Musaceae together with Musa textilis is cultivated, mainly in the Philip-
Strelitzia, Ravenala, Phenakospermum, Heli- pines and other South Pacific islands, for its strong
conia and often also Orchidantha. Orchidantha leaf fibers (Manila hemp), used to make ropes and
was separated by Ridley (1924) in a family of coarse textiles. Leaves of the Musaceae are also
its own, the Lowiaceae. Hutchinson (1934) se- widely used for thatch.
gregated Strelitzia, Ravenala, Phenakosper- Cultivars of both Musa and Ensete are often
mum and Heliconia as a separate family, the used in ornamental plantations.
300 Musaceae
KEY TO THE GENERA
1. Plants suckering; spathes caducous; free (inner) tepa!
simple 1. Musa
- Plants monocarpic; spathes persistent; free (inner) tepa!
3-lobed 2. Ensete
1. Musa L. Fig. 78A-G
Musa L., Sp. Pl.: 1043 (1753); Argent, Notes R. Bot. Gard.
Edinb. 35: 77-114 (1976), rev.
Suckering herbs, often growing in dense clumps;
lateral buds leaf-opposed. Bracts caducous;
flowers and bracts separately inserted on axis.
Flowers of proximal cymes usually functionally
female. Tepal lamina 5-toothed, not deeply di-
vided. Seeds usually ca. 9 mm diam. Chromo-
somes number: x = 7, 9, 10, 11. Thirty to 40 spp.,
from the Himalayas and S China to N Australia
and the Philippines; occurrence further east into
the Pacific (Pemba, Samoa and Hawaii) supposed
to be due to introductions.
The genus is often divided into five sections
(some species still with uncertain placement); (1)
Sect. Musfil, x = 11, robust plants with pendent
inflorescence, dull and waxy bracts, fruits never
orange-red; 13-15 spp. (2) Sect. Rhodochlamys,
x = 11, slender plants, with erect inflorescence,
opaque and mostly waxy, brightly coloured bracts;
5-7 spp. (3) Sect. Ingentimusa, x = 7, tall plants
with pendent inflorescence and dull cream bracts;
only one sp., M. ingens. (4) Sect. Australimusa, x
= 10, inflorescence usually horizontal to pendent,
bracts usually shiny, seeds subglobose to lenticu-
lar; 5-7 spp. (5) Sect. Callimusa, x = 10, inflores-
cence usually erect, bracts usually shiny, seeds
barrel- or top-shaped; 6-10 spp. The cultivated
bananas have a complex history amply reviewed
by Simmonds (1962, 1966). Obviously, they have a
multiple origin in SE Asia from where they have
been spread by man, both east- and westwards.
Two series of cultivars are discerned. All commer-
cially grown bananas belong to sect. M usa. Most
of them are triploids and derived either from
M. acuminata alone or from hybrids with M.
balbisiana. Diploid cultivars are grown locally in
India and Malaysia. The second series of cultivars
consists of the so-called Fe'i bananas. These be-
long in sect. Australimusa and lack commercial
importance. They are grown locally in E Indone-
sia, New Guinea and the Pacific.
2. Ensete Horan. Fig. 78H-K
Ensete Horan., Prodr. Scitam.: 40 (1862); Cheesman, Kew Bull.
1947: 97-106 (1947), rev.
Physocaulis Baker (1893).
Gigant, monocarpic herbs. Lateral (adventitious)
buds developing only on damage. Bracts and
flowers integral with each other and with axis, ab-
scission layer lacking. Flowers of proximal cymes
predominantly bisexual; the 5 fused tepals divided
nearly to the base. Seeds usually > 10 mm diam. 2n
= 18. Six spp., three in Central Africa from the
Cameroons to Ethiopia and Transvaal, three in SE
Asia from NE India and Burma through China to
the Philippines, New Guinea, and Java.
Selected Bibliography
Bisson, S., Guillemet, S., Hamel, J.-L. 1968. Contribution a
!'etude caryo-taxonomique des Scitaminees. Mem. Mus.
Hist. Nat., II B, 18:59-145, pl.1-19.
Bouharmont, J. 1963. Evolution de !'ovule feconde chez Musa
acuminata Colla subsp. burmannica Simmonds. Cellule 63:
262-279, pl. 1-4.
Cheesman, E.E., Larter, L.N.H. 1935. Genetical and cytological
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Musaceae. J. Genet. 30: 31-52.
Cronquist, A. 1981. See general references.
Dahlgren, R. et al. 1985. See general references.
Desler, B. 1994. Vergleichende Untersuchung der Anatomie
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Diploma Thesis. Kaiserslautern: Faculty of Biology, Univer-
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Eichler, A.W. 1875. Bllithendiagramme construiert und
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Fahn, A. 1953. The origin of the banana inflorescence. Kew
Bull. 1953: 299-306.
Fahn, A., Kotler, M. 1972. The development of the flower and
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1972: 153-169.
Fisher, J.B. 1978. Leaf-opposed buds in Musa: their develop-
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Friedrich, W.L., Strauch, F. 1975. Der Arillus der Gattung
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Humphrey, J.E. 1896. The development of the seed in the
Scitamineae. Ann. Bot. 10: 1-40.
Hutchinson, J. 1934. Families of flowering plants, 2nd edn.,
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Kaplan, D.R. 1973. The problem of leaf morphology and evo-
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2490-2508.
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 Musaceae/Najadaceae
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301
Najadaceae
R.R. HAYNES, L.B. HoLM-NIELSEN, and
D.H. LES
Najadaceae Juss., Gen. Pl.: 18 (1789}, nom. cons.
Monoecious or rarely dioecious, annual or rarely
perennial, erect, caulescent, glabrous herbs of
fresh or brackish waters, growing entirely sub-
mersed. Roots fibrous, few, nonseptate, at base of
stem and lower nodes. Stems slender, much
branched, sometimes armed with spinulose teeth
on the internodes; turions and tubers never devel-
oped. Leaves spiral, subopposite or pseudover-
ticillate, sessile, with sheathing base; blade linear,
flattened, 1-veined, sometimes abaxially toothed
on midrib, the margins serrulate to dentate, the
apex acute to acuminate, with 1-3 teeth per side;
the teeth multicellular, formed by layers of cells
decreasing in cell number distally and terminated
by a large, sharp-tipped cell, or unicellular;
sheaths variously shaped, each enclosing a pair
of tiny intravaginal squamules, the margins
usually spiny-toothed, with 1-15 teeth per side.
Flowers axillary, solitary, or clustered, imperfect,
perianth absent. Staminate flowers mostly
subtended by 1 or 2 clear to pale purple, mem-
branous, spathe-like envelopes, each flower
consisting of a single stamen; pedicels at first
short, elongating at anthesis, pushing the anther
through the outer envelope; anther sessile, 1- or
4-sporangiate, dehiscing by an apical slit.
Carpellate flowers sessile; a clear to pale purple
membranous envelope rarely present; gynoecium
ending in a short style with 3 or 4 branches;
ovary 1-loculed; wall delicate, 2 cell layers thick;
placentation basal; ovule solitary, bitegmic, anat-
ropous. Fruit achenelike; gynoecial wall closely
enveloping the seed, opening by decay. Seed fusi-
form to elliptic; testa areolate; embryo elongate,
macropodous, straight or rarely curved; en-
dosperm developing nuclear, absent in the mature
seed.
A monogeneric, nearly cosmopolitan family
with ca. 40 species.
VEGETATIVE MORPHOLOGY. The Najadaceae are
all aquatic herbs which grow entirely submersed
in fresh or brackish waters (Fig. 79). The roots
arise from the base of the stem or along the lower
302 Najadaceae

Fig. 79A-G. Najadaceae. Najas podostemon. A Habit. B Leaf. except that the margins and apices are lined with
C Enlarged leaf apex. D Leaf axils with immature staminate prickle hairs. The prickles are composed either
flower in axil on left, mature staminate flower after release of
pollen in lower right axil, mature ovulate flower in middle of only one enlarged yellowish to brown cell ( 1-
right axil, and immature ovulate flower in upper right axil. cellular) (Fig. 79C), or this may be situated on a
E Transverse section of anther. F Seed. G Enlargement of base formed by layers of smaller cells, which de-
exotesta with 4-6-sided areolae. (Orig.) crease distally in cell number (multicellular).
Leaves with unicellular prickles (or teeth) mostly
appear entire to the unaided eye but normally
nodes, especially those nodes that contact the have 50 or more marginal teeth, 2 or 3 at the leaf
bottom. apex. Leaves with multicellular teeth appear ser-
The stem varies in length up to 1m, depending rate to coarsely dentate to the unaided eye and
on water depth. It is usually highly branched, ei- mostly have 25 or fewer teeth along each margin
ther near the apex or near the base. When branch- and only 1 or rarely 2 at the apex. Occasionally,
ing is from the base, the lateral branches normally those with multicellular teeth also have prickles
grow long and erect, taking on the appearance of along the lower surface of the midvein and more
the main stern. The lateral branches and main rarely along the upper surface.
stern often branch profusely near the apex. This The sheath is wider than the blade and truncate,
branching along with reduced internode length rounded or sloping, rarely auricled. The margins
gives the distal part of the plant a bushy appear- have 1-15 teeth similar to those of the blade. There
ance. In subgen. Najas, the internodes normally are 2 intravaginal squamules, which are mostly
have prickles; these are absent in subgen. subulate with entire margins but vary in shape,
Caulinia. being occasionally nearly oval. They are 3 cell
The leaves are subopposite on the stern, 1 leaf of layers thick and lack vascularization. They pre-
each pair being slightly lower than the other. sumably secrete mucilage which may prevent the
These subopposite pairs are essentially spiro- growth of microorganisms.
distichous. Leaf arrangement often appears to be
whorled due to a short-shoot, which arises in the VEGETATIVE ANATOMY. Raphides and silica bod-
axil of the lower leaf of each pair of cauline leaves. ies are lacking. Vessels are completely lacking.
The short-shoot consists of only 2 leaves, a lower Tannin cells and crystals are little developed.
one which is reduced to a scale and an upper The anatomy is fairly simple and has been
which develops a blade. An axillary bud is present summarized by Tomlinson (1982). The roots
only in the upper of these leaves. The 3 foliose are copiously supplied with root hairs, which
leaves, the 2 stem leaves and 1 short-shoot leaf are produced only by specialized short, densely
make up the pseudowhorl. cytoplasmatic cells of the piliferous layer.
The leaf blade is flat, linear to linear-lanceolate, In the stem, stomata are absent and the cuticle
and tapers into an acute apex. It is glabrous, is thin. The epidermis contains chlorophyll. The
 Najadaceae
cortex is fairly wide, including a 1- or 2-layered
hypodermis of compact cells, a middle lacunose
region with several enlarged longitudinally ex-
tended intercellular air spaces forming a single
series of air canals, and a compact innermost layer
with slightly thickened walls. The endodermis en-
circling the stele is composed of cells with thin
walls or rarely with slightly thicker outer walls in
the older stems. The stele is composed of a fairly
large protoxylem lacuna surrounded by a single
layer of parenchyma cells and a cylinder of ph-
loem, which is often in contact with the endoder-
mis. The vascular and cortical tissues are poorly
developed. The phloem has sieve plates in all parts.
The leaves are mostly composed of two epider-
mal layers with occasional narrow intercellular
spaces. A few species, e.g., N. gracillima, have 1 or
rarely more layer(s) of mesophyll between the epi-
dermallayers. The epidermal cells contain chloro-
phyll; stomata are lacking. Lacunae are mostly
restricted to the midrib area, except that those
species with a layer of mesophyll often have some
intercellular spaces. The midvein consists of a
single vascular bundle, which is surrounded by
mesophyll tissue. The vascular tissue is reduced,
with the tracheids mostly replaced by protoxylem
lacunae.
INFLORESCENCE STRUCTURE. The inflorescence
consists of a solitary flower produced in the axils
of a pseudowhorl. The axillary meristem, which
gives rise to the lateral branch of the pseudowhorl,
divides laterally early in its inception, one side
producing the vegetative branch. Occasionally,
there are 2-several flowers in the axils of a
pseudowhorl. The additional flowers are pro-
duced by the axillary meristem of the short-shoot
of the pseudowhorl repeating the lateral bifurca-
tion which gave rise to the lateral branch origi-
nally. This process may be repeated until up to 5
flowers per axil are produced (de Wilde 1962).
FLORAL STRUCTURE. The flowers of the Najada-
ceae are so reduced that it is difficult to determine
the nature as well as the number of floral parts.
Both staminate and carpellate flowers are usually
enclosed by 1 or 2 translucent floral envelopes
(Fig. 79D). The nature of these envelopes is not
fully understood. Tomlinson (1982) referred to
the inner one as the perianth and the outer one as
the spathe. Posluszny and Sattler (1976), however,
referred to them as floral envelopes since they
are initiated as girdling primordia on the floral
axis after the initiation of the reproductive
structures.
303
The staminate flower is subsessile and com-
posed of one stamen, which is almost always sur-
rounded by 2 floral envelopes. The outer floral
envelope is initiated after initiation of the stamen
and elongates more rapidly than does the floral
apex. When the outer envelope surpasses the
floral apex, the inner envelope is initiated. Both
envelopes overgrow the stamen and form a short
neck at the apex of the flower. The inner envelope
adheres to the stamen so closely as to become
invisible except for the small bilobed neck which
extends past the stamen. The neck of the outer
envelope extends past that of the inner and is ir-
regularly 3- to 5-toothed. The flower stalk begins
to elongate at the time the pollen is mature and
pushes the anther upwards, irregularly rupturing
the floral envelopes and then becomes slightly
recurved. The anther dehisces by an apical pore
or slit, releasing the pollen into the surrounding
water.
There have been several reports in the literature
that the staminate flowers are in the upper
axils and the carpellate flowers are in the lower.
Developmental data by Posluszny and Sattler
(1976), however, indicate that there is no apparent
pattern to the arrangement of the flowers on a
plant.
The carpellate flowers are sessile and apparently
composed of one ovule surrounded by a transpar-
ent gynoecial wall. Posluszny and Sattler (1976)
have demonstrated that the translucent floral
envelope on carpellate flowers is actually the
gynoecial wall. This is initiated in a manner
similar to that of the outer floral envelope of the
staminate flowers. During floral development,
the ovule is initiated in a terminal position on the
floral apex, followed by the initiation of the
gynoecial wall. This overgrows the developing
ovule and remains 2 cell layers thick throughout
its development. The apex narrows into a short
neck or style with 2 stigmas. Often these stigmas
divide to form 4lobes. The 2 integuments are ini-
tiated after the gynoecial wall overgrows the floral
apex. Once the integuments are initiated, the
ovule turns down into an anatropous position.
The exact terminal position of the ovule
during development led Posluszny and Sattler
(1976) to the conclusion that Najas is acarpellate.
This concept, however, has not received wide
acceptance.
EMBRYOLOGY. The anther wall formation is of the
Reduced type (Davis 1966) in which only 2 parietal
layers are formed. The fully developed anther wall
is composed of only the endothecium, which lacks
304 Najadaceae
fibrous thickenings and is soon crushed. The
tapetum soon becomes amoeboid. The tapetal
cells remain 1-nucleate. Their walls eventually
break up and a periplasmodium is formed
(Venkatesh 1956).
Microsporogenesis is of the Successive type. The
tetrads are tetrahedral or isobilateral, rarely linear
or T-shaped. These varying types often occur in
the same species (Venkatesh 1956). The pollen
grains are 2-celled when the tetrads break up, but
become 3-celled shortly thereafter. The mature
pollen grains are filled with starch grains.
The ovules are always anatropous and bitegmic.
A parietal cell is formed; this divides to form 1 or
2 parietal layers below the nucellar epidermis
(Davis 1966). The micropyle is formed by the
inner integument. Embryo-sac formation follows
the Polygonum type. Endosperm formation is
nuclear, but walls are initiated later near the mi-
cropylar pole, and the tissue becomes cellular
throughout.
Embryogeny follows the Caryophyllad type, in
which the basal cell does not divide but enlarges
as the terminal cell of the short suspensor.
The ripe seeds lack endosperm and perisperm.
The embryo is cylindrical, linear, macropodous,
tapering at each end and mostly straight, although
a few species have slightly curved embryos. The
embryo fills the entire seed cavity. It has 1 or 2
bands of chlorophyll.
POLLEN MORPHOLOGY. The pollen grains are
globose to ellipsoid and inaperturate monads. The
exine is without sculpturing, or occasionally has
shallow reticulations, and it so thin that it has been
observed only with transmission electrom micros-
copy, and then only with certain staining proce-
dures (Pettitt and Jermy 1975). In contrast, the
intine is fairly thick.
KARYOLOGY. Chromosome numbers are reported
for 15 species of Najas. These records include 2n
= 12, 14, 16, 24, 34, 36, 42, 46, 48, 54, and 60.
Viinikka (1973) reported the presence of B chro-
mosomes in nine European populations of Najas
marina.
PoLLINATION. According to Cox (1985), the
pollen germinates in the anther prior to dehis-
cence. The long tube plus the pollen grain allows a
much larger surface with which to contact a
stigma as the pollen moves through the water.
Rendle (1899) indicated the pollen grains fall,
weighted by their starch contents, onto stigmas of
carpellate flowers lower in the water column.
FRUIT AND SEED. The fruit of Najadaceae does
not fit the classical definition of any fruit type. The
fruit never actually dries; its wall adheres closely
to the seed and dehisces by decay.
The seeds are quite variable in the family. The
exotesta may be deeply or more rarely shallowly
pitted with areolae (Fig. 79F,G). These are ar-
ranged in 20-50 longitudinal rows and vary in
shape from isodiametric to rectangular and if rect-
angular are usually longer than wide, rarely wider
than long. They are separated by walls that usually
are slightly raised from the testa surface. The junc-
tures of these walls are often raised additionally
into a papilla, giving the exotesta a papillose ap-
pearance. Finally, a few species are so deeply
pitted that centers of the areolae seem to form a
funnel.
The mesotesta consists of stone cells that
provide the seeds with mechanical strength. Dif-
ferences in the expression of this tissue have been
used in the classification of Najas. In N. marina
(subgen. Najas) the mesotesta is 10-15 cell layers
deep, while in subgen. Caulinia it is composed of
1 or 2 layers. The inner layer of the tegmen
(endotegmen) is provided with minute tubercles
(endotegmen tuberculae). Najas shares these
features with at least 11 genera of the Hydro-
charitaceae (the genera of the Limnanthemum
group, the Vallisneria group, some of the Elodea
group and Enhalus and Blyxa; see Hydrocharita-
ceae account), and these endotegmen tuberculae
are unknown elsewhere in flowering plants
(Shaffer-Fehre 1991a,b).
Starch grains are not accumulated in the seed.
DISPERSAL. Dispersal mechanisms are poorly un-
derstood. The seeds mostly remain in the leaf axils
until the plant dies. The dying plants are uprooted
and usually remain suspended in the water for a
short period before sinking to the substrate.
During this time they may be transported by
currents before sinking to the bottom, where the
seeds become embedded in the substrate and
germinate.
Short-range dispersal by adhesion to birds and
other animals is likely, but long-range dispersal by
birds eating the seeds is unlikely, as the seeds are
too delicate to pass through the digestive system
undamaged.
PHYTOCHEMISTRY. Saponins and alkaloids are
absent in the family. The flavonoid spectrum,
however, is comparatively diverse. Roberts and
R.R. Haynes (unpubl.) have isolated leucoantho-
cyanidins and 9 flavonoids from the leaves and
 Najadaceae 305

stems of Najas. These include both flavones and
flavonols. Najas subgen. Najas contains only
flavones, whereas Najas subgen. Caulinia contains
both flavones and flavonols. Sulfated flavonoids
have not been observed.
1981) of both England and the former USSR.
For species of Najas subgen. Caulinia from the
Miocene and Pliocene of the former USSR, distin-
guishable by their seed-coat anatomy, see
Dorofeev (1978).

AFFINITIES. Extreme structural reduction and EcoNOMIC IMPORTANCE. The family is economi-
adaptations to water pollination have complicated cally important in three respects. As a source of
efforts to determine the phylogenetic position of food for waterfowl, Martin and Uhler (1939) rank
the genus Najas. Otherwise, a distinctive array of it eighth in importance in the United States and
features including nuclear endosperm formation, Canada, and Gaevskaya (1966) lists 18 species of
straight embryos, basal placentation, solitary animals that have been known to feed on it.
carpels, and solitary stamens have facilitated the Secondly, populations of Najas have often been
circumscription of this genus in the Alismatidae. abundant enough to be regarded as weeds,
The specialized features of Najas have inspired warranting the use of various methods of control.
most authors to recognize the genus within a sepa- Finally, Najas guadalupensis is utilized in the
rate family, Najadaceae. This family has been as- aquarium trade.
sociated phylogenetically with Scheuchzeriaceae,
Juncaginaceae, Potamogetonaceae, Posidonia- Only one genus:
ceae, Cymodoceaceae, Ruppiaceae, Zannichellia-
ceae, and Zosteraceae to comprise an order
Najas L. Fig. 79
referred to as Najadales or Zosterales (Dahlgren
and Clifford 1982; Dahlgren et al. 1985). Miki Najas L., Sp. Pl.: 1015 (1753); Rendle, Pfianzenreich IV. 12
(1937), however, had already considered that (1901); Haynes, Sida 8: 34-56 (1979), N and Central
Najas could have originated from submerged American spp.; Lowden, Aquat. Bot. 24: 147-184 (1986),
Neotrop. spp.; Triest, Mem. Acad. R. Sci. d'Outre mer, Cl.
Hydrocharitaceae. Evidence obtained from seed Sci. Nat. (8) 22: 1-172 (1988), Old World spp.
coat anatomy (see Fruit and Seed, above), and
molecular studies have now indeed established About 40 spp., characters as for family. Two sub-
that Najas is not closely related to any of the genera: subgenus Najas, dioecious, internodes
families traditionally included in Najadales or and leaf undersurface prickly, only N. marina L.;
Zosterales, but is derived from within the family subgenus Caulinia (Willd.) Aschers., monoecious,
Hydrocharitaceae (Shaffer-Fehre 1991b; Les et al. without prickles, ca. 40 species, most of which in
1993; Les and Haynes 1995; D.H. Les, unpubl.). the eastern hemisphere.
The presence of endotegmen tuberculae and a
thin exotesta represent reliable morphological
synapomorphies linking Najas to the Hydro- Selected Bibiography
charitaceae.
The Najadaceae have been maintained as a dis- Casper, S.J., Krausch, H.-D. 1980. Pteridophyta und
Anthophtya. 1. Teil: Lycopodiaceae bis Orchidaceae. In:
tinct family in this treatment at the request of the Ettl, H., Gerloff, E., Heynig, H. (eds.) Sii6wasserfiora von
Editor, regardless if such recognition renders Mitteleuropa, Band 23. New York: Gustav Fischer.
Hydrocharitaceae a paraphyletic family. The pre- Cook, C.D.K. 1990. Aquatic plant book. The Hague: SPB
ponderance of phylogenetic evidence indicates Academic Pub!.
that Najas is embedded in the Hydrocharitaceae; it Cox, P.A. 1985. Noodles on the tide: why is water-borne pollen
so long and stringy? Nat. Hist. 94: 36-41.
clearly belongs within the order Alismatales. Daghlian, C.P. 1981. A review of the fossil record of monocoty-
ledons. Bot. Rev. (Lancaster) 47: 517-555.
DISTRIBUTION AND HABITATs. The Najadaceae Dahlgren, R.M.T., Clifford, H.T. 1982. See general references.
have a subcosmopolitain distribution with most Dahlgren, R.M.T. eta!. 1985. See general references.
Davis, G.L. 1966. See general references.
species in tropical Asia and tropical Africa. They Dorofeev, P.l. 1978. On the taxonomy of the Neogene genus
are aquatic and occur especially in substrates with Caulinia Willd. Bot. Zh. 63: 1089-1101.
a high organic content. A few species grow in Gaevskaya, N.S. 1966. The role of higher aquatic plants in the
brackish waters, although most occur in fresh- nutrition of the animals of fresh-water basins. 3 vols.
water. No species are marine. (Trans!. from Russian by D.G. Maitland Muller 1969).
Yorkshire: National Lending Library for Science and
Technology.
PALEOBOTANY. The earliest records are seeds of Haynes, R.R. 1977. The Najadaceae in the Southeastern United
Najas and Najadites from the Oligocene (Daghlian States. J. Arnold Arbor. 58: 161-170.
306 Najadaceae/Palmae
Les, D.H., Haynes, R.R. 1995. Systematics of subclass
Alismatidae: a synthesis of approaches. In: Rudall, P.J.,
Cribb, P.J., D.F. Cutler, C.J. Humphries (eds.) Monocotyle-
dons: systematics and evolution. Royal Botanical Gardens,
Kew.
Les, D.H., Garvin, D.K., Wimpee, C.F. 1993. Phylogenetic
studies in the monocot subclass Alismatidae: evidence for a
reappraisal of the aquatic order Najadales. Mol. Phylogenet.
Evol. 2: 304-314.
Magnus, P. 1870. Beitrage zur Kenntnis der Gattung Najas.
Berlin: Georg Reimer.
Martin, A.C., Uhler, F.M. 1939. Food of game ducks in the
United States and Canada. U.S.D.A. Tech. Bull. No. 634.
Miki, S. 1937. The origin of Najas and Potamogeton. Bot. Mag.
(Tokyo) 51: 472-480.
Pettitt, J.M., Jermy, A.C. 1975. Pollen in hydrophilous an-
giosperms. Micron 5: 377-405.
Posluszny, U., Sattler, R. 1976. Floral development of Najas
flexilis. Can. J. Bot. 54: 1140-1151.
Rendle, A.B. 1899. A systematic revision of the genus Najas.
Trans. Linn. Soc. Lond. II. 5: 379-444.
Shaffer-Fehre, M. 199la. The endotegmen tuberculae: an
account of little-known structures from the seed coat of the
Hydrocharitoideae (Hydrocharitaceae) and Najas (Najada-
ceae). Bot. J. Linn. Soc. 107: 169-188.
Shaffer-Fehre, M. 199lb. The position of Najas within the
subclass Alismatidae (Monocotyledones) in the light of
new evidence from seed coat structures in the Hy-
drocharitoideae (Hydrocharitales). Bot. J. Linn. Soc. 107:
189-209.
Tomlinson, P.B. 1982. Anatomy of the monocotyledons VII.
Helobiae (Alismatidae). Oxford: Clarendon Press.
Venkatesh, C.S. 1956. Structure and dehiscence of the anther
in Najas. Bot. Not. 109: 75-82.
Viinikka, Y. 1973. The occurrence ofB chromosomes and their
effect on meiosis in Najas marina. Hereditas 75: 207-212.
Wilde, W.J.J.O. de 1962. Najadaceae. Flora Malesiana I. 6:
157-171.
Palmae
J. DRANSFIELD and N.W. UHL
Palmae Juss., Gen. Pl.: 37 (1789), nom. cons.
Arecaceae Schultz Sch. (1832), nom. cons. alt.
Small, medium-sized, or large, solitary or clus-
tered, armed or unarmed, hapaxanthic or
pleonanthic, hermaphroditic, polygamous, mono-
ecious, or dioecious plants. Stems "woody",
slender to massive, very short to very tall, creep-
ing, subterranean, climbing, or erect, usually
unbranched, rarely branching dichotomously,
lacking cambium but sometimes increasing in di-
ameter by primary growth, sometimes ventricose;
internodes very short to elongate; leaf scars con-
spicuous or not; stilt roots present or absent, roots
sometimes modified into spines. Leaves alternate,
spirally arranged, rarely distichous or tristichous;
sheath initially always tubular at the base, later
frequently splitting, unarmed or armed with
spines or prickles, glabrous or variously scaly or
hairy, sometimes with a ligulelike appendage on
either side of or in front of the petiole, sheaths
sometimes forming a crownshaft; petiole usually
present, terete, or variously adaxially channeled or
ridged, unarmed or bearing spines or teeth, gla-
brous or variously scaly or hairy; hastulae present
or absent; blade palmate, costapalmate, pinnate,
bipinnate, or bifid, or entire and pinnately veined,
plicate in bud, splitting along the adaxial folds
(induplicate), or abaxial folds (reduplicate), rarely
splitting between the folds, or not splitting; seg-
ments or leaflets lanceolate or linear to rhomboid
or wedge-shaped, V-shaped (induplicate), or /\-
shaped (reduplicate), single-fold or many-fold, a
midrib and numerous parallel secondary veins
usually present, segments very rarely splitting
further between the secondary veins, tips acute,
acuminate, truncate, oblique or bifid, praemorse,
or irregularly toothed or lobed, sometimes armed
with spines or bristles along the margins and/or
main veins, variously scaly and hairy, transverse
veinlets conspicuous or obscure; proximal leaflets
sometimes modified as spines (acanthophylls),
rachis prolonged distally into a climbing whip
(cirrus) in many climbing palms, sometimes also
bearing acanthophylls. Inflorescences axillary,
solitary or multiple, infrafoliar, interfoliar, or ag-
 Palmae
gregated into a suprafoliar compound inflores-
cence, spicate or branched up to 6 orders, usually
maturing acropetally, rarely basipetally, in some
species of Calamus inflorescence modified as a
climbing whip (flagellum); peduncle short to long;
prophyll usually 2-keeled, very varied in shape
and size, rarely subtending a first-order branch;
peduncular bracts 0-many, very varied in shape
and size; rachis shorter or longer than the pe-
duncle; rachis bracts similar to peduncular bracts,
or dissimilar, or much reduced; rachillae (flower-
bearing branches) short to long, slender to
massive, rachilla bracts conspicuous to minute or
apparently lacking, sometimes connate laterally
and adnate to the rachilla to form pits containing
the flowers. Flowers hermaphrodite or unisexual,
then similar or dimorphic, sessile or stalked,
borne singly or in cincinni of various forms in-
cluding dyads, triads, or acervuli, or rarely in
short monopodia! clusters; perianth rarely of
similar parts, usually clearly differentiated into
sepals and petals, rarely uniseriate with a variable
number of lobes; sepals (2)3 (rarely more), dis-
tinct or variously connate, usually imbricate or
basally connate, rarely valvate or widely sepa-
rated; petals (2)3 (rarely more), distinct or vari-
ously connate, valvate, imbricate, or imbricate
with brieflyvalvate tips; stamens (3)6 (or many to
950 or more), filaments erect or inflexed in bud,
free, or variously connate, or adnate to the petals,
or both connate and adnate, anthers basifixed or
dorsifixed, rarely didymous, or with widely sepa-
rated anther sacs, straight or rarely twisted, in-
trorse, latrorse, extrorse, or rarely opening by
pores; pollen including 17 aperture types, sulcate
or derived therefrom, exine intectate or tectate,
very varied in ornamentation; staminodes ranging
from toothlike to well developed, distinct or
connate, sometimes adnate to petals or gyno-
ecium, rarely absent; gynoecium apocarpous with
(1-2)3(4) carpels, or variously syncarpous with 3
or rarely with more (to 10) locules or pseudo-
monomerous with 1 fertile locule, carpels follicu-
lar or rarely ascidiform, glabrous, variously hairy,
or covered with imbricate scales, styluli distinct or
not clearly differentiated or connate into a style,
stigmas erect or recurved, rarely indistinct; ovule
solitary in each locule, anatropous, hemiana-
tropous, campylotropous, or atropous, basally,
laterally, or apically attached, crassinucellate, in-
teguments 2, the outer often wide, the inner
narrow, the outer or the inner or both integu-
ments forming the micropyle; pistillode present or
absent in the staminate flower, ranging from
minute and often trifid to large, bottle-shaped,
307
exceeding the stamens. Fruit usually 1-seeded,
sometimes 2-3-10-seeded, ranging from small to
very large, stigmatic remains apical, lateral, or
basal; pericarp smooth or hairy, prickly, corky-
warted, or covered with imbricate scales, meso-
carp fleshy, fibrous, or dry, endocarp not
differentiated or thin, sometimes with an opercu-
lum over the embryo, or thick and then often with
3 or more pores at, below, or above the middle.
Seed adhering to the pericarp or free, with thin or
sometimes fleshy testa (sarcotesta), endosperm
homogeneous or ruminate, sometimes penetrated
by the testa; endosperm storing hemicellulose,
aleurone, fats and oils but no starch; embryo api-
cal, lateral or basal. Germination adjacent-ligular,
remote-ligular, remote-tubular; eophyll simple
and entire, bifid, palmate, or pinnate.
A family of ca. 190 genera and 2000 species,
almost wholly confined to the tropics of both
hemispheres.
VEGETATIVE MORPHOLOGY. The morphological
diversity of palms is greater than that of any other
monocotyledonous family; indeed, they may rep-
resent one of the most diverse families of seed
plants as a whole (Fig. 80).
Roots. In the seedling a small primary root devel-
ops and functions for a short time. Subsequent
roots are adventitious, and usually borne laterally
near the base of the stem, often forming large
masses near ground level. The adventitious roots
lack secondary growth. Several kinds of aerial
roots also occur in the family (Tomlinson 1990);
these include stilt or prop roots, spine roots, and
pneumatophores (including pneumathodes and
pneumatorhizae).
Stem. The terms tree palms, shrub palms, acaules-
cent palms, and climbing palms (Dransfield 1978)
are useful when referring to different expressions
of the palm stem. The vegetative body of a palm is
a stem that terminates in a crown ofleaves. Shoots
are either solitary or in clusters. Clustering palms
exhibit a sympodial habit, which Holttum (1955)
considered characteristic of monocotyledons.
Each new shoot develops from an axillary bud that
in most palms is located near the base of the stem.
As each shoot then produces a new axillary shoot
in turn, a clustered habit results. Many genera
include both species with solitary stems (monopo-
dia!) and species that are clustering. No palms are
truly herbaceous, and none develops bulbs or
corms, although enlarged bases of stems such as
the saxophone-shaped underground stem in Sabal
308 Palmae
Fig. 80A-L. Growth forms. A Roystonea regia, large tree palm
with well-defined crownshaft. B Corypha umbraculifera,
hapaxanthic with a suprafoliar mass of inflorescences forming
a compound terminal inflorescence. C Washingtonia filifera,
stem partially covered with persistent marcescent leaves. D
Colpothrinax wrightii, stem ventricose. E Hyphaene thebaica,
stem dichotomously branched. F Chamaedorea pinnatifrons,
a dwarf palmlet of the forest understory. G Dypsis lutescens,
caespitose habit. H Acoelorraphe wrightii, densely caespitose
habit. I Sabal etonia, underground stem. J Salacca
wallichiana, acaulescent habit. K Serenoa repens, stem pros-
trate and branching. L Calamus sp., climber with cirrate
leaves. (Takhtajan 1982)
and a few other genera (Fig. 89A) perhaps serve as
underground storage organs.
The development of branching shows greater
variation in palms than in any other monocotyle-
donous family. The usual pattern is the develop-
ment of a single bud in each leaf axil; the basal
ones vegetative and the more distal producing in-
florescences. However, many different growth
forms occur (Fisher 1973; Fisher and Tomlinson
1973 ). Dichotomous branching, an actual division
of the stem apex (Fig. 80E, 87X), has been de-
termined in a number of unrelated genera
(Tomlinson 1990).
 Palmae
The climbing habit has apparently developed in-
dependently several times in the family. Among the
Calamoideae (Fig. SOL) it probably has originated
three or more times and also occurs in two other
subfamilies, the Ceroxyloideae and Arecoideae.
Halle et al. (197S) proposed about 20 architec-
tural models for trees, of which four are found in
palms. Unbranched monocarpic palms, such as
Metroxylon salomonense, they called Holttum's
model. Unbranched pleonanthic palms, Metro-
xylon amicarum and many other palms, were
called Corner's model. Branched palms illustrate
Tomlinson's model, and single stems with di-
chotomous branching Schoute's model. Differ-
ent individuals of the same species of palms may
display Corner's or Tomlinson's models. Other
growth forms occur (Bernal 199S). Whether
the stem is determinate or indeterminate (i.e.,
whether the palm is hapaxanthic or pleonanthic)
may be of more fundamental architectural
significance.
Leaf The leaf is the most conspicuous and dis-
tinctive organ of the Palmae (Fig. Sl). Its charac-
teristic plication and splitting into segments or
leaflets are paralleled only in some members of
Cyclanthaceae and in Curculigo seychellensis
(Hypoxidaceae). The palm leaf is characterized by
a sheath that encircles the stem and that may split
abaxially at the base in some Coryphoideae or may
be a very prominent tubular structure, a crown-
shaft, in certain Arecoideae. The leaf also usually
has a distinct petiole, and a blade that has a single
stout central axis (the rachis), which is short or
absent in truly palmate leaves, better developed in
costa palmate leaves, and prominent in the pinnate
leaf.
Leaves differ enormously in size at maturity.
Among the smallest are the bifid or undivided
blades of Chamaedorea tenella and C. tuerck-
heimii with blades sometimes less than 15 em
long; in contrast are the blades of Raphia with a
length of about 25m (Halle 1977), the largest
leaves in the plant kingdom. Leaves also differ
markedly in size and shape in a progression from
eophylls (the first leaves with blades) of seedlings
throughout juvenile stages to the mature blade
(Tomlinson 1990).
The blade is always folded. Whether the folds or
plications originate by splitting or differential
growth has been long debated (Kaplan et al.
19S2a). However, detailed studies have shown that
differential growth is totally responsible for plica-
tion development in the pinnate leaf of Dypsis
(Dengler et al. 19S2) and in the palmate leaf of
Rhapis (Kaplan et al. 19S2b).
309
The blades of mature palms may be undivided
or only bifid at the apex, or much more often regu-
larly or irregularly divided into 1-ribbed or
several-ribbed units each consisting of a single
fold or of several folds. These units are called seg-
ments in palmate and costapalmate leaves, and
leaflets or pinnae in pinnate leaves. One genus,
Caryota, is characterized by bipinnate leaves in
which the pinnae have a prominent midrib but he
pinnules lack midribs (Fig. S1 T-V).
Palm leaves differ in another way. Some blades
are divided so that the leaflets or segments appear
to be V-shaped and the blade is then termed
induplicate (Fig. S1P). Induplicate leaves charac-
terize subfamily Coryphoideae, with a few excep-
tions, and all genera of tribe Caryoteae of the
Arecoideae. All other groups of palms have leaves
which are divided so that the leaflets appear to be
tent-shaped and the blade is termed reduplicate
(Fig. SIN).
In its simplest but not necessarily most primi-
tive form, the blade is undivided except slightly
along the margins as in some of both indupli-
cate and reduplicate leaves. It is bifid at the apex
in such reduplicate entire leaves as those of
Asterogyne martiana (Fig. S1L). When the blade is
divided the segments or leaflets may each have
only a single fold, or in both palmate and pinnate
leaves the blade may be unequally divided into
segments or leaflets, some or all with several folds.
The simplest eophyll is linear to elliptic and
entire. Undivided eophylls of both induplicate
(e.g., Phoenix) and reduplicate genera (e.g.,
Clinostigma haerestigma, Roystonea) are basically
similar. Bifid eophylls are most common in palms
with reduplicate leaves, and usually with an obvi-
ous rachis. Compound eophylls occur in Latania
(Coryphoideae: Borasseae) and several species of
Chamaedorea, e.g., C. elegans (Ceroxyloideae).
These suggest further stages in evolution that may
correspond with levels of specialization suggested
for major categories. In Nypa, the eophyll may
sometimes appear imparipinnate.
The eophylls are followed by an often long pro-
gression of juvenile leaves, where the size and
nature of the mature blade develops through in-
crease in the number of lateral ribs and through
the division of the blade into segments or leaflets.
Different patterns of shape and division appear
in group after group. The great diversity of leaf
form in the least specialized subfamily, Cory-
phoideae, has only recently been appreciated
(Dransfield et al. 1990). The palmate blade varies
from flabellate to cuneate in outline, and from
nearly lacking a rachis to possessing a long rachis,
310
Fig. SlA-W. Leaf structure in palms. A-D Serenoa repens. A
Induplicate palmate leaf. B Two segments. C Base of lamina
showing adaxial hastula. D Cross-section of lamina showing
induplicate folding. E, F Mauritia flexuosa. E Reduplicate pal-
mate leaf. F Cross-section of blade showing reduplicate seg-
ments. G Chelyocarpus ulei, induplicate palmate leaf with
short marginal splits and deep divisions, producing wedge-
shaped segments. H, I Saba/ palmetto. H Induplicate
costapalmate leaf. I Group of segments showing intersegment
fibers. J Copernicia rigida, palmate leaf with large adaxial
hastula. K ]ohannesteijsmannia altifrons, entire leaf. L
Asterogyne martiana, entire-bifid leaf. M Nypa fruticans, re-
duplicate pinnate leaf. N Butia capitata, portion of rachis
Palmae
showing insertion of reduplicate leaflets. 0 , P Phoenix
roebelenii. 0 Tip of imparipinnate induplicate leaf. P portion
of rachis showing insertion of induplicate leaflets. Q
Chamaedorea schiedeana, irregularly pinnate leaf with multi-
fold leaflets. R, S Socratea salazarii. R Pinnate leaf with
praemorse segments. S Detail of single leaflet. T-V Caryota
mitis. T Bipinnate leaf with induplicate leaflets. U Detail of
single, praemorse leaflet. V Juvenile, once-pinnate leaf. W
Ceratolobus pseudoconcolor, a rattan showing spiny leaf
sheaths, swelling on the sheath below petiole (knee), armed
rachis, praemorse leaflets, and cirrus armed with grouped,
reflexed spines. (Takhtajan 1982)
 Palmae
then approaching a pinnately ribbed form. Cir-
cumscribed to include Phoenix, the subfamily
includes palmate, costapalmate, pinnate, and
entire-leaved taxa.
Several different forms of pinnate leaves also
occur. The pinnate blade in large groups, such
as the tribes Calameae, Areceae, and Cocoeae,
exhibits nearly the full range of dissection from
undivided to regularly pinnate. Leaflets may be
in groups in one plane, or in groups in several
planes, sometimes in the same genus as
in Salacca (Calamoideae). Distal leaflets are
modified into straight to recurved hooks
(acanthophylls) in climbing species of three
subfamilies (Calamoideae, Ceroxyloideae, Are-
coideae). Other elaborate adaptations for
climbing have evolved in many species of Calamo-
ideae, where the rachis extends distally beyond the
leaflets as a specialized climbing organ (cirrus),
armed with recurved grapnel-like spines (Fig.
81W).
Two further structures associated with the leaf
are noteworthy. The leaf of many, but not all,
members of tribes Corypheae and Borasseae
(Coryphoideae) bears an adaxial and sometimes
an abaxial crest or hastula at the junction of the
petiole and blade (Fig. 81C,J). The origin and
adaptive nature of the hastula are as yet not
clear.
Another structure in the position of a ligule at
the top of the sheath in front of the petiole, or
sometimes lateral to the petiole, is found in the
leaves of many palms, as in Livistona. In
Calamoideae (where it is termed an ocrea), such a
structure often harbors ants.
The evolution of the palm leaf remains in ques-
tion. Both palmate and pinnate leaves in palms
may be derived by the differential expansion,
elongation, and dissection of a primitively simple
leaf. The various forms of the palm leaf appear to
have arisen more than once. The palmate leaf
appears to have developed separately in the
coryphoid and calamoid subfamilies, especially
since it is reduplicate in the latter.
Similarly, the pinnate leaf appears to have
evolved three or more times - once in Phoenix
within the Coryphoideae, once in the Caryoteae,
and again in the palms with reduplicately folded
leaves, perhaps separately in each of the calamoid,
nypoid, ceroxyloid, and arecoid subfamilies. A
main trend in the evolution of the leaf in palms
seems to have been toward greater complexity.
Armature. Armature of various kinds occurs
throughout the family and can be divided into
three kinds: modified organs, true emergences,
311
and forms derived internally, i.e., single or mul-
tiple, sharp, vascular or fibrous bundles exposed
by disintegration of sheath or lamina (Tomlinson
1990). The genus Phoenix (Coryphoideae) can be
distinguished immediately from all other palms
with pinnate leaves by the stout, spinelike,
induplicate leaflets on the basal portion of the
blade. In the Corypheae and Borasseae large teeth
are often present along the margins of the petiole
and sometimes of the blade. In several coryphoid
genera, e.g., Rhapidophyllum, Zambia, and in
Trithrinax acanthocoma, Guihaia argyrata, and
Maxburretia furtadoana, the leaf sheath dis-
integrates into long stiff spines derived from its
fibers. In two subtribes, Oncospermatinae and
Bactridinae (Arecoideae), presence of spines is
diagnostic for the group, and many genera of
Calamoideae are ferociously armed.
VEGETATIVE ANATOMY. Roots. The anatomy of
palm roots conforms to the usual pattern of alter-
nating bands of xylem and phloem with a central
pith (Tomlinson 1990); however, larger roots may
be polystelic and exhibit complex stelar patterns.
Seubert (1996a,b, 1997) investigated root anatomy
in a number of genera in all subfamilies except
Arecoideae and found that such characters as
variation in the rhizodermis (velamen) and the
exodermis, the nature of the cortex, monostely or
polystely, lignification of parenchyma, occurrence
of bundles in the pith, and distribution of scleren-
chyma may be characteristic of genera or species
and useful in phylogenetic studies. Contrary to
some previous reports that palms lack root hairs,
Seubert (1997), in a survey of 36 genera of
Coryphoideae, found that they are frequent.
The Stem Wood. Palm "wood" differs from the
wood of dicotyledonous trees, which is largely sec-
ondary and formed from a ringlike meristem, the
cambium. In palms, as in monocotyledons gener-
ally, such a cambium is lacking, and the wood
consists of primary tissues, which originate from
the growing tip. Once fully formed, the stems of
many palms do not increase in diameter. Vascular
bundles are variously scattered in a parenchyma-
tous ground tissue with fibers of two kinds, vascu-
lar fibers, which are associated with the bundles of
xylem and phloem, and nonvascular fibers, which
occur singly or in bundles but not associated with
the conducting tissues.
Most palm stems are divided into a narrow
cortex and an inner zone containing scattered vas-
cular bundles. Within the inner zone different
palms exhibit different distributions of vascular
strands and fibers (Parthasarathy and Klotz 1976).
312 Palmae
In the Coryphoideae, most climbing palms, and
the coconut, the vascular bundles are rather
evenly distributed, giving a homogeneous appear-
ance to the center of the stem in cross section; in
some other palms such as Raphia (Pyykko 1985)
the peripheral vascular bundles are closer together
and may be surrounded by more sclerenchyma in
the form of larger fibrous bundle sheaths and
separate fibrous bundles, forming a stem with a
soft center and an extremely tough or hard pe-
riphery. The character of the ground tissues also
affects the "wood"; the parenchyma cells may be
sclerotic or lignified with few intercellular spaces
or thin-walled with large intercellular spaces. In
some palms (e.g., Metroxylon) they contain abun-
dant starch, which is often exploited by man.
Thus, the distribution of the vascular bundles and
of the vascular and nonvascular fibers, and the
nature of the ground parenchyma give palm wood
its character. Palm stems do not produce lumber
comparable to that of dicotyledonous trees, but
they show great elasticity in tropical storms. Their
toughness and flexibility make some of them valu-
able as construction materials and others useful
for basketry.
Silica Bodies. The hardness of palm stems and
leaves results partly from the presence of silica
bodies in specialized cells called stegmata
(Tomlinson 1961) which are associated with vas-
cular or nonvascular fibers. The form of the silica
bodies, either hat-shaped or spherical, is some-
times of diagnostic value. Ultrastructure of silica
bodies in Calamus axillaris (Schmitt et al. 1995)
has shown that silica bodies form in vacuoles of
the immature cells and that subsequently the walls
thicken, becoming extremely thick and envelop-
ing nearly half of the silica body, and the cyto-
plasm of the stegma degenerates after thickening
of the wall.
Xylem. The vascular bundles of palm stems and
other organs have certain common characteris-
tics. In each bundle there are 1, 2, or several large
xylem elements or vessels (for discussion of termi-
nology see Klotz 1978a), one or more phloem
area(s), and usually a fibrous bundle sheath. In
cross-section the number of large vessels and the
often distinctive shape of the fibrous sheaths may
aid in the identification of a sample as a palm
(Tomlinson 1961, Klotz 1978b). These characters
are especially useful when dealing with fossil
materials.
The relative lengths and certain features of the
end walls of the vessel elements may also be useful
in determining relationships and evolutionary
trends. The variously sculptured end walls of
vessels are considered least specialized if they bear
long scalariform (ladderlike) perforation plates,
intermediate in specialization, if shorter with
fewer bars, and most advanced when porous. A
survey by Klotz (1978b) of large vessels in the
roots, stems, and petioles of 209 species in 169
genera of palms has shown that vessels are most
advanced in roots, intermediate in stems, and
least advanced in leaves. This agrees with trends
established for monocotyledons as a whole
(Cheadle 1943, 1944). Nypa, one of the earliest
palms in the fossil record, is among the palms with
the most primitive vessels and represents the only
subfamily lacking simple perforation plates.
Phloem. A study of 374 species in 163 genera of
palms by Parthasarathy (1968) found sieve ele-
ments in roots to have compound sieve plates on
very oblique to oblique end walls, while those in
stems and petioles have end walls varying from
very oblique to transverse with compound to
simple sieve plates. In the species studied, com-
pound sieve plates predominate except among the
Calamoideae and Nypoideae, which exhibit a high
percentage of simple sieve plates on transverse to
oblique end walls of stems and petioles. Other-
wise, in palms the inclination of the end wall of
sieve-tube elements shows a progressive change
from oblique in the roots to transverse in the leaf
lamina. In general, observations on palms support
the conclusions about sieve tubes in other mono-
cotyledons (Cheadle 1948).
A striking correlation is found in the distribu-
tion of 1 or 2 phloem strands in central vascular
bundles of the petiole. One phloem strand occurs
in certain genera of Corypheae and in Phoeniceae,
Borasseae, Nypoideae, Ceroxyloideae, Caryoteae,
and Iriarteeae; 2 phloem strands occur in other
genera of coryphoid palms and in Calamoideae,
Areceae, Cocoeae, Geonomeae, and Phyteleph-
antoideae. The single strand is associated with less
specialized groups or with less specialized genera
in the coryphoid palms except for perhaps
Corypha, Nannorrhops, and Sabal. Two strands
are generally associated with more specialized
groups and with more specialized genera within
the Corypheae. Thus, there seems a clear sugges-
tion of an evolutionary trend toward the develop-
ment of 2 phoem strands in the central vascular
bundles of the petiole.
It is noteworthy also that sieve elements are
long-lived, presumably functioning for the life of
the palm, perhaps for decades or a century or
more, with little change in structure except for an
apparent decrease in the amount of endoplas-
matic reticulum and increased degeneration of
 Palmae 313

mitochondria and plastids in the older elements studies have proved diagnostically useful. Read
(Parthasarathy 1974). The sieve elements differ (1975) found anatomical characters to distinguish
structurally from those of most dicotyledons in all four species of Thrinax. The species of Chelyo-
absence of obvious quantities of P-protein and in carpus (Uhl 1972b) and of Hyophorbe (Uhl 1978)
failure to produce definitive callose except after can also be identified from transections of the
leaf fall (Parthasarathy and Klotz 1976). lamina. A large number of characters may be
Vasculature. Patterns of vascular bundles in helpful with diagnoses or phylogenetic studies.
palm stems have been extensively studied by These include epidermal features, presence or
Zimmermann and Tomlinson (1965, 1974). Like absence and number of hypodermal layers and
many researchers on palms, the investigators had palisade tissue, patterns and characteristics of
to develop techniques appropriate to the difficult sclerenchyma, and vein structure.
material, in this case the very large number of Developmental studies have also been impor-
bundles and their great length. The coconut stem tant in understanding the leaf. Studies of how
has about 20 000 bundles. The investigators chose the plications developed proved to have broad
a smaller palm, Rhapis excelsa, with only about implications for developmental processes in
1000 bundles in the stem, and adapted the tech- biology as a whole (Dengler et al. 1982; Kaplan
nique of cinematography, the photographing of et al. 1982a,b).
serial sections or sequentially cut surfaces on
movie film, for the task. From initial application to INFLORESCENCE STRUCTURE. Inflorescences,
the stem of Rhapis, it has been subsequently flower clusters, and flowers are remarkably varied,
applied to the rhizome, juvenile phase, apex, but the many patterns can be related to relatively
inflorescence, root and branch, flower, the vessel simple basic patterns of structure and maturation.
network, vessel length distribution in the stem, Inflorescence in palms may be regarded in two
xylem structure of the leaf meristem, and finally to ways - in the physiological sense of a reproductive
transport. Cinematography has also made pos- event or flowering, and in the strictly morphologi-
sible extensive research on the conducting cal sense of an axillary branch system. In the first
systems, transport problems, and crown structure sense, palms appear primitively and predomi-
of other large monocotyledons (see Tomlinson nantly to be of indeterminate growth, i.e.,
1990 and references therein). pleonanthic, but determinate growth (the hapax-
When a single bundle is followed through a anthic condition) has developed several times and
palm stem, a relatively simple pattern can be dis- in different groups.
cerned (Zimmermann and Tomlinson 1974). The In a morphological sense, a remarkable series
bundle curves from the center of the stem towards of variations is based on an axillary monopodia!
its periphery and, before entering a leaf, branches branch system as characteristic of unspecialized
in an upward direction, producing a continuing Corypheae (Tomlinson and Moore 1968). This
axial bundle. During its course the original bundle system is composed of a main axis or branch on
may also produce several short branches (bridges) which is borne an adaxially 2-keeled prophyll, and
that unite with neighboring bundles, and other usually 1-several empty peduncular bracts, which
branches that become traces to axillary buds and are followed by rachis bracts, each of which sub-
inflorescences. Because this vasculature was deter- tends a branch of the first order. These branches,
mined by extensive study of Rhapis excelsa, its which may sometimes also bear prophylls, may be
form has become known as the Rhapis principle. again branched one to several times, each succes-
The Rhapis principle is of primary importance in sive branch subtended by a bract and the ultimate
interpreting vascular patterns in other palms and axes or rachillae bearing solitary flowers each
monocotyledons. subtended in turn by a small bract and some-
Despite considerable research on the histology times bearing a bracteole. A sympodial or rarely
of palm stems (Tomlinson 1961, 1990) and on (Phytelephantoideae) monopodia! system devel-
their vascular systems, many palms remain to be ops to produce the diverse flower clusters consid-
studied. At present, although it is relatively easy to ered below. Except for the prophyll and empty
recognize a fossil palm stem by the large bundles peduncular bracts, there is a 1: 1 relationship
of characteristic form, it is difficult or impossible between bracts and branches including, with a few
to relate the palm to a modern genus. exceptions (see flowering units), the flowers. A
Leaf Tomlinson (1961) described the anatomy of simple system for referring to the orders of
the segments or leaflets of many genera and out- branches and of bracts was devised for Nan-
lined trends within the family. A few subsequent norrhops (Tomlinson and Moore 1968).
314 Palmae
In Calamus, the inflorescence is often modified
by sterilization into a specialized climbing organ
(flagellum) which is adnate (fused) to the leaf
sheath above the sheath which subtends it. In
other Calamoideae, adnation between the inflo-
rescence and internode above occurs. The amount
of inflorescence adnation usually tends to be
greater in presumed specialized species than in
unspecialized ones, but, in general, adnation in
the inflorescence occurs in the less specialized
subfamilies. Substantial basal adnation of axes
to those of the internode above occurs in the
Corypheae (Coryphoideae), Nypoideae, and Cala-
moideae, but adnation of branches is not found
in inflorescences of Ceroxyloideae, Arecoideae,
or Phytelephantoideae (Uhl and Dransfield
1987).
The most common position of the inflorescence
is among the leaves (interfoliar), but in many
Arecoideae it is below the crown (infrafoliar),
being exposed in bud by the fall of the deciduous
subtending leaf which has a long tubular sheath.
When inflorescences are produced in the axils of
bracts above the crown, as in Corypha and
Metroxylon, they are suprafoliar.
Most inflorescences are solitary in the leaf axil,
but several inflorescences rather than the usual
one at each node are produced in two subfamilies.
Multiple inflorescences occur in some members of
Caryoteae, Iriarteeae, and Areceae (Arecoideae),
and in Ceroxyleae and Hyophorbeae (Ceroxy-
loideae).
By studies of development, Fisher and Moore
(1977) showed that multiple inflorescence buds
develop from an axillary meristem. The multiple
inflorescences appear to have several adaptive fea-
tures. Since many of them are staminate, they in-
crease the number of staminate flowers and the
amount of pollen produced. They also extend the
flowering period for both seasonal and nonsea-
sonal species, and they offer increased protection
of reproductive structures against herbivory
(Fisher and Moore 1977).
The Prophylls. Bracts occur in many different
forms. The unvarying presence of an adaxially
2-keeled prophyll on the base of the peduncle of
the inflorescence is, along with the leaf, a distinc-
tive feature of the Palmae. The prophyll always
completely encloses the inflorescence in early
stages and in its simplest form is tubular and 2-
keeled. In members of Areceae, the prophyll may
enlarge with the peduncular bract, the 2 bracts
reaching nearly equal size and opening at about
the same time. Rarely, in some genera of
Ceroxyloideae and Arecoideae, the prophyll is
incomplete, only partly encircling the peduncle
(Moore 1973).
In some Coryphoideae, Calamoideae, and in
Nypa, prophylls are borne on lateral branches as
well as on the main axis. Their occurrence on
lateral branches in these less specialized groups
suggests the possibility that a prophyll associated
with each branch may represent a basic, primitive
condition.
Peduncular Bracts and Rachis Bracts. One or a
few to many empty bracts, termed peduncular
bracts, may be borne on the peduncle above the
prophyll. Since even the prophyll may sometimes
subtend a first-order branch (Tomlinson and
Moore 1968; Tomlinson and Soderholm 1975), it is
probable that these peduncular bracts have
evolved through suppression of the subtended
branch and specialization in various ways. In
many coryphoid palms they are small, tubular,
sheathing appendages, opening early at the apex.
In many palms of the Arecoideae, they are large,
closed, exceedingly hard, often beaked structures
that enclose the entire inflorescence in bud, split-
ting abaxially and becoming marcescent or more
often deciduous at anthesis.
The number of peduncular bracts is important
in delimiting subfamilies and tribes. Three to
more than 10 empty peduncular bracts character-
ize most Coryphoideae, Hyophorbeae (Ceroxylo-
ideae), Caryoteae, Iriarteeae, and Podococceae
(Arecoideae) and some calamoid genera (e.g.,
Mauritia). In the Arecoideae there is a clear trend
toward reduction in size or, infrequently, com-
plete loss (at maturity) of those bracts that
subtend branches.
Bracts on the inflorescence in tribes Areceae,
Cocoeae, and Geonomeae of Arecoideae and in
Phytelephantoideae are usually only 2: a some-
times large prophyll and a usually much enlarged
single peduncular bract, which is particularly
striking in subtribes Butiinae and Attaleinae
(Arecoideae: Cocoeae). Here the peduncular bract
is woody, often cowl-like, markedly plicate in bud,
long persistent above the inflorescence, and
sometimes brilliant yellow or white internally at
anthesis, when it may serve as an attractant to
pollinators. Other large peduncular bracts occur
in many arecoid genera, where they are thinner,
fibrous, and deciduous.
In whatever ways the prophyll, peduncular
bracts, and rachis bracts may be modified, they
appear to function as protective organs for the
developing inflorescence and during pollination.
In the few cases studied, the structure of peduncu-
lar bracts has been important in pollination. The
 Palmae 315

timing of bract opening can be correlated with sympodial unit. A remarkable series of such units
both floral structure and pollination events (Uhl occurs in Calamoideae (Uhland Dransfield 1987).
and Moore 1977; Henderson 1986). The Acervulus. The monoecious genera of tribe
Vascular Supply. The extensive cine analysis of Hyophorbeae (Ceroxyloideae) have unisexual
the inflorescence of Rhapis excelsa by Tomlinson flowers in reversed cincinni adnate to the rachillae
and Zimmermann (1968) established a direct vas- (Uhl and Moore 1978), forming distinct lines of
cular linkage between the inflorescence and the closely two-ranked flowers, called acervuli (Fig.
leaf that subtends it. Inflorescence anatomy re- 88H); the proximal flower is usually pistillate, the
sembles that of the stem, but at each branch a part distal staminate, and the bracteoles are reduced or
of the vascular supply of the parent axis passes not evident at anthesis.
directly into the branch without any connections The Triad. The most common flowering unit in
with the subtending bract. The vascular supply is palms, the triad (Fig. 83H), characterizes all 124
reduced in the smaller distal branches. genera of the Arecoideae. It was described as
a monopodia! unit, but clearly represents a
FLOWER CLUSTERS. Flowers are borne in several cincinnus of 2 lateral staminate and a central pis-
kinds of clusters that have a characteristic form in tillate flower. The bracteoles of the 3 flowers are
each subfamily. The basic flowering unit seems sometimes large and distinctive. The sympodial
clearly to be a single flower, sessile, or borne on a nature of this unit has been determined by de-
pedicel which is subtended by a bract and bears a tailed studies in Caryoteae, Iriarteeae, Ptychosper-
more or less 2-keeled bracteole or prophyll. The matinae, and Geonomeae (Uhl1966, 1976, Uhland
pedicel, the bract subtending it, or the bracteole Moore 1980).
may be reduced or absent. Solitary flowers are Monopodia[. Flower clusters that develop mo-
usually spirally inserted; often they are nearly nopodially characterize only the three genera of
decussate basally on rachillae, but elsewhere they subfamily Phytelephantoideae. In these palms,
can be closely crowded and somewhat irregularly splendid series representing reduction in mo-
arranged. In Borasseae (Coryphoideae), Calamo- nopodia! branching systems are evident (Uhl and
ideae, and perhaps elsewhere, some flowers have 2 Dransfield 1984).
bracts which may represent a prophyll and a floral
bracteole. FLOWER STRUCTURE. The basic flower structure
Sympodial. Clusters of several to many flowers in palms is trimerous, with 3 slightly imbricate
are either sympodial in development and form sepals, 3 slightly imbricate petals, 6 stamens in 2
cincinni\ or in phytelephantoid genera they are whorls, and 3 distinct uniovulate carpels. This
condensed monopodia! branching systems. Cin- simple pattern is modified in many ways, even
cinni result when a second flower develops in the among the least specialized coryphoid palms,
axil of the floral bracteole, the bracteole below the through connation and/or adnation, increase,
second flower then subtending a third flower, etc. reduction, or loss of parts, and by differential
This type of growth produces flower clusters of elongation of the receptacle.
various forms, depending on the relative positions While trimery is the principal pattern, tetramery
of the bracteole and its subtended flower. The occurs regularly in the perianth of staminate
form of the cincinnus is characteristic in each of flowers of phytelephantoid palms (Barfod 1991),
four subfamilies (Uhl and Dransfield 1987). The though largely obscured by the time of an thesis. A
trend in Coryphoideae seems to be from short ir- 4- to 7-parted gynoecium may occur in some
regular cincinni to longer, regularly 2-ranked species of Attalea, but the perianth is trimerous.
forms. Several to 10 carpels are present in the pistillate
The Dyad. A pair of flowers, a dyad (Fig. 86K), is flowers of phytelephantoid palms. Rarely, in
a basic form and appears to represent a 2-flowered Thrinacinae and in the staminate flowers of
Phytelephas, the entire perianth is reduced to an
irregularly lobed whorl with minute segments.
1 Although the prophylls on the palm inflorescence and its Reduction from the hermaphrodite flower to the
branches are adaxial, as usually in monocots, the floral far more common unisexual flower is by abortion
bracteoles from which the cincinnus develops are shifted into or loss of the gynoecium in staminate flowers and
a transversal position. The floral bracteole which subtends the the androecium in pistillate flowers. Staminate
second flower is borne at a right angle to the bract which
subtends the first flower of the cluster, and bracteoles of sub-
and pistillate flowers may be very similar and
sequent flowers are the same, resulting in the characteristic modified only late in ontogeny (N.W. Uhl,
branching system of the cincinnus. unpubl.), as in some Coryphoideae and in most
316
Calamoideae, or they may be markedly dimor-
phic, as in Nypa and the phytelephantoid genera.
The receptacle of palm flowers is remarkably
variable. It may be elongate below all floral organs
or between any 2 whorls of organs. In those groups
where flowers are borne in pits, the receptacle of
the staminate flower elongates between the inser-
tions of the calyx and corolla (Welfia), or between
the corolla and stamens (Lodoicea) by late
intercalary growth (N.W. Uhl, unpubl.) to push
the stamens out of the pit at anthesis (Fig. 85C).
The pistillate flower of Podococcus is similarly ex-
serted, at least after fertilization, but in pistillate
flowers of Geonomeae, the style rather than the
receptacle elongates to exsert the stigmas, and the
fruit develops within the pit and ultimately forces
it open. Flowers of Chuniophoenix are exserted
from tubular bracts by elongation of a recep-
tacular stalk (J. Dransfield, unpubl). Remarkable
plasticity of the floral receptacle as well as the
floral organs is evident.
The size and shape of sepals varies from group
to group; connation occurs frequently, especially
in the staminate flower. Petals vary similarly,
common trends being toward the valvate state at
maturity in the hermaphrodite and staminate
flower and toward extreme imbrication in the
pistillate flower. Gamopetalous, often valvately
lobed corollas are present in some genera in all
subfamilies except Nypoideae, frequently with
connation and/or adnation of stamen filaments or
staminodes in the Coryphoideae and Geonomeae.
Such adnation occurs more frequently in the
pistillate than in the staminate flower.
The vasculature of perianth parts varies from
none to many bundles, and their histology from
parenchymatous to extremely tanniniferous and
fibrous, with fibers present in bundle sheaths
or as separate strands. Raphides may also occur
in specific locations, often in abundance.
Stamens and staminodes have !-several vascular
bundles and may be fibrous or histologically
specialized.
The palm stamen typically has a relatively
slender, distinct filament and a moderately elon-
gate latrorse anther. The primary vascular supply
consists of 1 or 2 traces, but several bundles may
be present in larger, more specialized filaments.
Expansion of filaments and increase in number of
traces per stamen appear to be characteristic of
advanced groups. When stamens are reduced in
pistillate flowers, they may simply have abortive
anthers or they may be reduced to filaments, to
minute teeth, or to a cupule, sometimes large and
fibrous (Attalea, Cocoeae), at the base of the
Palmae
ovary; sometimes staminodes fail to develop. In
some Geonomeae (Calyptrogyne, Calyptronoma),
staminodes form a unique, distally expanded tube
around the stigmas, which in Calyptrogyne serves
as food for the bat pollinators.
The usual number of stamens among the palms
is 6 in 2 whorls. Reduction to three opposite the
sepals occurs in Nypa, where the filaments are
fused to form a column (Uhl 1972a). Three
stamens also characterize one species of Syne-
chanthus (Ceroxyloideae), and some species of
Wallichia, Orania, Dypsis (where the 3 stamens
may be antesepalous or antepetalous), Areca, and
Geonoma (Arecoideae). More than 6 stamens
occur in some 70 genera, including members of all
subfamilies except Nypoideae. In multistaminate
taxa, the number of stamens ranges from 7 or 8 to
mostly 9-many. The largest numbers of stamens
within the palms are found in the Phyteleph-
antoideae where samples of 849-954 have been
counted in Phytelephas aequatorialis, and over
1000 in Ammandra.
Increase in stamen number appears to have oc-
curred in different ways and perhaps in response
to different factors in different groups of palms.
Androecial development has been studied in all
but two of the groups where many stamens occur
(Uhl and Moore 1980). After petal inception, the
floral apex expands in diameter and/or height in a
different way in each group to accommodate the
larger number of stamens. In Phytelephantoideae
stamens arise centrifugally on a circular apex.
Elsewhere, stamens develop in different arrange-
ments in antesepalous and antepetalous positions.
A larger number of stamens in Thrinax, Co-
ccothrinax, and Zambia (Thrinacinae) seems as-
sociated with wind pollination, but the numerous
stamens in Chelyocarpus and Itaya of the same
subtribe must be explained differently.
In the development of multistaminate and-
roecia, the mode of the apical expansion and the
resulting stamen patterns appear to conform to
pressures exerted by bracts and perianth seg-
ments. Thus, it seems reasonable to suggest that
specialization of the inflorescence bracts and peri-
anth occurred before the advent of multistaminy
in the palms as a whole. The presence of a trimer-
ous perianth and gynoecium in the multis-
taminate taxa (except in Phytelephantoideae)
supports this conclusion.
Trimery in the gynoecium is basic, as it is also
elsewhere in the flower, but there are exceptions.
Among the 16 genera of apocarpous palms, five
genera, each specialized in some way, develop
only a single carpel. Among the 17 subtribes of the
 Palmae
syncarpous tribe Areceae (Arecoideae), all except
five triovulate genera in four subtribes exhibit
pseudomonomery (where only one carpel devel-
ops completely and bears a fertile ovule, but parts
of two other carpels are present). More than three
carpels occur in only five genera: Chelyocarpus
(Coryphoideae), Attalea (Arecoideae: Cocoeae),
and all three genera of Phytelephantoideae.
The least specialized carpel among the apo-
carpous coryphoid and phoenicoid palms is
comparable to that considered primitive in
dicotyledons in follicular shape, conduplicate
nature, open ventral suture, and to a large extent
in vascular supply. The carpel of Nypa, however,
with its conduplicate margins, cupular shape, 2-
crested distal opening, diffuse internal stigmatic
surfaces and laminar to submarginal placentation,
may represent a unique type of carpel among the
angiosperms. Specialization of the carpel includes
modifications of the base or stalk, of the ovarian
part, and of the stylulus and stigma (Uhl and
Moore 1971).
The vasculature of the carpel usually consists of
a dorsal and two ventral bundles, and one to
several pairs of lateral bundles. In thick-walled
carpels accessory bundle systems may also be
present.
Anatropous, hemianatropous, campylotropous,
and atropous ovules occur in palms. All of these
forms are found in the unspecialized Corypheae.
In Coryphoideae and in the Cocoeae, anatropous
ovules characterize taxa less specialized in other
ways, suggesting, as might be expected, that anat-
ropous ovules are primitive within the family.
Atropy occurs within Borasseae, and Cocoeae, and
hemianatropy and campylotropy within the
Ceroxyloideae and various Arecoideae. A notable
feature of some palm ovules is their vasculariza-
tion with the numerous bundles entering the large
outer integument. The inner integument is usually
narrow, frequently 2 cells wide. The ovule also
varies in its position within the locule, in the pres-
ence or absence of an aril, in the size and fusion of
the 2 integuments, and in nucellar structure.
EMBRYOLOGY. Embryological features of palms
include (1) a secretory anther tapetum; (2) mostly
simultaneous cytokinesis of the pollen mother
cell; (3) 2-celled pollen grains; (4) bitegmic,
crassinucellate ovules with well-developed integu-
mentary vascular bundles; (5) Polygonum and
Allium type embryo sacs; (6) Nuclear type of en-
dosperm, rumination at maturity; (7) Asterid and
Onagrad type of embryogeny (J ohri et al. 1992,
who provide more detailed information).
317
POLLEN MORPHOLOGY. The remarkable variation
in pollen morphology as compared to all other
monocotyledonous families was evident from
early light microscopy studies (Thanikaimoni
1971; Sowunmi 1972) and has been documented
and confirmed by a large survey of approximately
1150 specimens, using SEM and TEM and includ-
ing all but about five genera (Harley 1996).
Extraordinary variation is found in apertures and
in appearance of the tectum surface. Seventeen
different aperture types or conditions have been
identified, a selection of which is shown in Fig.
82. The exine may be tectate, semitectate, or
intectate; 12 forms of exine architecture have been
described (Harley 1996). However, the least spe-
cialized sulcate aperture and simple tectum are
the most common forms throughout the family.
Grains range widely in size. The smallest are
found in the Thrinacinae and Phoeniceae
(Coryphoideae), and the largest in the Borasseae
(Coryphoideae), Attaleinae (Arecoideae), and in
the Phytelephantoideae.
KARYOLOGY. Chromosome numbers have been
published for about 120 of the 190 genera of palms
(Uhl and Dransfield 1987; Roser 1994). Numbers
range between 2n = 36 and 2n = 26, thus in-
cluding every gametic number from 13 to 18, a
dysploid series. Species of the same genus and
members of some groups have identical chromo-
some numbers. Most palms of the subfamily
Coryphoideae, which we consider on other
grounds to be the least specialized group, have n
= 18. All evidence available points ton= 18 as the
most likely ancestral chromosome number, with
descending series of dysploid numbers as far as n
= 13 derived from that, probably independently
in several different evolutionary lines. Rearrange-
ments of chromosome parts and dysploid lower-
ing of the gametic number, one step at a time,
has often accompanied evolution in various
lines leading to other groups of palms, culmi-
nating independently in n = 13 in both Cala-
mus (Calamoideae) and Chamaedorea (Ceroxy-
loideae). Polyploidy is of minor significance,
but Voanioala, a Madagascan endemic, has 2n =
606 ± 3, possibly the highest number in the mono-
cotyledons (Johnson 1989; Roser 1994).
The family is highly differentiated in the
structure of interphase nuclei and longitudinal
differentiation of prophase and metaphase chro-
mosomes (Roser 1994). Alteration of interphase
chromatin structure and condensation behavior
can be correlated with the differentiation of major
groups of palms including subfamilies, tribes, and
318
Fig. 82A-L. SEM micrographs of palm pollen, illustrating 12
of the 17 major aperture types recognized for palm pollen
by M.M. Harley. A Aperture type 1: symmetric sulcate,
Gronophyllum sp., X 1900. B Aperture type 2: asymmetric sul-
cate, Geonoma macrostachya, X 1360. C Aperture type 3: ex-
tended sulcate, Pinanga maculata, X 1430. D Aperture type 4:
brevisulcate, Ammandra decasperma, X750. E Aperture type
5: equatorial disulcate, Metroxylon salomonense, X 1080. F
Aperture type 6: distal disulcate, Iriartella setigera, X2800.
Pal mae
G Aperture type 7: symmetric trichotomosulcate, Bactris
concinna, X 1250. H Aperture type 9b: symmetric monoporate
oblate, Areca caliso, X 1570. I Aperture type 12: subequatorial
diporate, Daemonorops sparsiflora, X 1600. JAperture type 14:
subapical triporate, Sclerosperma mannii, X950. K Aperture
type 15: incomplete zonosulcate, Areca abdulrahmannii,
X 1180. L Aperture type 16: zonosulcate, Nypafruticans, XSOO.
(Courtesy M.M. Harley, Palynological Unit, Royal Botanic
Gardens Kew)
 Palmae 319

subtribes of Coryphoideae and may date back to The endocarp of primitive palms is not special-
early radiations. Some common features link all ized, but a very thick endocarp is developed in
subfamilies except Coryphoideae, Ceroxyloideae the Borasseae, in Eugeissona (Calamoideae),
with Arecoideae, and Nypoideae with Phyteleph- Nypa (Nypoideae), Ptychococcus and the Cocoeae
antoideae. Roser (1994, 1995) states that nuclear (Arecoideae), and in the Phytelephantoideae. Four
characters found in subtribe Thrinacinae (Cory- types of endocarp have been described: the
phoideae) can be considered to link palms with coryphoid type differentiates from the inner part
other monocotyledons. of the fruit wall; the chamaedoreoid type forms
solely from the locular epidermis; and the cocoid-
PoLLINATION. It is only relatively recently (Uhl arecoid type develops from the locular epidermis,
and Moore 1977) that the myth (Delpino 1870) bundle sheaths, and intervening parenchyma
that palms are wind-pollinated has been dispelled. (Murray 1973). A fourth type occurs in Eugeissona
During the past decade, it has been shown that (Calamoideae). Different developmental patterns,
many palms are insect-pollinated (Henderson described as continuous, discontinuous, and basi-
1986), or, even if primarily anemophilous, that petal are correlated with the endocarp types.
entomophily occurs, as in Aiphanes (Listabarth Several other different forms of endocarp occur
1992). True anemophily does indeed occur (e.g., in the family. In a few palms, pyrenes, hard
in Thrinax) but even the date palm, Phoenix seedlike bodies, are formed when an endocarp
dactylifera, may in fact be secondarily anemophil- develops separately in the wall of each carpel of
ous. Large quantities of pollen, a feature of the a syncarpous ovary (see genera of Lataniinae).
syndrome of wind pollination, seem, at least in The pyrene forms a hard, sometimes sculptured
some palms, to be an adaptation to predation by outer layer around each seed; in Latania the
insects. Different pollination syndromes have sculpturing of pyrenes is diagnostic. Characteris-
been observed for many insects including flies, tic features of the heavy en do carps of Cocoeae are
bees, ants, and beetles (Henderson 1986 and the "pores", that are thinner portions in the en-
others). The large incidence of beetle pollination docarp. Usually there are three of these, but some-
is noteworthy. In some cases, exine sculpturing times (Attalea) more, with the embryo or embryos
appears correlated with pollination syndromes. opposite them. When only one embryo develops,
Iriartea, shown by Henderson ( 1985) to be bee- its pore is usually larger than the others. Another
pollinated, has a finely reticulate exine whereas specialization is found in the brittle, sometimes
beetle-pollinated Socratea has spiny pollen grains highly sculptured endocarps in the Iguanurinae
(Harley 1996). and Oncospermatinae (Areceae) where an opercu-
lum (a thin cap) develops opposite the embryo.
FRUIT AND SEED. The most common type of palm The usually ovoid, ellipsoidal, or globose seed,
fruit has a fleshy wall and 1 seed, or much less like the fruit, occurs in a wide range of sizes. It
frequently 2 to 10 seeds. Size varies from fruits is always provided with abundant nuclear endos-
about 4.5 mm in diameter in species of Geonoma perm that is most often homogeneous but may be
to very large as in Lodoicea, where they may be ruminate even in species of the same genus. In a
50cm long and weigh 18kg. Although most few genera (e.g., Cocos) it is hollow, The seed coat
fruits are smooth, scales cover those of all Cala- is heavily vascularized in Iriarteeae, in some
moideae, hairs appear on those of Rhapido- Areceae, and in the Cocoeae, and a sarcotesta,
phyllum and Wettinia, prickles are present on a fleshy outer seed coat, is well developed in
the fruits of some species of Astrocaryum and many Calamoideae. Sometimes the endosperm is
Bactris, while warty processes that arise by invaginated by the seed coat in Corypheae and
cracking of the pericarp characterize fruits of Phoeniceae, or it may be dissected (Coccothrinax,
one species of Chelyocarpus, and of Itaya, Zambia) or perforate (Thrinax). The seed of
]ohannesteijsmannia, a few species of Licuala, Lodoicea, the largest in the plant kingdom, is
and all but one species of Pholidocarpus (Cory- bilobed, while seeds of some genera in the
pheae), Manicaria, Lemurophoenix, Pelagodoxa, Ptychospermatinae, some Borasseae, and the
Sommieria (Areceae), and all three genera of calamoid genus Eugeissona are angled or ridged
Phytelephantoideae. Crystals, fibers, fiber- and grooved. Seeds of some genera of the
sclereids, sclerenchyma, and tannin are common Iguanurinae are elaborately sculptured in confor-
constituents of the mesocarp, presumably serve a mity with the endocarp.
protective function, and may be diagnostic (Essig The mostly small embryos are cylindrical to
1998). conical and late-maturing. All palms have hypo-
320 Palmae
geal germination, as the cotyledon does not
emerge from the seed. Depending on the extent of
elongation of the cotyledonary axis (or petiole),
three types of germination are recognized: adja-
cent, remote-ligular, and remote-tubular. The less
specialized palms have remote-tubular germina-
tion. Remote-ligular germination is largely associ-
ated with some genera of advanced groups and
appears to represent an adaptation to germination
in a moist environment. Germinating palms, for
the most part, require considerable moisture and
protection from excessive light, and in this respect
they are most vulnerable when the tropical forests
are cut. Though palms are often left standing in
open fields, reproduction is very rarely successful
in such a habitat.
DISPERSAL. Information concerning the disper-
sal of palms is very scanty and consists mostly
of casual observations. A few species have fruits
dispersed by water. The best-known examples are
Nypa fruticans and the coconut (Cocos nucifera).
The very thick, fibrous mesocarps of these two
species are seen as adaptations to floating. How-
ever, the recently described Ravenea musicalis
(Dransfield and Beentje 1995) is noteworthy as
both fruit and seed are adapted to dispersal by
water and the seeds germinate under water. The
greatest number of palms seem to be adapted to
dispersal by animals. A survey of the literature
(Zona and Henderson 1989) found that birds,
bats, nonflying mammals, reptiles, insects, and
fish feed on palm fruits and scatter the seeds.
Either obligate or opportunistic frugivores may
disperse seeds.
PHYTOCHEMISTRY. In two palms, presence ofUV
fluorescence of unlignified cell walls has been
proven (Harris and Hartley 1980). Oxalate and
silica are present as raphide cells and in stegmata.
The family is rich in polyphenols such as C-
glycosyl flavones (also as sulfate), tricin, and con-
densed tannins including catechins, proantho-
cyanidins, and catechins. Steroid saponins have
variously been found, but alkaloids are rare
(Areca). The dragon's blood produced by some
Daemonorops spp. is a resinous mixture of
diterpenes and sesquiterpenes with flavan-derived
pigments. More detailed information and refer-
ences are given in Hegnauer (1963, 1986). So far,
secondary chemistry has hardly been used in the
context of systematic studies within the family.
DISTRIBUTION AND HABITATS. Palms are con-
spiCUOUS and important components of many veg-
etation types in the tropics and subtropics. They
show a great diversity in ecological adaptations,
but generalizations are difficult to make.
Palms are most diverse in the lowland rain-
forests of the western Amazonian regions in the
Colombian Choc6, Ecuador, Peru, and the Iquitos
region of Brazil, and in the Old World in New
Guinea and the Sunda shelf, particularly parts of
the island of Borneo. A few taxa are subtropical
with the limits of palm distribution 44 °N for
Chamaerops humilis in Europe, and 48°18'S for
Rhopalostylis sapida in New Zealand.
Several palms occur in such vast natural stands
as to dominate the vegetation completely. In the
eastern tropics Nypa fruticans often occurs in
dense stands on estuarine muds with dicotyle-
donous mangrove species almost excluded by the
palm; in other situations it may occur mixed with
dicotyledonous mangrove trees. Pure stands of
Nypa may extend for several hundred hectares in
extent, especially in parts of Borneo and eastern
Sumatra. Such Nypa forests are of immense im-
portance, not only for the wealth of products that
Nypa can provide, but also in terms of gas ex-
change and mud stabilization. On the landward
fringe of the mangrove in West Malesia occurs
another vegetation type almost completely domi-
nated by palms. This is the habitat of Oncosperma
tigillarium and Calamus erinaceus; in Borneo
Daemonorops longispatha also occurs in this habi-
tat. Mauritia flexuosa occurs in vast numbers,
dominating wetlands of northern South America;
Copernicia alba in Paraguay, and adjoining parts
of Brazil, Argentina, and Bolivia forms huge
stands on areas subject to periodic flooding
and periodic drought; the largest populations
have been estimated to contain half a billion
individuals (Markley 1955). These are a few
examples of vegetation types where palms are
the most conspicuous and apparently dominant
plant. When the particular palms are useful, these
natural stands become immensely significant
either in the local economy or in international
economy.
Tropical rainforest developed on white sands
in Borneo is normally physiognomically very
distinctive. Known locally as kerangas, this
forest type is developed on extremely poor soils.
Kerangas carries a varied, peculiar, and important
palm flora. Many of the palms of kerangas are
confined to this vegetation type or to the catena
between kerangas and the neighboring diptero-
carp forests developed on richer soils. ]ohannest-
eijsmannia altifrons is confined to kerangas in
Sarawak, yet the same species occurs in diptero-
 Palmae
carp forest in Malaya. One of the delights of palm
hunting in kerangas is the unpredictable nature of
the palm flora from place to place. Elsewhere in
Southeast Asia, white sands seem not to support
such an unusual palm flora as in Borneo. Indeed,
on white sands in Sumatra and Malaya, palms are
very poorly represented.
Ultramafie rocks with their usually toxic soils
rich in heavy metals, such as Mn, Cr, Fe, and Cu,
usually support a restricted but peculiar palm
flora. Thus, in Sabah and Palawan, ultramafic soils
carry a curious assemblange of rattans and, rarely,
species of Pinanga and Areca not found on the
surrounding nonultramafic rocks. In New
Caledonia, an island rich in ultramafics, the palm
flora of the ultramafics is quite different from that
of the non ultramafic rocks.
Within the humid tropics of Southeast Asia
and Malesia it may be observed that palms
are rather rare on limestone but what palms do
occur are often of considerable interest (e.g.,
Maxburretia). In drier climates there seems to
be a greater abundance of calcicolous palms.
In the New World there is greater diversity
of palms occurring on limestone, such as
Brahea, Gaussia, Pseudophoenix, Coccothrinax,
and Thrinax spp.
Rheophytes are found in several genera in both
New and Old Worlds.
Tropical montane forests may support a rich
diverse assemblage of palms. In West Malesia
there is usually a striking vegetational change at
about lOOOm altitude, where the tall forests domi-
nated by dipterocarps are replaced by usually
shorter, even crowned forest dominated by mem-
bers of the Lauraceae and Fagaceae, in the change
from Hill Dipterocarp forest to Lower Montane
forest. This change occurs more or less at the alti-
tude above which coconuts do not fruit. The palm
flora also changes dramatically. Very few species
of the lowland and Hill Dipterocarp forest occur in
the montane forests and vice versa. Plectocomia
elongata, a plant apparently adapted to several
forest types, is one of these very few species.
Above 1000 m the genus Korthalsia does not
occur; ]ohannesteijsmannia, Eugeissona, Plecto-
comiopsis, Myrialepis, and many other genera
only very rarely transgress this vegetational
boundary. However, although there appear not
to be any genera confined to the montane
forest, there is a rich assemblage of species of
Pinanga, Areca, Calamus, Daemonorops, Caryota,
Livistona, and others. The highest elevations re-
corded for palms in the Malesian region are on G.
Kinabalu in Borneo, where Pinanga capitata
321
peters out at about 2900 m and Calamus gibbianus
reaches just over 3000 m.
There is a greater diversity of palms at high
elevations in South America than in Southeast
Asia and Malesia, and there is one genus confined
to the montane forests of the Andes, Ceroxylon.
Not only is Ceroxylon the genus with the tallest
species of palm (C. quindiuense ), it is also the ge-
nus with the highest altitudinal record, C. utile,
recorded at over 4000 m elevation in Colombia
(Dugand 1974). Besides the two genera confined
to the montane forest, there is a rich assemblage
of species of Geonoma, Aiphanes, Prestoea,
Euterpe, Dictyocaryum, Chamaedorea, Socratea,
Catoblastus, and other genera.
In contrast, there is only one species of palm,
Phoenix reclinata, occurring in montane forest in
Africa.
SuBDIVISION. The family is divided into six
subfamilies which vary widely in size. Subfamily
Coryphoideae, generally considered to include the
least specialized palms, has 40 genera and
embodies most of the palmate and costapalmate
genera and the genus Phoenix, which has a pin-
nate but induplicate leaf. The second subfamily,
the Calamoideae, is a group of 22 genera, well
established as a monophyletic group with several
synapomorphies, the best known being scaly
fruits. The third subfamily, the Nypoideae, in-
cludes only the genus Nypa, the mangrove palm.
Subfamily IV, the Ceroxyloideae, has only ten
genera divided into three distinctive and mono-
phyletic tribes, although the monophyly of the
subfamily is in question (Uhl et al. 1995). The larg-
est subfamily, the Arecoideae, includes 75 genera.
Finally the Phytelephantoideae has only three
genera which are morphologically disjunct and
add a new dimension to the family.
AFFINITIES. In many ways, palms epitomize the
monocotyledons, but their broader affinities have
never been clear. In the past, they have frequently
been associated with the Cyclanthaceae or
Pandanaceae, less commonly with Lilianae or
Arales, and more recently with the commelinoids.
Phylogenetic and molecular analyses have found
the palms a sister group to a diverse clade
that includes Bromeliales, Commelinales, and
Zingerberales (Linder and Kellog 1995). Chase
et al. (1995a) and Stevenson and Loconte (1995)
found the palms isolated as a sister group to the
Poales within the Commelinidae. Recent sequenc-
ing studies of atpA (J.I. Davis, pers. comm.) also
322 Palmae
put the palms as sister to or in some analyses in a
basal position within the Commelinidae.
FosSILS. Palms have one of the richest fossil
records among monocotyledons (Daghlian 1981;
Herendeen and Crane 1995). There are numerous
records for leaves, stems, and pollen (Harley
1996). Roots and flowers have also been found.
The oldest records for palms are from the
Magothy Formation and consist of leaves, appar- II.
ently costapalmate (Berry 1905), and stems (Berry
1916) from the late Santonian. Earliest pollen
records are of Nypa-like grains of Late Cretaceous
origin which are widespread, and reported to
occur in South America, Venezuela, and Colom-
bia, Brazil, Africa, Senegal, and the Ivory Coast.
There are many records of sulcate grains which
have recently been assessed using SEM and TEM
(Harley 1996). Some of the earliest supposedly
monocotyledonous grains have characters that
show possible affinities with palms.
ECONOMIC IMPORTANCE. In economic impor-
tance palms are second only to grasses and possi-
bly legumes. Harold E. Moore, Jr. wrote in 1973:
"The versality of palms in the hands of man is
astonishing. Houses, baskets, mats, hammocks, III.
cradles, quivers, packbaskets, impromptu shel- IV.
ters, blowpipes, bows, starch, wine, protein from
insect larvae, fruit beverages, flour, oil, orna-
ments, loincloths, cassava graters, medicines,
magic, perfume - all are derived from palms ... to
whatever extent man has been involved in the
tropical ecosystem, palms have certainly been a V.
major factor in making possible this involve-
ment ... " Information on utilization of palms can
be found in numerous books including Balick
et al. (1990) and Johnson (1991, 1996); Dransfield
and Manokaran (1993) provide information on
rattans.
CoNSERVATION. Some 224 species belonging to
63 genera are endangered (Johnson 1996). Habitat
destruction and over-exploitation are the main
threats to palms. Those endemic on islands, as in
Madagascar, are particularly at risk. An action
plan has been developed by the IUCN/SSC palm
specialist group.
CONSPECTUS OF CLASSIFICATION OF PALMS
I. Subfamily Coryphoideae Griff. (1844}.
1. Tribe Corypheae Mart. (1837}.
a. Subtribe Thrinacinae Becc. (1907}.
Genera 1-14.
b. Subtribe Livistoninae Saakov (1954}.
Genera 15-26.
c. Subtribe Coryphinae Becc. (1907}.
Genera 27-30.
d. Subtribe Sabalinae Mart. (1837}.
Genus 31.
2. Tribe Phoeniceae Drude (1881}.
Genus 32.
3. Tribe Borasseae Mart. (1837}.
a. Subtribe Lataniinae Meisner (1842}.
Genera 33-36.
b. Subtribe Hyphaeninae Becc. (1924).
Genera 37-40.
Subfamily Calamoideae Griff. (1844).
1. Tribe Calameae Drude (1881}.
a. Subtribe Ancistrophyllinae Becc. (1918}.
Genera 41-42.
b. Subtribe Eugeissoninae Becc. (1918).
Genus 43.
c. Subtribe Metroxylinae Bl. (1843}.
Genera 44-45.
d. Subtribe Calamineae Meisner (1842).
Genera 46-53.
e. Subtribe Plectocomiinae J. Dransf. & N. Uhl (1986).
Genera 54-56.
f. Subtribe Pigafettinae J. Dransf. & N. Uhl (1986).
Genus 57.
g. Subtribe Raphiinae J.D. Hook. (1883).
Genus 58.
h. Sub tribe Oncocalaminae J. Dransf. & N. Uhl (1986}.
Genus 59.
2. Tribe Lepidocaryeae Mart. ( 183 7).
Genera 60-62.
Subfamily Nypoideae Griff. (1850).
Genus 63.
Subfamily Ceroxyloideae Drude (1881).
1. Tribe Cyclospatheae O.F. Cook (1902}.
Genus 64.
2. Tribe Ceroxyleae Satake (1962}
Genera 65-68.
3. Tribe Hyophorbeae Drude (1881).
Genera 69-73.
Subfamily Arecoideae Griff. (1845).
1. Tribe Caryoteae Drude (1881}.
Genera 74-76.
2. Tribe Iriarteeae Drude (1881).
Genera 77-82.
3. Tribe Podococceae J. Dransf. & N. Uhl (1986}.
Genus 84.
4. Tribe Areceae Drude (1881}.
a. Subtribe Oraniinae J. Dransf. & N. Uhl (1986).
Genus 83.
b. Subtribe Manicariinae J. Dransf. & N. Uhl (1986).
Genus 84.
c. Subtribe Leopoldiniinae J. Dransf. & N. Uhl (1986}.
Genus 85.
d. Subtribe Malortieinae J.D. Hook. (1883}.
Genus 86.
e. Subtribe Dypsidinae Becc. (1914}.
Genus 87.
f. Subtribe Lemurophoenicinae J. Dransf. (1995).
Genus 88.
g. Subtribe Euterpeinae J. Dransf. & N. Uhl (1986).
Genera 89-93.
h. Subtribe Roystoneinae J. Dransf. & N. Uhl (1986}.
Genus 94.
i. Subtribe Archontophoenicinae J. Dransf. & N. Uhl
(1986).
Genera 95-100.
 Palmae
j. Subtribe Cyrtostachydinae J. Dransf. & N. Uhl
(1986).
Genus 101.
k. Subtribe Linospadicinae J.D. Hook. (1883).
Genera 102-105.
I. Subtribe Ptychospermatinae J.D. Hook. (1883).
Genera 106-114.
m. Subtribe Arecinae. J.D. Hook. (1883).
Genera 114-122.
n. Subtrie Iguanurinae J.D. Hook. (1883).
Genera 123-149.
o. Subtribe Oncospermatinae J.D. Hook. (1883).
Genera 150-157.
p. Subtribe Sclerospermatinae J. Dransf. & N. Uhl
(1986).
Genus 158.
q. Subtribe Masoalinae Beentje & J. Dransf. (1995).
Genera 159-160.
5. Tribe Cocoeae Mart. (1837).
a. Subtribe Beccariophoenicinae J. Dransf. & N. Uhl
(1986).
Genus 161.
b. Subtribe Butiinae Saakow (1954).
Genera 162-171.
c. Subtribe Attaleinae Drude (1887).
Genus 172.
d. Subtribe Elaeidinae J.D. Hook. (1883).
Genera 173-174
e. Subtribe Bactridinae J.D. Hook. (1883).
Genera 175-180.
6. Tribe Geonomeae Drude (1881).
Genera 181-186.
VI. Subfamily Phytelephantoideae Drude (1887).
Genera 187-189.
KEY TO THE SUBFAMILIES OF PALMAE
The following key uses only characters which will
distinguish all members of each subfamily. More
comprehensive diagnoses and commentary are
given under the separate treatments.
1. Leaves palmate or costapalmate, rarely entire; induplicate,
rarely reduplicate (but then apocarpous), or mixed
induplicate-reduplicate (Licuala), or pinnate but indu-
plicate and the leaflets with entire tips (Phoenix); flowers
solitary or clustered, never in triads of a central pistillate
and 2 lateral staminate I. Coryphoideae
- Leaves pinnate, bipinnate, or entire and pinnately ribbed,
or rarely palmate but then reduplicate and flowers syncar-
pous, reduplicate, or rarely induplicate, but then leaflets
with praemorse tips; flowers solitary or clustered,
frequently in triads 2 family exhibits traits that are considered basic
2. Ovary and fruit covered in imbricate scales; flowers
hermaphrodite or unisexual but only rarely dimorphic,
arranged singly or in dyads II. Calamoideae
- Ovary and fruit glabrous or with peltate or basifixed scales,
hairs, corky warts, or spines, but not with imbricate scales;
flowers hermaphrodite or unisexual, often dimorphic,
borne singly or in triads, or in pairs derived from triads 3
3. Pistillate flower borne in a terminal head, each flower with
3( -4) free, large, asymmetrical carpels and 6 minute peri-
anth segments; staminate flowers crowded on spikes at the
tips of inflorescence branches below the pistillate head, each
flower with 6 linear distinct perianth segments and 3 an-
thers borne on a solid stalk III. Nypoideae
323
- Pistillate flowers not borne in a terminal head or, if so, then
plants dioecious and flowers multiparted; staminate flowers
with stamen filaments free or variously connate, very rarely
forming a solid stalk 4
4. Pistillate flowers borne in a head on a short peduncle, very
large, each with numerous, spirally arranged sepals and
petals and an elongate style, gynoecium 5-10-locular and
ovulate; fruit corky-tuberculate with 5-10 seeds; staminate
flowers large, sessile or stalked, receptacle fiat or clublike,
with reduced uniseriate perianth or perianth scarcely dis-
cernible, stamens very numerous VI. Phytelephantoideae
- Pistillate flowers not borne in a head, both staminate and
pistillate flowers with sepals and petals in 2 whorls, style
usually short or not differentiated, locules 1-3, rarely more
5
5. Flowers usually unisexual, rarely hermaphrodite, borne sin-
gly or in lines (acervuli), very rarely in groups of3, but then
staminate flowers above the pistillate and peduncular bracts
numerous IV. Ceroxyloideae
- Flowers always unisexual, borne in triads or in pairs derived
from triads, very rarely the staminate flowers above the
pistillate, but then peduncular bract 1 V. Arecoideae
I. Subfamily Coryphoideae Griff. (1844).
Leaves palmate or costapalmate, only rarely un-
divided or pinnate, almost always strictly in-
duplicate, but interfold splits occurring in two
genera and leaves reduplicate in Guihaia and par-
tially so in Licuala; flowers solitary or in cincinni,
never borne in triads; carpels free or variously
connate.
Within this subfamily there are very few
differences in leaf form or venation. The only
exceptions in form are the undivided leaves of
]ohannesteijsmannia and a few species of Licuala,
and, of course, the regularly pinnate but in-
duplicate leaf of Phoenix. There are a few excep-
tions to induplicate folding. The leaves of Guihaia
are palmate and reduplicate; and those of Rhapis,
Rhapidophyllum, and Licuala show anomalous
splitting superimposed on a basic induplicate
structure. (It is noteworthy that elsewhere in the
family the only palmate leaves are reduplicate and
belong to three genera of Calamoideae: Lepido-
caryum, Mauritia, and Mauritiella.)
In inflorescence and flower structure the sub-
in palms and to some extent also in monocotyle-
dons. The interfoliar or suprafoliar, often highly
branched inflorescences show considerable adna-
tion of branches and reduction to spicate forms.
Some of the simplest of all palm flowers belong
to the first tribe, the Corypheae. These truly
apocarpous flowers, having 3 sepals, 3 similar
petals, 6 stamens in 2 whorls of 3, and 3 follicular
carpels, embody a structure considered unspeci-
alized within the monocotyledons as a whole.
With the exception of Nypa, all apocarpous palms
belong to the Coryphoideae.
324
KEY TO THE TRIBES OF THE CORYPHOIDEAE
1. Leaves palmate, costapalmate, or entire; acanthophylls
absent 2 - Leaves reduplicate. China, Vietnam
- Leaves pinnate; basal leaflets modified as spines
2. Phoeniceae (p. 333)
2. Hermaphroditic, or polygamodioecious, rarely strictly dio-
ecious; if dioecious, flowers not or only slightly dimorphic;
rachillae lacking deep pits; endocarp usually thin, crusta-
ceous, or cartilaginous 1. Corypheae
- Dioecious; flowers usually strongly dimorphic; staminate
and sometimes pistillate flowers borne in deep pits formed
by connation and adnation of rachilla bracts; endocarp very
thick and hard 3. Borasseae (p. 334)
1. Tribe Corypheae Mart. (1837).
Leaves palmate, costapalmate, or undivided,
induplicate (except reduplicate in Guihaia); inflo-
rescence with many peduncular bracts, rarely with
prophyll only, rachis bracts usually well devel-
oped; rachillae lacking deep pits; fruit usually 1-
seeded, if more then bi- or trilobed; endocarp
usually thin, crustaceous or cartilaginous, or not
well differentiated.
The members of this tribe are distinct from
other palms in the combination of induplicately
palmate or costapalmate leaves and hermaphro-
ditic or, when unisexual, only slightly dimorphic
flowers borne singly or in cincinni. Systematically,
these 31 genera are of primary importance for two
reasons: first, because they include the palms with
the simplest floral morphology, and second,
because they provide a microcosm of the struc-
tural diversity seen elsewhere in the family. Some
exceptions are the absence of pinnate leaves, of
infrafoliar and multiple inflorescences, of flowers
multiparted in all organs, and of adjacent-ligular
germination of the seed. The four subtribes
are based on the nature of the gynoecium. Not
surprisingly, cladistic analyses of anatomy and
morphology (Uhl et al. 1995; Barrow 1996) have
found that the 14 apocarpous genera in the
Thrinacinae do not form a monophyletic group.
Realignments of the genera of Corypheae can be
expected.
KEY To THE SuBTRIBES oF THE CoRYPHEAE
1. Carpels distinct, usually 3(1-4) Ia. Thrinacinae
- Carpels connate
2. Carpels basally distinct, united by their styles only
I b. Livistoninae
- Carpels united basally or throughout
3. Carpels united basally, styluli free or united, if united then
with separate stylar canals I c. Coryphinae
- Carpels united throughout, the style with a common stylar
canal I d. Sabalinae
Palmae
KEY TO THE GENERA OF THE CORYPHEAE
1. Leaves induplicate 2
13. Guihaia
2. Gynoecium of 1-3 free carpels, stamens 5-25 3
- Gynoecium of 3 carpels, connate by thin erect styles alone,
or up to their midregions, or completely connate; stamens
6 15
3. Perianth biseriate; carpels 1-3; stamens 5-25 4
- Perianth uniseriate, usually 6-toothed; carpel 1; stamens
5-15 13
4. Carpel! 5
- Carpels 2, 3, or 4 6
5. Flowers with long stalklike base, proximal flowers on each
rachilla hermaphroditic, distal flowers staminate; stamens
6. Belize 5. Schippia
- Flowers lacking a long stalklike base, all hermaphroditic;
stamens 18-24. Peru, Brazil 4. Itaya
6. Stamen filaments distinct 7
- Stamen filaments connate in a tube or upper 3 filaments
epipetalous 10
7. Flowers solitary 8
- Flowers in cincinni of 2-4 9
8. Leaf blade not split centrally to the rachis (but deeply
divided in Trithrinax biflabellata); sepals united ca. 1/3 of
their length; stamens with distinct elongate filaments, long
exserted, more than twice as long as the corolla; seed with
lobed intrusion of the seed coat. Brazil to Argentina
1. Trithrinax
- Leaf blade with the central split reaching to the rachis,
segments wedge-shaped; sepals imbricate, free nearly to
the base; stamens with fleshy wide filaments, only slightly
exserted; seed without conspicuous lobed intrusion of the
seed coat. Amazonian Bolivia, Peru, Brazil, and western
Colombia 2. Chelyocarpus
9. Leaf sheaths unarmed, fibrous, blade splitting along
adaxial ribs to varying lengths; inflorescences exserted
from leaf sheaths; seed kidney-shaped or oblong and
grooved on the rapheal side, seed coat deeply intruded.
Temperate and subtropical E Asia 9. Trachycarpus
- Leaf sheaths armed with stout needlelike fibers; leaf blade
usually splitting between the folds; inflorescences hidden
among the leaf sheaths; seed oblong, the seed coat thick-
ened below the raphe but not conspicuously intruded.
Southeastern USA 10. Rhapidophyllum
10. Flowers polygamodioecious; stamen filaments connate in
a flared cupule; endosperm ruminate. Mediterranean
11. Chamaerops
- Flowers polygamodioecous or hermaphroditic; stamen
filaments connate in an erect cupule or epipetalous;
endosperm homogeneous 11
11. Stem bearing conspicuous root spines at the base; calyx
lobes nearly as long as the petals; stamen filaments con-
nate in an elongate erect cupule not attached to the petals,
anthers reflexed and radiating at anthesis; carpels with
attenuate styles. Central America and Colombia
3. Cryosophila
- Stem without root spines; calyx lobes much shorter than
2 the petals; stamen filaments distinct, epipetalous or the
antesepalous filaments free, or all filaments connate in an
epipetalous cupule but anthers scarcely exserted; carpels
3 without attenuate styles 12
12. Stems slender, reedlike; leaves divided between the folds
into many-fold, truncate segments; perianth whorls fleshy,
sepals connate; stamen filaments free, epipetalous. S
China, Indochina, northern Sumatra 14. Rhapis
 Palmae
- Stems scarcely reedlike, stouter; leaves divided along
adaxial folds into single-fold, acute or bifid segments; peri-
anth whorls coriaceous; sepals distinct, imbricate; stamen
filaments connate in an epipetalous cupule (M. gracilis, M.
furtadoana), or separate and epipetalous (M. rupicola).
Thailand and Malay Peninsula 12. Maxburretia
13. Base of leaf sheath split in petiolar region, inflorescence
emerging through the split; seed umbilicate, excavate, or
completely perforate below the raphe, endosperm not
furrowed. Caribbean 6. Thrinax
- Base ofleaf sheath not split in petiolar region; seed deeply
perforated by furrows 14
14. Leaf sheaths disintegrating into a regular fibrous network,
fiber tips reflexed, spinelike; flowers subdistichous on
the rachillae; fruit white at maturity; seed deeply
bilobed, the lobes again irregularly bilobed. Hispaniola
8.Zombia
- Leaf sheath fibers not reflexed or spinelike; flowers borne
spirally; fruit tawny yellow, pink, red, or black; seed deeply
furrowed, not bilobed. Caribbean 7. Coccothrinax
15. Gynoecium of 3 carpels, free at the base, connate by their
styles alone 16
- Gynoecium of 3 completely fused carpels or carpels con-
nate basally and styluli free 27
16. Leaf blades undivided, diamond-shaped, pinnately
nerved, the costa prominent; petiole and basal margins of
the blade armed with hooked teeth; inflorescences each
with several inflated peduncular bracts; fruit warty;
endosperm deeply and irregularly intruded by the seed
coat. Malay Peninsula, Sumatra, Borneo
17. ]ohannesteijsmannia
- Leaf blades divided, or, if undivided, not diamond-shaped
and the basal margins of the blade unarmed; fruit smooth
or, if corky-warted, then leaf divided 17
17. Leaf blades divided along abaxial folds to the petiole into
single-fold (rarely) to several-fold, usually wedge-shaped,
truncate segments, or undivided. China, SE Asia, Malesia
to Australia 18. Licuala
- Leaf blades divided along adaxial folds into single-fold or
several-fold but not truncate segments 18
18. Inflorescence with long peduncle, branched only at its tip
or branched at the very base to produce 2-4 more or Jess
equal long axes, also branched only at their tips, giving the
appearance of more than 1 inflorescence per axil; corolla
lobes caducous at anthesis exposing the prominent
androecial tube; seed coat not intruded. Fiji, Tonga, Dan-
ger Islands, Hawaiian Islands 20. Pritchardia
- Inflorescence branching not confined to the tip; corolla
lobes not caducous at anthesis, very rarely tardily
deciduous 19
19. Rachis bracts split on one side and pendulous, sword-
shaped; petals large, flat, chaffy, without grooves on the
inner surfaces, becoming strongly reflexed at anthesis;
style elongate, about 3 times as long as the ovarian part of
the gynoecium; flowers and fruit pedicellate; fruit with
persistent chaffy calyx. Southwestern USA and northwest-
ern Mexico 26. Washingtonia
- Inflorescence bracts tubular, not splitting into sword-
shaped blades at anthesis; petals neither flat nor chaffy and
with grooves matching anthers on inner surfaces; styles as
long as the ovarian part or rarely 2-2 1/ 2 times as long
(Serenoa) 20
20. Fruit large and smooth (ca. Scm in diameter at maturity);
mesocarp spongy with short fibers near and closely
adhering to the endocarp, endocarp with a triangular
basal projection rounded on 1 side, keeled on the other 3
325
sides; embryo eccentrically apical, endosperm with a
conspicuous, lobed intrusion of the seed coat. New
Caledonia 19. Pritchardiopsis
- Fruit various, if large, then usually corky-warted;
endocarp smooth, not keeled, without a basal projection;
embryo lateral or basal, seed coat intrusion present or not
21
21. Petioles and often margins and/or veins of leaves with
stout teeth; abortive carpels apical in fruit; seeds with
ruminate endosperm. West Indies, S America
5. Copernicia
- Petioles armed or not, leaves lacking teeth; abortive
carpels usually basal in fruit; seeds with homogeneous
endosperm, sometimes penetrated by a lobed intrusion
of the seed coat 22
22. Endosperm intruded by the seed coat 25
- Endosperm not intruded by the seed coat 23
23. Trunk solitary, sometimes ventricose at or near the
middle; petiole unarmed; androecial tube projecting
beyond the corolla lobes. Cuba, Guatemala, Panama
21. Colpothrinax
- Trunks clustered, erect, or creeping and branching; petiole
spiny, armed with fine or coarse teeth; androecial tube not
projecting beyond the corolla lobes 24
24. Stems erect, densely clustered; flowers in cincinni, but
paired or solitary towards the apex of the rachillae; sepals
distinct or basally shortly connate, imbricate; stamen fila-
ments united in a basal ring, free portions short, abruptly
narrowed; fruit globose, raphe short, scarcely more than
1/2 as long as the seed. Southern USA, Central America
22. Acoelorrhaphe
- Stems prostrate, rarely erect; flowers solitary or occasion-
ally paired; calyx tubular, shortly 3-lobed; stamen fila-
ments shortly united at the base, free portions gradually
narrowed; fruit ellipsoidal; raphe extending the length of
the seed. Southern USA 23. Serenoa
25. Sepals distinct, imbricate. Western USA, Mexico, Guate-
mala 24. Brahea
- Sepals united basally 26
26. Blade divided by deep splits into compound segments,
further divided by shallow splits into single-fold segments;
androecial ring conspicuous, thick, almost free at the base,
filaments, where separate, slender not broadly rounded;
gynoecium top-shaped; fruit very large (at least 6cm in
diameter at maturity) usually corky-warted (smooth in P.
kingianus) Malesia 16. Pholidocarpus
- Blade regularly divided into single-fold segments, very
rarely into compound segments (L. saribus); androecial
ring broadly scalloped apically, epipetalous; gynoecium
widest above the locules, abruptly narrowed to the style;
fruit small (rarely exceeding 4 em at maturity), smooth. NE
Africa, Arabia, Ryukyu Islands, SE Asia to Australia
15. Livistona
27. Dioecious; leaf blade dark green on upper surface, gray-
white beneath; all inflorescence axes with a thick layer
of thick layer of multicellular hairs; petals inserted on a
floral stalk well above the calyx; carpels basally connate by
ovarian regions only; styles free, widely separated; en-
dosperm shallowly ruminate. Thailand 30. Kerriodoxa
- Hermaphroditic; leaves not gray-white beneath; rachillae
variously tomentose or glabrous; floral stalks present or
poorly developed; carpels connate by their ovarian regions
and usually through the styles; endosperm homogeneous
or ruminate 28
28. Clustering palms; hastulae absent; rachilla bracts tubular
29
326 Palmae

- Solitary palms; adaxial hastula present; rachilla bracts
minute, triangular 30
29. Slender pleonanthic palms; inflorescences spicate or
branching to 2 orders; endosperm homogeneous or
ruminate. Indochina and China 29. Chuniophoenix
- Moderate hapaxanthic palm, acaulescent or with erect,
often branching stems; inflorescence paniculate, highly
branched; endosperm homogeneous. Western Asia and
Arabia 28. Nannorrhops
30. Massive hapaxanthic palms; petiole margins armed with
teeth; flowers in adnate cincinni; gynoecium 3-grooved,
abruptly narrowed to the style, stylar canals 3. SE Asia to
Australia 27. Corypha
- Acaulescent to robust, erect, pleonanthic palms; petioles
unarmed; flowers solitary; gynoecium shallowly 3-lobed,
style elongate, broad, only slightly narrower than the ova-
rian part, stylar canal single. Southern USA to Venezuela
31. Saba!
Solitary or clustering, hermaphroditic; distal
sheath fibers becoming sharp spines; blade fan-
shaped to nearly circular, costapalmate, divided to
beyond the middle (T. flabellata almost to base)
into single-fold stiff or flexible, bifid segments;
prophyll and the 2( -3) peduncular bracts inflated.
Flowers solitary, sepals 3, connate basally; petals 3,
longer than sepals, acute; stamens 6, filaments
twice as long as petals, anthers versatile; carpels
attenuate to a long stylulus, stigma apical; ovule
hemianatropous with aril. Postament large. About
four spp., subtropical S America in Bolivia, west-
ern and southern Brazil, Paraguay, Uruguay, and
Argentina.

2. Chelyocarpus Damm. Fig. 81G, 83D-F

la. Subtribe Thrinacinae Becc. (1907). Chelyocarpus Damm., Notizbl. Bot. Gart. Berlin-Dahlem 7:
395. (1920); Moore, Principes 16: 67-88 (1972); Kahn &
Pleonanthic, leaves palmate or costapalmate, Mejia, Principes 32: 69-72 (1988).
induplicate but reduplicate in Guihaia; adaxial
hastula well developed, often deltoid, abaxial small Solitary or clustering, unarmed, hermaphroditic.
or lacking; inflorescences interfoliar, each bearing Leaves palmate, or costapalmate, petiole not split-
a tubular, bicarinate prophyll and 0 to usually sev- ting basally; blade deeply divided, each half
eral peduncular bracts; anthers latrorse, pollen el- divided into paired or irregularly grouped, single-
liptic, sulcate (pantoperculate in Chamaerops, fold segments, or divided into many-fold seg-
occasionally trichotomosulcate in Cryosophila), ments, split distally into bifid, single-fold,
with finely reticulate or rarely otherwise, tectate 1-ribbed segments. Rachillae sometimes fascicu-
exine; carpels free, usually 3(1-4). Endosperm ho- late. Flowers solitary; sepals 2, 3, or 4, slightly con-
mogeneous, ruminate in Chamaerops. Eophylls nate or imbricate basally; petals like the sepals;
simple, bifid in Chelyocarpus. stamens 5, 6, 7, 8, or 9, filaments distinct, fleshy;
A group of 14 genera, largely confined to the carpels 3(1-4), follicular, ovule hemianatropous
subtropics, and including some of the most with aril. Epicarp smooth or coarsely corky-
drought- and cold-tolerant palms. Many genera warted. Germination remote-tubular. Four spp.,
grow only on limestone. Within this highly diverse Amazonian Bolivia, Brazil, Peru, and western
group are genera exhibiting some of the least Colombia in understory of upland forests or on
specialized features in the family. Two main lines alluvial deposits left by flooding rivers.
of evolution seem evident. One is entirely New
World, including genera 1-8, and has been found
monophyletic by cladistic analysis of morphologi- 3. Crysophila Blume
cal and molecular datasets (Uhl et al. 1995, Barrow Crysophila Blume, Rumphia 2: 53 (1838, "1836"); Evans, Syst.
1996). Evolutionary trends are in the reduction to a Bot. Monographs 46: 1-70 (1995).
single carpel and to a cupular perianth. The
second line includes genera 9-14 and, except for Solitary, hermaphroditic. Stem bearing root
Rhapidophyllum, is strictly Old World. In this spines. Leaves palmate, sheaths splitting basally;
group the main trends of specialization are to blade divided centrally nearly to base, each half
dioecy and elaboration and fusion of perianth divided into many-fold segments, further divided
members. Guiaha is unusual in having reduplicate into 2 or more single-fold acute or bifid segments;
leaves, and Rhapis and Rhapidophyllum have white-tomentose abaxially. Peduncular bracts 6 or
leaves irregularly split in reference to plications. more, much larger than the prophyll. Flowers soli-
tary; sepals 3, connate basally; petals 3, distinct;
filaments flat, connate to 1/2 their length; carpels
1. Trithrinax Mart.
3, follicular, styluli attenuate; ovule campy-
Trithrinax Mart., Hist. Nat. Palm. 2: 149 (1837); Henderson lotropous with aril. Postament small. Germination
et al., Field Guide: 37 (1995). adjacent-ligular. Ten spp., western Mexico to
northern Colombia.
 Palmae 327

4. Itaya H.E. Moore

Itaya H.E. Moore, Principes 16: 85 (1972); F. Kahn, Palms of
Eldorado: 112 (1997).

Moderate, solitary, unarmed, hermaphroditic.

Leaves palmate; petiole split basally; blade orbicu-
lar, divided centrally to ca. 3/4 the radius at the
middle, each half divided into 4-7 wedge-shaped,
4-7-fold segments, these divided apically into
bifid, 1-fold segments. Flowers solitary, sepals
3, connate; petals 3, connate ca. 1/2 their length;
stamens 18-24, filaments fleshy, connate basally;
gynoecium of 1 carpel, eccentrically ovoid, ovule
hemianatropous, aril large, oblique. Epicarp
minutely granular-roughened; mesocarp thick,
white, splitting the epicarp when ripe. Only one
sp., I. amicorum Moore, known from one locality
in Amazonian Peru and one in Brazil, occurring in
seasonal rainforest at low elevations.

5. Schippia Burret
Schippia Burret, Notizbl. Bot. Gart. Berlin-Dahlem 11:867-869
(1933); Henderson eta!., Field Guide: 42 (1995).

Solitary, unarmed, hermaphroditic or polyga-

momonoecious. Leaves palmate to costapalmate;
sheath split basally, disintegrating into a fibrous
network; blade divided into narrow single-fold
segments, with pointed, unequal, bifid apices.
Flowers solitary, either hermaphroditic or
staminate, calyx and receptacle forming a
pseudopedicel, calyx tubular basally with 3
triangular-lanceolate lobes; petals 3, distinct,
much larger than calyx; stamens 6; gynoecium
unicarpellate, ovoid, style elongate, tubular, open
distally; ovule probably hemianatropous. Only
one sp., S. concolor Burret in Belize and possibly
adjacent Guatemala.
Fig. 83A-Q. Palmae-Coryphoideae. A- C Chamaerops
6. Thrinax 0. Swartz Fig. 83G humilis. A Pistillate flower; note free styluli. B Staminate
flower. C Staminate flower opened out, showing androecial
Thrinax 0. Swartz, Prod. 4: 57 (1788); Read, Smithsonian ring. D-F Chelyocarpus ulei. D Flower. E Vertical section of
Contr. Bot. 19: 1-98 (1975); Borhidi & Muniz, Acta Bot. flower. F Fruit showing pericarp split into corky warts. G
Hung. 31: 225-230 (1985). Thrinax radiata, flower showing single perianth whorl and
Hemithrinax Hook.f. (1883). single carpel with funnel-shaped stigma. H, I Nannorrhops
ritch iana. H 3-flowered cincinnus. I flower. J-R Sabal pal-
metto. J Habit. K Stem with persistent split leaf bases. L First-
Solitary, unarmed, hermaphroditic. Leaves pal-
order branch of inflorescence showing tubular bracts; note
mate, often irregular; sheath split opposite petiole prophylls on first- and second-order branches. M Flower. N
and abaxially; blade often irregularly folded; seg- Pistil. 0 Ovary in cross-section showing three locules. P
ments unsplit in basal about 1/2, bifid apically. Rachilla with two fruit. Q Seed. R Seed in vertical section
Inflorescences slender, emerging through petiole showing lateral embryo, homogeneous endosperm, and very
shallow postament. (Takhtajan 1982)
split. Flowers solitary; perianth a 6-lobed cupule;
stamens 6-12 (5-15), filaments slender or lacking,
sometimes partly united basally; gynoecium of 1
328
carpel, unilocular, uniovulate; ovule campy-
lotropous, with basal aril. Fruit white. Germina-
tion remote-tubular. Seven spp., two endemic to
Jamaica, T. radiata widely distributed in littoral
habitats of Atlantic northern Caribbean. The three
species formerly of Hemithrinax are all very local
endemics in Cuba.
7. Coccothrinax Sargent
Coccothrinax Sargent, Bot. Gaz. 27:87 (1899); Bailey & Moore,
Gentes Herb. 8:93-205 (1949); Read, Phytologia 46:285-287
(1980); Henderson eta!., Field Guide: 45-50 (1995).
Solitary or clustering, hermaphroditic. Leaf
sheaths disintegrating into a regular fibrous net-
work, or long fibers, or stout spines. Leaves,
palmate; petiole unsplit basally; blade often ir-
regularly folded, divided about midlength into
pointed segments. Flowers solitary; perianth shal-
lowly cupshaped with several (5-9) short points;
stamens 9(6-13), filaments connate basally; gyno-
ecium of 1 carpel, unilocular, uniovulate, globose
basally; style long, terminating in a cuplike stigma;
ovule basal, erect, nearly atropous. Fruit purplish
black. About 49 spp. described but probably fewer,
on islands of the West Indies, with the greatest
diversity (about 34 spp.) in Cuba.
8. Zombia L.H. Bailey
Zambia L.H. Bailey, Gentes Herb. 4: 240 (1939); Henderson,
Field Guide: 50-51 (1995).
Clustering, hermaphroditic. Leaves palmate;
sheath a network of fibers, distal fibers reflexed
forming spines; blade irregularly divided 1/2-
2/3 length into single-fold, bifid segments, dis-
colorous. Flowers solitary; perianth cup-shaped
with 6 points; stamens 9-12; filaments distinct;
pollen with exine foveolate; gynoecium obpyri-
form, unicarpellate, stigma cuplike, ovule basal,
atropous. Fruit white, large. Seed deeply bilobed,
the lobes again divided; embryo central, en-
dosperm homogeneous within lobes. Only one sp.,
Z. antillarum L.H. Bailey, Hispaniola.
9. Trachycarpus H. Wendl.
Trachycarpus H. Wend!. in J. Gay, Bull. Soc. Bot. de France 8:
429 (1863? "1861"), Beccari, Ann. Roy. Bot. Gard. Calcutta
13: 1-356 (1931); Kimnach, Principes 21: 155-160 (1977);
Gibbons & Spanner, Principes 42: 24-29 (1998).
Solitary or clustering, acaulescent or erect,
unarmed or armed, dioecious or polygamous.
Palmae
Leaves palmate; petiole toothed or unarmed; blade
fan-shaped to circular, segments single-fold, bifid.
Prophyll inflated, peduncular bracts 1-3. Flowers
solitary or in clusters of 2-3, similar in both sexes;
sepals 3, connate basally; petals 3, distinct, imbri-
cate; stamens 6, filaments distinct, fleshy; sta-
minodes present or not; carpels 3, distinct,
follicular, styluli short, ovule hemianatropous,
with fleshy aril; pistillodes similar to carpels. Fruit
kidney-shaped to oblong. Postament present. Ger-
mination remote-tubular. Six spp., ranging from
the Himalayas to northern Thailand and China.
10. Rhapidophyllum H. Wendl. & Drude
Rhapidophyllum H. Wend!. & Drude in Drude, Bot. Zeit. 34:
803 (1876); Shuey & Wunderlin, Principes 21:47-59 (1977).
Clustering, dioecious, polygamodioecious, rarely
monoecious. Stem decumbent or erect, upper 3/4
with stout needlelike spines and fibers from leaf
sheath. Leaves shortly costapalmate; divided
between the folds nearly to base into linear
segments. Inflorescences short, robust. Flowers
2-4 in cincinni. Staminate flowers globose, sepals
3, distinct to slightly connate at the base; petals 3,
distinct, fleshy, imbricate; stamens 6, pistillodes 3,
resembling the carpels. Pistillate flowers similar,
carpels 3, distinct, follicular, basally tomentose,
styles short, recurved, ovule hemianatropous.
Fruit ovoid, with deciduous hairs. Only one sp., R.
hystrix (Pursh) H. Wendl. & Drude, southeastern
United States.
11. Chamaerops L. Fig. 83A-C
Chamaerops L., Sp. Pl.: 1187 (1753); Uhl & Dransf., Gen. Palm.:
184-186 (1987).
Clustering, acaulescent or shrubby, polygamous
or dioecious. Leaves palmate; sheath disintegrat-
ing into fine fibers; petiole with spines pointing
distally; blade divided 2/3-3/4 into single-fold,
deeply bifid segments. Staminate, pistillate, and
hermaphroditic inflorescences and flowers simi-
lar; peduncular bracts absent; rachillae short,
crowded, glabrous. Flowers solitary, sepals 3,
triangular, basally connate; petals 3, imbricate,
basally connate; stamens 6, filaments broad trian-
gular, connate in a conspicuous staminal ring;
pollen pontoperaculate; carpels 3, distinct, follicu-
lar; ovule hemianatropous. Endosperm ruminate;
postament conspicuous. Germination remote-
tubular. Only one sp., C. humilis L., native to
coastal areas of the western Mediterranean, both
 Palmae
in Europe and north Africa, becoming rarer east-
wards to Malta.
12. Maxburretia Furtado
Maxburretia Furtado, Gard. Bull., Singapore 11: 240 (1941);
Dransfield, Gentes Herb. 11: 187-199 (1978).
Small, clustering, acaulescent or shrubby, her-
maphroditic or dioecious. Leaves palmate; sheaths
fibrous, fibers sometimes becoming spines; petiole
unarmed; blade divided to ca. 2/3 into bifid single-
folds. Flowers small, solitary or in groups of 2-3;
staminate and pistillate similar; sepals 3, distinct,
imbricate; petals 3, basally connate; stamens 6,
adnate to petals; filaments forming a cupule, or
distinct; carpels 3, briefly joined basally, styluli
triangular; ovules anatropous to hemianatropous
with basal funicular arils; pistillode minute, 3-
lobed. Postament thin. Three spp., Malaya and
Peninsular Thailand.
13. Guihaia J. Dransf., S.K. Lee & F.N. Wei
Guihaia J. Dransf., S.K. Lee & F.N. Wei, Principes 29:7 (1985),
Dransfield eta!., Principes 29: 3-12 (1985).
Clustering, dioecious. Stem short, decumbent or
erect. Leaf sheath disintegrating into erect, spine-
like fibers or a tongue-shaped lattice of flat fibers;
petiole unarmed. Blade orbicular or cuneate,
divided to 3/4 to 4/5 into single or rarely 2-fold
reduplicate, bifid segments; discolorous. Stami-
nate and pistillate inflorescences and flowers
similar; peduncular bracts absent. Flowers
solitary, extremely small; sepals 3, united at base;
petals longer, basally connate; stamens 6, the
filaments distinct, adnate to corolla, anthers
didymous; pistillode absent; carpels abruptly
narrowed. Germination remote-tubular. Two
spp., south China and northern Vietnam.
14. Rhapis L. f. ex W. Aiton
Rhapis L. f. ex W. Aiton, Hort. Kew. 3: 473 (1789); Beccari,
Ann. Roy. Bot. Gard. Calcutta 13: 1-356 (1931).
Clustering, unarmed, dioecious or polygamo-
dioecious. Stems reedlike, leaf sheaths persistent,
fibrous. Leaves palmate or deeply bifid, divided
between the folds into segments varying in num-
ber of ribs. Inflorescences short; peduncular
bracts absent. Flowers solitary, staminate flowers
with calyx thick, cupshaped; corolla fleshy tubu-
lar, appearing stalked basally; stamens 6; filaments
329
elongate, adnate along most of corolla tube;
pistillode minute. Pistillate and hermaphrodite
flowers similar; carpels 3, distinct, wedge-shaped;
ovule hemianatropous with basal aril. Postament
lateral. Germination remote-tubular. About 12
spp., southern China, south through Indochina to
peninsular Thailand and northern Sumatra.
lb. Subtribe Livistoninae Saakov (1954).
Pleonanthic, hermaphroditic or rarely unisexual
by abortion; leaf sheath disintegrating into a
brown, often clothlike mass of interwoven fibers;
adaxial hastula well developed (ephemeral in
]ohannesteijsmannia), abaxial hastula absent or
very small; stamens 6; pollen sulcate (rarely
trichotomosulcate, Pritchardia, Copernicia), exine
finely reticulate (rarely scabrate (Johannest-
eijsmannia, Pritchardia), tectate; carpels 3, wedge-
shaped, distinct in ovarian regions, connate
distally to form a slender style with apical dotlike,
or briefly 3-lobed stigma; ovules anatropous; stig-
matic remains apical; endosperm homogeneous,
but ruminate in Copernicia; a postament present
in all but Pritchardia, Colpothrinax, and
Copernicia.
A subtribe of 12 genera widely distributed
through the tropics, but with so many disjunctions
as to suggest an ancient origin. Differences
between the genera are small. Leaves are usually
palmate or costapalmate, but there are two dis-
tinctive leaf forms: the diamond-shaped leaf of
]ohannesteijsmannia, and wedge-shaped seg-
ments with reduplicate margins in most species of
Licuala. Washingtonia is unique in having more
or less woody, swordlike bracts and large chaffy
perianth segments.
15. Livistona R. Brown
Livistona R. Brown, Prod. Flor. Nov. Hoi!.: 267 (1810); Rodd,
Revision of Livistona (Arecaceae) in Australia (in press).
Slender to robust, solitary. Leaves palmate or
costapalmate; petiole unarmed or toothed; blade
of single (multiple)-fold segments. Inflorescences
branched to 5 orders; peduncular bracts 1-few.
Flowers solitary or in cincinni; calyx tubular 3-
lobed; corolla tubular with 3 valvate lobes; sta-
mens epipetalous, filaments connate in fleshy
ring. Fruit globose to ovoid, pyriform or ellipsoi-
dal. Seed ellipsoid. Germination remote-tubular;
eophyll lanceolate. About 25 spp., Africa and
Arabia, to the Himalayas and Rykyu Islands,
330
through Indochina and Malesia to New Guinea,
the Solomon Islands, and to its greatest diversity
in Australia.
16. Pholidocarpus Blume
Pholidocarpus Blume in J.A. & J.H. Schultes, Syst. Veg. 7: 1308
(1830); Beccari, Ann. Roy. Bot. Gard. Calcutta 13: 1-356
(1931); Dransf. & Uhl, Kew Bull. 38: 197-198 (1983).
Robust, solitary trees. Leaves costapalmate, peti-
ole with robust, bulbous-based spines; blade
deeply divided into 3-4-fold segments. Inflores-
cences branching to 4 orders, peduncular bracts l-
ea. 5. Flowers, solitary or in cincinni of 2-3; calyx
cup-shaped, 3-lobed; corolla deeply divided into 3
triangular, valvate petals; filaments slender,
united basally to form a free tube. Fruit large, glo-
bose; epicarp smooth (P. kingiana), or cracked
into corky brown warts. Germination remote-
tubular; eophyll entire. Six or fewer spp., south
Thailand, Malaya, Sumatra, Borneo, Sulawesi, and
the Moluccas.
17. Johannesteijsmannia H.E. Moore Fig. 81K
Johannesteijsmannia H.E. Moore, Principes 5: 116 (1961);
Dransf., Gard. Bull., Singapore 26: 63-83 (1972).
Solitary, acaulescent or short-trunked. Stem de-
cumbent or erect. Leaves diamond-shaped; petiole
toothed; blade subpinnately ribbed, lower mar-
gins toothed. Inflorescences short, curved; pedun-
cular bracts to 7, slightly inflated. Flowers solitary
or in cincinni of 2-4; calyx cup-shaped with 3
lobes; corolla divided almost to base into 3 valvate
lobes; stamens epipetalous; filaments broad,
fleshy, connate basally to form a ring; ovule anat-
ropous. Fruit rounded, mesocarp cracking to
produce thick, corky warts. Seed globose. Germi-
nation remote-tubular; eophyll simple. Four spp.,
one widespread but very local in south Thailand,
Malay Peninsula, Sumatra, and the western part
of Borneo, the other three endemic to the Malay
Peninsula.
18. Licuala Thunb.
Licuala Thunb., Kongliga Vetenskaps Acadamiens Nya
Handlingar 3: 286 (1782); Beccari, Ann. Roy. Bot. Gard.
Calcutta 13: 1-356 (1931); Furtado, Gard. Bull. Singapore
11: 31-73 (1940).
Solitary, clustered, acaulescent, shrubby, or rarely
treelike. Leaves palmate; petiole spiny or
unarmed; blade entire or of single to multiple-fold
Palmae
reduplicate segments, the central sometimes
stalked. Inflorescences spicate to branched to 3
orders; peduncular bracts 0-5 or more. Flowers
solitary or in cincinni of 2-3; calyx truncate,
irregularly splitting, or 3-lobed; corolla tubular
basally, 3-lobed; stamens epipetalous, filaments
distinct, or united into a tube or a 3-lobed
androecial ring, with 3 anthers with short fila-
ments, 3 sessile at sinuses. Epicarp smooth, rarely
corky-warted. Germination remote-tubular;
eophyll strap-shaped. About 108 spp., from India
and southern China through SE Asia to Malesia,
Queensland, the Solomon Islands and New
Hebrides, the greatest diversity being in Malaya,
Borneo, and New Guinea.
19. Pritchardiopsis Becc.
Pritchardiopsis Becc., Webbia 3: 131 (1910); Moore & Uhl,
Allertonia 3: 313-402 (1984).
Moderate, solitary, unarmed trees. Leaves briefly
costapalmate; blade divided to middle into single-
fold bifid segments. Inflorescences branched to
4 orders; pecuncle short, peduncular bracts
lacking. Flowers in cincinni of 3 or solitary
distally; calyx tubular, adnate basally, 3-lobed
distally; corolla tubular with 3 long valvate lobes;
stamen filaments briefly connate, adnate to petal
tube. Fruit large, globose, endocarp woody,
keeled. Germination remote-tubular; eophyll
entire. Only one sp., P. jeanneneyi Becc., New
Caledonia.
20. Pritchardia Seemann & H. Wendl.
Pritchardia Seemann & H. Wend!. ex. H. Wend!., Bonplandia
10: 197 (1862), nom. cons.; Becc. & Rock, Mem. Bernice P.
Bishop Mus. 8(1): 1-77 (1921); Hodel, Principes 24(22): 25-
81 (1980).
Moderate, solitary, unarmed trees. Leaves
costapalmate; blade divided 1/3 to 1/2 into single-
fold bifid segments; inflorescences solitary or 2-4
with common prophyll in each axil, branched to 3
orders; peduncular bracts several; rachillae
crowded at end oflong peduncle. Flowers solitary;
calyx tubular, 3-lobed; corolla tubular basally with
3 valvate lobes forming a deciduous cap; filament
bases connate, tubular. Germination remote-
tubular; eophyll entire. About 38 spp., Fiji, Tonga,
Danger Islands, and Hawaii. All but five spp. are
Hawaiian endemics; many are extremely rare and
endangered, or not seen in the wild for several
years.
 Palmae
21. Colpothrinax Griseb. & H. Wendl. Fig. SOD
Colpothrinax Griseb. & H. Wend!., Bot. Zeit. 37: 148 (1879);
L.H. Bailey, Gentes Herb. 4(10): 357-360 (1940); Read,
Principes 13(1): 13-22 (1969).
Moderate, solitary, unarmed trees. Stem columnar
or ventricose. Leaves costapalmate; sheath disin-
tegrating into fibrous network or long fibers; blade
orbicular, irregularly divided into single-fold bifid
segments. Inflorescences branched to 4 orders;
peduncular bracts 4-9; peduncle and rachis long,
equal. Flowers on tubercles; calyx cuplike, 3-
lobed; corolla longer, shortly tubular basally, free
lobes 3, valvate; filaments connate in epipetalous
cup, adnate to and ± equalling corolla tube.
Germination not recorded. Eophyll probably
simple. Two spp., Cuba, Guatemala, and along the
Atlantic slopes of the mountains in Panama.
22. Acoelorraphe H. Wendl. Fig. SOH
Acoelorraphe H. Wend!., Bot. Zeit. 37: 148 (1879); L.H. Bailey,
Gentes Herb. 4(10): 361-365 (1940).
Moderate, clustered. Stems with brown fibrous
sheaths and petiole bases. Leaves briefly
costapalmate; petiole spiny; blade nearly orbicu-
lar, divided to below the middle into deeply bifid
single-fold segments, abaxially silvery. Inflores-
cences exceeding the leaves, branched to 4 orders;
peduncular bracts 2. Flowers on low spurs in
cinicinni of 2-3; sepals 3, slightly connate basally;
petals 3, connate for 1/4length; filaments connate
in shallow cup. Fruit small, black. Germination
not recorded, eophyll simple. Only one sp., A.
wrightii (Griseb. & H. Wendl.) H. Wendl., south-
ern Florida, the West Indies, and parts of the
Caribbean coast of central America.
23. Serenoa Hook. f. Fig. SOK
Serenoa Hook. f. in Bentham & Hook., General Pl. 3: 879, 926
(1228); L.H. Bailey, 3(6): 234-339 (1934).
Clustered, shrubby. Stem subterranean or pros-
trate, rarely erect, leaf sheaths persistent; axillary
buds developing as inflorescences or vegetative
shoots. Leaves palmate; petiole margin with small
teeth; blade nearly orbicular, regularly divided 1/2
into stiff bifid, single-fold segments. Inflores-
cences curved, branched to 3-4 orders; peduncu-
lar bract 1 or lacking. Flowers solitary or in pairs;
calyx tubular, 3-lobed; corolla tubular, split 2/3
length into 3 valvate lobes; filaments adnate to
corolla tube. Fruit dark blue to black. Germination
331
remote-ligular; eophyll entire. Only one sp., S.
repens (Bartram) Small, southeastern United
States.
24. Brahea Mart. ex Endl.
Brahea Mart. ex End!., Genera Pl.: 252 (1837); L.H. Bailey,
Gentes Herb. 4(3): 119-125 (1937); Henderson eta!., Field
Guide: 54-56 (1995).
Moderate, rarely clustered. Sheath becoming
fibrous, eventually splitting basally; petiole
toothed or unarmed; blade divided nearly 1/2 into
single-fold, deeply bifid, stiff or flexible segments.
Inflorescences branched to 4 orders; peduncular
bracts 0-several. Flowers solitary or in cincinni of
2-3; sepals 3, distinct, imbricate; petals 3, tubular
basally, valvate apically; stamens epipetalous, fila-
ments connate in 6-lobed ring. Germination
remote-ligular; eophyll entire. About 16 spp., Baja
California, Guadalupe Island, Mexico, and
Guatemala.
25. Copernicia Mart. ex Endl. Fig. S1J
Copernicia Mart. ex End!., Genera Pl.: 253 (1837); Dahlgren &
Glassman, Gentes Herb. 9(1): 3-40 (1961), 9(2): 43-232
(1963); Henderson eta!., Field Guide: 57-61 (1995).
Solitary trees. Leaves palmate to shortly costapal-
mate, petiole lacking, short, or elongate, margins
toothed; adaxial hastula short to long, unarmed or
spiny; blade wedge-shaped to orbicular divided
1/4-1/3 into single-fold segments, outermost bifid.
Inflorescences branched to 6 orders; peduncular
bracts 0-1; subsequent bracts tubular, often
densely tomentose. Flowers solitary or in cincinni
of 2-4; sepals 3 connate; corolla tubular, 3-lobed,
thick-tipped, valvate; filament bases united in en-
pule. Endosperm ruminate. Germination remote-
tubular; eophyll entire. Twenty-five or fewer spp.,
three in S America, two in Hispaniola, the remain-
der in Cuba.
26. Washingtonia H. Wendl. Fig. SOC
Washingtonia H. Wend!., Bot. Zeit. 37: !xi, 68, 148 (1879),
nom. cons.; L.H. Bailey, 4(2): 52-82 (1936).
Robust, tall, solitary trees. Leaves costapalmate;
sheath split below petiole; margins fibrous; petiole
toothed; blade divided 1/3 into bifid, single-fold
segments, pendulous, interfold filaments con-
spicuous. Inflorescences branched to 3-4 orders;
rachis bracts splitting becoming swordlike, coria-
ceous. Flowers solitary; calyx chaffy, tubular with
332 Palmae

3 tattered lobes; corolla tubular for 1/4 of its recognized spp., but probably fewer, ranging from
length, lobes valvate, pointed; stamens adnate southern India and Sri Lanka to the Bay of Bengal,
to corolla tube. Germination remote-ligular; and Indochina through Malesia to northern
eophyll entire. Two closely related spp., W. filifera Australia; distribution probably much influenced
(Linden) H. Wendl., southeastern California, hyman.
western Arizona, and Baja California, W. robusta
H. Wendl., Baja California and Sonora, Mexico.
28. Nannorrhops H. Wendl. Fig. 83H,I
Nannorrhops H. Wend!., Bot. Zeit. 37: 148 (1879); Moore,
lc. Subtribe Coryphinae J. Dransf. & N.W. Uhl Flora Iranica 146 (1980).
(1986).
Shrubby, clustered, hapaxanthic, hermaphroditic.
Leaves induplicate; pollen elliptic, sulcate (rarely Branching in erect stems dichotomous. Leaves
trichotomosulcate in Corypha), exine finely or costapalmate; sheath splitting; hastulae absent;
coarsely (Kerriodoxa) reticulate, tectate; carpels 3, blade divided into single-fold, bifid segments.
united basally, styluli free or united into a style but Inflorescences above the leaves, branched to 4
stylar canals separate. orders, branches subtended by leaves with
A heterogeneous subtribe of four Old World reduced blades or tubular bracts; peduncular
genera. Species of Corypha are most striking bracts 0 to several. Flowers in cincinnus of
palms because of their massiveness. The inflores- 1-3(-7), calyx tubular, 3-lobed; corolla stalklike
cence is the largest among seed plants; the number basally, 3-lobed distally, stamens 6, anthers
of flowers has been estimated as ten million latrorse; carpels 3, connate except at base, ovule
(Tomlinson and Soderholm 1975). Hapaxanthic anatropous. Stigmatic remains basal; endosperm
shoots and rather similar flowers distinguish homogeneous. Eophyll undivided. Only one, vari-
Nannorrhops and Corypha, and Chuniophoenix able species (N. ritchiana (Griff.) Aitch.) in semi-
and Nannorrhops lack a hastula and have tubular desert areas of the Middle East (Iran, Afghanistan,
rachilla bracts. Otherwise, the genera are rather Pakistan, and Arabia).
disjunct morphologically. However, two investi-
gations using cladistic analyses of morpho-
logical and molecular characters (Uhl et al. 29. Chuniophoenix Burr.
1995, Barrow 1996) have resolved Nannorrhops, Chuniophoenix Burr., Notizb. Bot. Gart. Berlin-Dahlem 13:
Chuniophoenix, and Kerriodoxa in a monophyl- 583 and 582-606 (1937); Tang & Wu, Acta Phytotax. Sin.
etic group with Corypha usually in a sister posi- 15: 111-112 (1977).
tion. The subtribe is further resolved as related to
the Borasseae. Small, clustered, unarmed, pleonanthic, hermaph-
roditic or polygamodioecious. Leaves palmate,
sheath splitting, in C. hainanensis petiole split ba-
27. Corypha L. Fig. SOB sally, hastulae absent; blade deeply divided into
Corypha L., Sp. Pl.: 1187 (1753); Becc., Ann. Roy. Bot. Gard.
single to several-fold segments. Inflorescences in-
Calcutta 13: 1-356 (1931). terfoliar, spicate or branched to 2 orders, peduncu-
lar bracts several, rachilla bracts tubular. Flowers
Solitary, hapaxanthic, hermaphrodite, trees. solitary or in cincinnus of 1-7, stamens 6; anthers
Leaves costapalmate, sheath cleft below the peti- introrse; gynoecium tricarpellate, ovary with sep-
ole, petiole massive, margins toothed; blade tal nectaries basally, ovules anatropous; ovary or
divided to ca. 1/2 into single-fold, bifid segments. pollen aborting in polygamodioecious individuals.
Inflorescences above the leaves, subtended by Stigmatic remains apical. Endosperm ruminate or
leaves with reduced blades or by tubular bracts, homogeneous; embryo basal. Eophyll entire. Two
branched to the 4th order. Flowers in adnate spp., Vietnam, S China, and Hainan Island.
cincinni of up to 10; calyx tubular basally, 3-lobed;
petals basally imbricate, stamens 6, antesepalous
30. Kerriodoxa J. Dransf.
free, antepetalous adnate basally to petals; gyno-
ecium tricarpellate, syncarpous, triovulate, style Kerriodoxa J. Dransf., Principes 27: 4 (1983).
elongate, ovule hemianatropous. Fruit with basal
stigmatic remains; endosperm homogeneous. Solitary, acaulescent or erect, unarmed, pleo-
Germination remote-tubular; eophyll entire. Eight nanthic, dioecious. Leaves palmate, sheath split;
 Palmae
adaxial hastula conspicuous, abaxial absent; blade
divided 1/4-1/3 into single-fold segments, dis-
colorous. Inflorescences interfoliar; staminate
branched to 4 orders, pistillate to 2. Flowers in
pairs on low spurs, similar; calyx with tomentose
tubular base, 3 glabrous lobes; corolla stalklike
basally, 3-lobed, papillose; stamens 6, anthers
latrorse; carpels 3 (rarely 4), connate, tips distinct;
ovule anatropous. Stigmatic remains basal. En-
dosperm ruminate; embryo subbasal; germination
remote-ligular; eophyll apically lobed. Only one
sp., K. elegans J. Dransf., known from two locali-
ties in peninsular Thailand.
ld. Subtribe Sabalinae Mart. (1837).
Solitary, hermaphroditic, pleonanthic, unarmed;
leaves induplicate, costapalmate, carpels 3, united
throughout, with a common stylar canal.
The occurrence of prophylls on inflorescence
branches is useful in delimiting species. Saba[ is
ditinguished from other genera with costapalmate
leaves by the lack of spines on the petiole and by
the split leaf bases. Formerly used for making
brooms and locally as a source of thatch. Many
species are important ornamentals.
31. Sabal Adanson Fig. 801, 83J-R
Sabal Adanson, Families des Plantes 2: 495, 599 (1763); Zona,
Aliso 12: 583-666 (1990).
Acaulescent or erect. Sheath split, adaxial hastula
short, abaxial hastula obscure; blade divided ca.
1/2 into single-fold bifid segments. Inflorescence
interfoliar, branching to 4 orders; peduncular
bracts several. Flowers solitary; calyx and corolla
tubular, 3-lobed; stamens 6, filaments in tube ad-
nate to corolla basally, anthers latrorse; pollen el-
liptic, sulcate, exine finely reticulate, tectate;
carpels 3, completely connate, style elongate,
stigma capitate, ovule anatropous. Stigmatic scar
basal; endosperm homogeneous, postament shal-
low; embryo lateral. Germination remote-ligular;
eophyll entire. Sixteen spp. One of the larger
coryphoid genera confined to the central Western
hemisphere from Colombia to northeastern
Mexico and the southeastern United States.
2. Tribe Phoeniceae Drude (1881).
Dioecious, leaves pinnate, induplicate, leaflets
with acute tips, proximal leaflets modified as
333
spines; inflorescence bearing only a prophyll,
branching to 1 order; flowers solitary, dimorphic,
endocarp membranous, seed with a deep longitu-
dinal furrow.
The single genus, Phoenix, is easily distin-
guished from all other palms by the induplicate
pinnate leaf with the lower leaflets modified as
spines. The distinct carpels and relatively un-
specialized perianth and stamens are paralleled
in the apocarpous coryphoid genera of the
Corypheae. Cladistic analyses (Uhl et al. 1995; Bar-
row 1996) do indeed resolve Phoenix as close to
the Thrinacinae and Sabalinae. A modern devel-
opmental study of the unique leaf is greatly
needed. The fossil record of the genus is extensive
(Harley 1996).
Most of the 12 species are plants of semiarid
regions but grow near water courses, oases, or un-
derground water sources; a few species are found in
tropical monsoonal areas. P. paludosa Roxb. occurs
in the Asian perhumid regions, where it is confined
to the landward fringe of mangrove forest.
The genus is immensely important from an
economic point of view. Not only does it include
the date palm, the major crop of several Middle
Eastern countries and of lesser importance else-
where, but other species are widely used as a
source of fiber for weaving, starch, sugar, and a
multiplicity of things such as thatch and fuel.
Many species are widely grown as ornamentals.
Phoenix roebelenii O'Brien is commercially im-
portant as a pot plant. Species are known to hy-
bridize freely.
32. Phoenix L. Fig. 810,P, 84A-R
Phoenix L., Sp. Pl.: 1188 (1753); Barrow, Kew Bull. (in
press).
Solitary or clustered, pleonanthic, dioecious.
Leaves induplicate, pinnate; sheath fibrous;
leaflets single-fold, acute, regularly arranged or
grouped, proximal modified as spines (acantho-
phylls). Inflorescences interfoliar, branching to 1
order, the staminate and pistillate similar; pedun-
cular bracts absent. Flowers solitary, staminate
sepals connate, petals 3, valvate, stamens usually
6 (rarely 3 or 9), pollen elliptic or circular, sulcate,
exine reticulate, tectate; pistillate flowers globose;
sepals connate; petals imbricate, staminodes 6,
carpels 3, distinct, follicular, ovule anatropous.
Endosperm homogeneous; embryo lateral or
subbasal. Germination remote-tubular; eophyll
undivided. Twelve spp., Atlantic islands through
Africa, Crete, the Middle East, and India to
334 Palmae
Fig. 84A-Q. Palmae-Coryphoideae. A-N Phoenix dactylifera.
A Habit showing basal suckers and infructescences. B Stami-
nate inflorescence bearing 2-keeled prophyll and branching to
1 order only. C Staminate rachilla bearing solitary flowers. D
Pistillate rachilla bearing solitary flowers. E Staminate flower
bud. F Staminate flower in section, showing pistillode. G Pis-
tillate flower. H Apocarpous gynoecium. I Same in cross-
section at level of ovules. J Fruit. K Fruit in vertical section.
L Seed in ventral and M dorsal view. N Seed in cross-section at
level of embryo. 0 P. reclinata, crown showing leaves with
basal acanthophylls and interfoliar inflorescences. P P.
canariensis, detail of stem with spirally arranged leaf scars.
Q Phoenix sylvestris, stem with scars from incisions on the leaf
sheaths for drawing sugary sap. (Takhtajan 1982)
Hong Kong, Taiwan, Philippines, Sumatra, and
Malaya.
3. Tribe Borasseae Mart. (1837).
Dioecious; leaves palmate or costapalmate, in-
duplicate; staminate and sometimes pistillate
flowers borne in deep pits formed by connation
and adnation of rachilla bracts; staminate flowers
rarely solitary, usually in cincinni; stamens ex-
serted from pits by elongation of receptacle;
pistillate flowers solitary, bibracteolate; fruit 1-3-
seeded with thick hard endocarp, usually with an
apical pore.
The eight genera of Borasseae are confined to
the Old World. The tribe is divisible into two ap-
parently natural subtribes. The constant dioecy,
the staminate flowers in pits in catkinlike
rachillae, and fruits with well-developed endocarp
are characteristic features. Cladistic analyses us-
ing morphological and molecular datasets have
resolved this tribe as monophyletic (Uhl et al.
1995; Barrow 1996).
KEY TO THE SUBTRIBES AND GENERA OF
THE BoRASSEAE
1. Staminate and pistillate inflorescences dissimilar; stami-
nate flower clusters concealed in pits; bracteoles subtending
staminate flowers not bearing tufts of hairs; pistillate
flowers sessile in the axils of large leathery bracts; fruit
sessile, symmetrical, 1-3-seeded, endocarp forming
pyrenes, stigmatic remains apical (subtribe Lataniinae) 2
- Staminate and pistillate inflorescences similar; flowers of
both sexes sunken in pits; bracteoles subtending flowers
bearing tufts of hairs; fruit stalked, usually 1-seeded, if 2-3-
seeded, lobed, pyrenes not formed; stigmatic and carpellary
remain basal (subtribe Hyphaeninae) 5
2. Staminate flowers solitary or in cincinni of 2-6 flowers 3
- Staminate flowers numerous, in cincinni of 30-60 4
3. Bracts of staminate inflorescence imbricate at the margins;
pistillode nipplelike; pyrenes densely hairy with thick, in-
ternally ridged walls; seed grooved to conform to the ridges.
Malay Peninsula, Borneo 33. Borassodendron
- Bracts of the staminate inflorescence connate at the mar-
gins; pistil!ode columnar, prominent; pyrenes not hairy, but
thin, ridged, sculptured, or apically 3-lobed; seed not
grooved. Mascarene Islands 34. Latania
4. Staminate rachillae several, wide; stamens 6; pyrenes not
deeply bilobed; seeds usually 3, wider than thick, shallowly
to deeply bilobed. Africa, Madagascar, Arabia, India to New
Guinea and Australia 35. Borassus
- Staminate rachillae only 1 or 2, very wide; stamens 15-18;
pyrenes deeply bilobed, very thick; seed usually 1(-3), very
large, deeply bilobed. Seychelles Islands 36. Lodoicea
5. Endocarp conspicuously winged and crested, with a con-
spicuous vertical crest, the endocarp lacking an apical
pore and the 2 halves of the endocarp partially separating,
allowing the cotyledonary sheath to emerge; endosperm
ruminate. Madagascar. 40. Satranala
 Palmae
- Endocarp only faintly ridged if at all, with an apical pore;
endosperm homogeneous or ruminate
6. Seed deeply grooved. Madagascar 39. Bismarckia
- Seed not grooved
7. Fruit various in shape, distally expanded, often shouldered,
usually asymmetrical, rarely ovoid or spherical; seed irregu-
lar with endosperm homogeneous. Africa, Madagascar,
Arabia, India 37. Hyphaene
- Fruit ovoid; seed with endosperm ruminate. Egypt and
Sudan 38. Medemia
3a. Subtribe Lataniinae Meisner {1842).
Solitary; staminate and pistillate rachillae dissimi-
lar; pistillate flowers sessile, not borne in well-
defined pits; pits in staminate rachillae lacking
hairs; fruit unlobed, with 1-3 pyrenes and seeds;
stigmatic remains apical.
33. Borassodendron Becc.
Borassodendron Becc., Webbia 4: 359 (1914); Dransfield,
Reinwardtia 8: 351-363 (1972).
Unarmed. Leaves costapalmate; adaxial hastula
large, abaxial absent; blade with multifold seg-
ments, divided to 1/4-2/3 into single-fold bifid
segments. Staminate inflorescence branched to 2
orders; rachilla pits, each containing a cincinnus
of 2-6 flowers; stamens 6-15; pollen circular,
monoporate or sulcate, exine coarsely reticulate,
tectate. Pistillate inflorescence with 0-4 branches;
staminodes with empty anthers. Fruit 3-seeded;
endocarp internally ridged. Seed grooved; en-
dosperm homogeneous. Eophyll palmate. Two
spp., B. machadonis (Ridl.) Becc. in southern
Thailand and northern Malaya, B. borneense J.
Dransf. in Borneo.
34. Latania Comm. ex Juss.
Latania Comm. ex Juss., Genera Pl.: 39 (1789); Moore &
Gueho, Gentes Herb. 12(1): 1-16 (1984).
Mostly unarmed. Leaves costapalmate; adaxial
hastula short, abaxial absent; blade divided to 1/3-
1/2 into single-fold bifid segments. Staminate
inflorescence with 1-14 rachillae; flowers solitary
in pits; stamens 15-30 or more; pollen ::±:: elliptic,
sulcate, with foveolate-fossulate, tectate exine;
pistillate inflorescence with fewer branches;
staminodes 6-9. Fruit 3-(1-2) seeded. Pyrenes
sculptured. Seed smooth; endosperm homoge-
neous. Eophyll digitate. Three spp., Mascarene
Islands.
335
35. Borassus L.
6
Borassus L., Sp. Pl.: 1187 (1753); Becc., Palmae della Tribu
7 Borasseae (ed. U. Martelli) (1924).
Armed. Leaves costapalmate; adaxial hastula
large, abaxial low; blade suborbicular to flabellate,
divided to 1/2 into single-fold bifid segments.
Staminate inflorescence branched to 2 orders; pits
with reflexed cincinni of ca. 30 flowers; stamens
6; pollen elliptic, sulcate, with verrucate, tectate
exine, finely reticulate between the warts. Pistillate
inflorescence with 0-1 branch; ovule atropous.
Fruit 1-3-seeded; endocarp internally ridged. Seed
bilobed; endosperm homogeneous. Eophyll
undivided. Seven spp. recognized, but probably
only three or four to be maintained, Africa,
Madagascar, northeastern Arabia, through India
and SE Asia to New Guinea and Australia.
36. Lodoicea Comm. Fig. 85A-O
Lodoicea Comm. ex DC., Bull. Sci. Soc. Philomatique 2(46):
171 (1800); Bailey, Gentes Herb. 6(1): 3-48 (1942).
Unarmed. Leaves costapalmate, juvenile
diamond-shaped; hastulae absent; blade divided
to 1/4-1/3 into single-fold bifid segments. Stami-
nate inflorescence unbranched or 2-3 rachillae;
60-70 flowers in recurved cincinni in pits; stamens
17-22; pollen sulcate, with foveolate, tectate exine;
pistillate inflorescence unbranched; staminodial
tips to 11. Fruit very large 1( -3) seeded. Seed larg-
est known, bilobed; endosperm homogeneous.
Eophyll lobed. Only one species (L. maldivica
(Gmelin) Pers.), Seychelles Islands.
3b. Subtribe Hyphaeninae Becc. {1924).
Solitary or sometimes clustered in Hyphaene.
Staminate and pistillate rachillae similar; pistillate
flowers pedicellate; pits of both sexes filled with
hairs; fruit 1-seeded or 2-3-seeded and then 2-3-
lobed; stigmatic remains basal or central between
the lobes.
The four genera of this subtribe are closely
related. Bismarckia is endemic to Madagascar,
Hyphaene is widely distributed in Africa,
Madagascar, and Arabia with one or possibly two
species just reaching the Indian subcontinent;
Medemia has been rediscovered in the Sudan, and
Satranala was recently found in Madagascar. The
genera differ largely in details of the fruits.
Pyrenes are not formed; epicarp, mesocarp, and
endocarp develop more or less separately, result-
ing in a lobed fruit if more than one carpel rna-
336
~D
Fig. 85A-O. Palmae-Coryphoideae. A-0 Lodoicea maldivica.
A Habit. B Detail of crown of staminate plant showing split
leaf bases and staminate inflorescence with peduncular bract.
C Surface of staminate rachilla showing bracts and staminate
flowers emerging at anthesis. D Vertical section of cincinnus
of staminate flowers. E Staminate flower at an thesis. F Stamen
in dorsal view. G Staminate flower in section showing the
elongate receptacle and united filaments. H Detail of crown of
pistillate tree. I Tip of pistillate rachilla. J Pistillate flower with
imbricate sepals and petals and large pistil. K Fruit. L Pyrene.
M Pyrene and seed in longitudinal section, showing hollow
endosperm. N Germinating seed with emerging cotyledonary
sheath. 0 Seedling. (Takhtajan 1982)
Palmae
tures. In Bismarckia, the endocarp is irregularly
ridged and pitted within and the endosperm is
homogeneous; in Medemia the endocarp is not
ridged but the endosperm is ruminate, while in
Hyphaene the fruits have unridged endocarps and
homogeneous endosperm.
37. Hyphaene J. Gaertn. Fig. 80E
Hyphaene J. Gaertn., De Fructibus 1: 28 (1788}; Becc., Palme
della Tribu Borasseae (1924}.
Stem dichotomously branched, rarely un-
branched. Leaves costapalmate; petioles spiny,
 Palmae
adaxial hastula large, abaxial absent; blade divided
to 1/3 into single-fold bifid segments. Staminate
inflorescences with 1-6 rachillae, pistillate with
1-3; staminate flowers 3 in each pit; pistillate
solitary; stamens 6; pollen ± elliptic, sulcate, with
verrucate, tectate exine, finely reticulate between
the warts; staminodes 6-9. Fruit variable, borne on
enlarged pedicel. Seed smooth; endosperm homo-
geneous. Eophyll simple. Probably only about ten
spp., drier parts of Africa southwards to Natal,
Madagascar, Red Sea and Gulf of Eilat coasts,
coastal Arabia, and west coast of India.
38. Medemia Wi.irttemb. ex H. Wendl.
Medemia Wiirttemb. ex H. Wend!., Bot. Zeit. 39: 90,93 (1881};
Gibbons & Spanner, Principes 40: 65-74 (1996}.
Sparsely armed. Leaves costa palmate; hastulae ab-
sent; blade divided to 2/3 into single-fold bifid
segments. Staminate inflorescences with 1-7
rachillae, pistillate with 1; staminate flowers 3 in
each pit; pistillate solitary; stamens 6; pollen ellip-
tic, sulcate, with areolate, tectate exine. Fruit
ovoid or 2-lobed, pedicellate. Seed smooth; en-
dosperm ruminate. Eophyll simple. One or two
spp., Egyptian Nubia and northeastern Sudan.
39. Bismarckia Hildebrandt & H. Wendl.
Bismarckia Hildebrandt & H. Wend!., Bot. Zeit. 39: 90, 93
(1881}; Becc., Palme della Tribu Borasseae (1924).
Unarmed. Leaves costapalmate; adaxial hastula
large; abaxial absent; blade divided to 1/4-1/3 into
single-fold bifid segments. Inflorescences with 3-
7-1 rachillae, pistillate fewer; staminate flowers 3
in each pit; pistillate solitary; stamens 6; pollen
elliptic, sulcate, with areolate, tectate exine; ovule
atropous. Fruit ovoid or 2-lobed, pedicellate,
endo-carp flanged, endosperm homogeneous
grooved to match. Eophyll simple. Only one sp., B.
nobilis Hildebrandt & H. Wendl., drier parts of
Madagascar.
40. Satranala J. Dransf. & Beentje
Satranala J. Dransf. & Beentje, Kew Bull. 50: 67 (1995} and 59:
85-92 (1995}.
Leaves costapalmate; petiole toothed distally;
adaxial hastula present; abaxial absent; blade
divided to 1/4-1/3 into single-fold bifid segments.
Staminate inflorescence with partial inflores-
cences, bearing 3-9 rachillae; staminate flowers 3
337
in each pit; pollen ellipsoid, sulcate, with coarsely
rugulate-fossulate, tectate exine. Fruit globose,
endocarp woody with broad anastomosing
flanges, endosperm ruminate. Eophyll palmate.
Only one sp., S. decussilvae Beentje & J. Dransf.,
east coast rainforest of Madagascar.
II. Subfamily Calamoideae Griff. (1844).
Frequently fiercely armed; leaves pinnate, pin-
nately ribbed or palmate, reduplicate; inflores-
cence frequently displaying adnation, bracts of all
orders almost always tubular; flowers almost al-
ways borne in dyads or dyad derivatives; ovary
and fruit covered in reflexed scales, usually ar-
ranged in neat vertical rows; ovary incompletely
trilocular, ovules basal, anatropous, with the
micropyles facing center of the gynoecium;
stigmatic remains apical, pericarp usually thin
at maturity, endocarp not differentiated (except
in Eugeissona); seeds 1-3, usually with thick
sarcotesta.
The subfamily includes the largest palm genus,
Calamus. Four genera (Raphia, Mauritia,
Mauritiella, and Lepidocaryum) occur in the New
World, while the rest (including Raphia) occur in
the Old World. Members of the subfamily are im-
mediately distinguishable by the presence of re-
flexed scales on the pericarp. The scales are usually
arranged in neat rows that come to mimic
orthostichies and parastachies of a phyllotactic
spiral, thereby misleading some workers in the
past to regard them as being foliar organs. How-
ever, they are clearly epidermal emergences and
are produced basipetally in the developing ovary.
The tricarpellate, triovulate syncarpous gyno-
ecium has incomplete partitions and the anatro-
pous ovules are basally attached. Spines are
abundant but are restricted to leaves or other
foliar organs, apart from root spines, that are
present in Eugeissona and Mauritiella. There are
many climbing members of the group, and several
different features are associated with the climbing
habit. In particular, barbed whiplike structures
have evolved, providing means of support for the
flexible stems. These may be extensions of the leaf
rachis (cirri), armed with reflexed epidermal emer-
gences and/or acanthophylls (modified leaflets) or
sterile inflorescences (flagella). There is extraordi-
nary diversity in pollen structure. All recent phylo-
genetic analyses have confirmed the monophyletic
nature of the subfamily. However, the arrange-
ment of genera presented by Uhl and Dransfield
( 1987), that forms the basis of the present account,
338 Palmae
will almost certainly require modification (W.J.
Baker and J. Dransfield, in prep).
KEY TO THE GENERA OF THE CALAMOIDEAE
1. Leaves pinnate or, rarely, simply forked but with pinnate
venation (Calameae)
- Leaves palmate with usually numerous reduplicate seg-
ments (Lepidocaryeae) 21
2. Flowers hermaphroditic, borne singly or in dyads or triads
(climbing palms)
- Polygamous, monoecious, or dioecious palms (climbing,
acaulescent or treelike palms)
3. Pleonanthic rattan; rachilla bracts minute, incomplete;
floral bracteoles absent; corolla tubular, very thick, with 3
short lobes; filaments united in a fleshy epipetalous ring.
Africa 42. Eremospatha
- Hapaxanthic rattans; rachilla bracts conspicuous, com-
plete or incomplete; floral bracteoles present (but in
Korthalsia concealed by hairs); corolla coriaceous, tubular
at the base, with 3 boat -shaped lobes; filaments not united
in a fleshy ring
4. Flowers in dyads, rarely in triads; rachilla bracts disti-
chous, not filled with hairs; leaflets linear-lanceolate,
entire. Africa 41. Laccosperma
- Flowers solitary; rachilla bracts spiral, forming a tight
catkin, usually filled with hairs; leaflets lanceolate to
broad, diamond-shaped, praemorse. SE Asia and Malesia
45. Korthalsia
5. Polygamous palms; flowers borne in dyads of 1 hermaph-
roditic and 1 staminate
- Monoecious or dioecious palms
6. Flowers very large, borne in a tight cupule of 7-11 overlap-
ping bracts; petals woody, sharply pointed; stamens 20-70.
Fruit ovoid, beaked, covered with minute scales; mesocarp
fibrous; endocarp thick, woody, with 6 or 12 internal lon-
gitudinal flanges penetrating the seed. Malay Peninsula
and Borneo 43. Eugeissona
- Flowers small, each pair borne in the axil of a rachilla bract
filled with hairs; petals neither woody nor spine-tipped;
stamens 6. Fruit rounded, with large scales, not beaked;
mesocarp thick, sparsely fibrous; endocarp lacking.
Papuasia 44. Metroxylon
7. Monoecious palms
- Dioecious palms
8. Climbing; flowers borne in clusters of ca. 5-11 in the axils
of overlapping distichous bracts, the central 1 or 3 pistil-
late, the rest staminate in 2 lateral cincinni. Fruit small,
covered with small scales. Africa 59. Oncocalamus
- Massive acaulescent or tree palms; flowers solitary, each in
the axil of a rachilla bract, the proximal on the rachillae
pistillate accompanied by 2 bracteoles, the distal staminate
accompanied by 1 bracteole, very rarely odd dyads present
at the transition from staminate to pistillate. Fruit very
large with few large scales. Africa, Madagascar, Central
America, and northern S America 58. Raphia
9. Pleonanthic, acaulescent; inflorescences short, in bud
enclosed within the subtending leaf sheath, at anthesis
emerging from a vertical groove on the abaxial surface of
the sheath, terminal leaflets compound, or the whole leaf
flabellate; rachillae of both sexes catkinlike. SE Asia to W
Malesia 47. Salacca
- Acaulescent, tree or climbing; inflorescences not emerging
from a vertical groove on abaxial surface of sheath; termi-
nalleaflets usually single-fold; rachillae catkinlike or not
10
10. Pistillate rachillae bearing dyads of one pistillate flower
together with a sterile staminate flower (very rarely 2 pis-
tillate and 1 sterile staminate); staminate rachillae bearing
fertile staminate flowers, singly or in pairs 11
- Pistillate rachillae bearing solitary flowers; staminate
flowers solitary or paired 17
11. Acaulescent, hapaxanthic palm; rachillae catkinlike in
2 both sexes; staminate flowers borne in pairs; seed with
conspicuous wide chalaza! pit and adherent sarcotesta.
Thailand, Peninsular Malaysia, Sumatra, and Borneo
46. Eleiodoxa
3 - Acaulescent or climbing, hapaxanthic or pleonanthic
palms. Rachillae very rarely catkinlike, if so, then not in
5 both sexes; staminate flowers borne singly 12
12. Entire inflorescence enclosed within the prophyll 13
- Inflorescence not enclosed within the prophyll, or
prophyll sheathing only the base of the inflorescence 15
13. Prophyll opening by a pair of pores near the apex, splitting
along its length only after anthesis or accidentally. Malay
Peninsula, Sumatra, Java, Borneo 52. Ceratolobus
- Prophyll splitting along its length at anthesis 14
14. Prophyll much larger than any other inflorescence bract;
4 two erect earlike appendages (auricles) present at the
mouth of the leaf sheath; climbing whips absent; en-
dosperm homogeneous. Malay Peninsula, Borneo
51. Pogonotium
- Prophyll enclosing a series of similar rachis bracts, but
with their tips enfolded in the beak of the prophyll;
auricles absent; cirrus almost always present; endosperm
ruminate. India to New Guinea 48. Daemonorops
(subgenus Daemonorops)
6 15. Prophyll and rachis bracts splitting along their lengths to
7 the very base, usually neatly falling at an thesis except for
the prophyll which may persist long after anthesis; floral
bracteoles usually low, often rather inconspicuous; en-
dosperm always ruminate. India to New Guinea
48. Daemonorops (subgenus Piptospatha)
- Prophyll and rachis bracts strictly tubular, or, if splitting,
then persistent and with a tubular base; floral bracteoles
inconspicuous to conspicuous; endosperm ruminate or
homogeneous 16
16. Short, stout rattan palm without climbing whips; inflores-
cence very short with prophyll in normal position, rachis
bracts tubular only at the base, expanded, elongate, and
8 abruptly narrowed to a spiny beak, inserted distichously at
9 right angles to the prophyll; fruit usually 3-seeded; en-
dosperm homogeneous. Malay Peninsula 50. Calospatha
- Acaulescent to high-climbing rattans or thicket-forming
shrubs, cirri or flagella usually present; inflorescence short
to very long with strictly tubular bracts, or if bracts ex-
panded, then not abruptly beaked; fruit usually 1-seeded,
very rarely 3-seeded; endosperm homogeneous or rumi-
nate. Africa to the Pacific 49. Calamus
17. Very tall, pleonanthic, tree palm; leaf sheaths bearing
combs of slender, soft bristlelike spines; rachillae numer-
ous, very slender, well exserted from conspicuous sub-
tending bracts, densely tomentose, rachilla bracts and
bracteoles mostly obscured by hairs; fruit very small,
scarcely exceeding 1 em. Sulawesi to New Guinea
57. Pigafetta
- Pleonanthic or hapaxanthic, rattan palms, usually fiercely
armed; rachillae if slender, then without tomentum, or
enclosed by subtending bracts; fruit at maturity usually at
least 1.5 em wide 18
18. Pleonanthic palm, briefly climbing but lacking climbing
whips; inflorescence bracts distichous, fibrous, netlike,
obscuring the rachillae. Borneo 53. Retispatha
 Palmae 339

- Hapaxanthic palms, usually high-climbing, cirri present;
inflorescence bracts not netlike 19
19. First-order inflorescence branches pendulous, bearing
strictly distichous, conspicuous, overlapping or not, boat-
shaped bracts, often completely enclosing the rachillae;
rachilla bracts obscure (except in monstrous forms);
staminate flowers borne in pairs. Himalayas to Bali
56. Plectocomia
- First-order inflorescence branches pendulous or not, bear-
ing bracts scarcely differentiated from other inflorescence
bracts, not obscuring rachillae; rachilla bracts conspicu-
ous; staminate flowers borne singly 20
20. Leaf sheath without an ocrea; inflorescences of both sexes
diffusely branched, usually to 3 orders; flowers with mem-
branous perianth whorls; anthers borne on long, fine,
inflexed filaments, pendulous from the lobes of the
androecial ring; fruit covered with innumerable, minute,
irregularly arranged scales. Indochina to Malay Peninsula
and Sumatra 54. Myrialepis
- Leaf sheath bearing an ocrea, sometimes disintegrating;
inflorescences of both sexes branching to 2 orders only,
very rarely diffusely branched to 3 orders; flowers with
thick coriaceous perianth whorls; anthers sessile, pendu-
lous from the tips of the lobes of the androecial ring; fruit
covered in large, regularly arranged scales. Thailand,
Burma, Malay Peninsula, Sumatra, Borneo
55. Plectocomiopsis
21. Slender, unarmed, clustered, undergrowth palms; leaf
usually irregularly divided into 1-several-fold segments;
rachillae short and compressed; staminate flowers in
dyads borne distichously on the rachillae. S America
62.Lepidoca~um
- Moderate to robust, solitary or clustered, armed or un-
armed palms; leaf regularly divided into single-fold seg-
ments; staminate rachillae and sometimes pistillate
rachillae catkinlike, bearing spirally arranged flowers,
singly or in pairs 22
22. Moderate, clustered palms with spiny stems; pistillate
flowers borne on short catkinlike branches; staminate
flowers solitary in the axil of each rachilla bract. S America
61. Mauritiella
- Massive, solitary palms with unarmed stems; pistillate
flowers very few or single, borne on scarcely catkinlike
branches; staminate flowers borne in dyads. S America
60. Mauritia
1. Tribe Calameae Drude (1881).
Erect, acaulescent or climbing, usually hapax-
anthic, hermaphroditic, polygamous, monoecious
or dioecious palms; leaves reduplicate, pinnate.
41. Laccosperma (G. Mann & H. Wendl.) Drude
Laccosperma (G. Mann & H. Wendl.) Drude, Bot. Zeit. 35:632,
635 (1877); Dransf., Kew Bull. 37: 455-457 (1982).
Calamus subgenus Laccosperma G. Mann & H. Wendl. (1864).
Hapaxanthic. Sheath spiny; ocrea split opposite
leaf, petiole present, leaflets entire. Inflorescence
prophyll enclosed within sheath; peduncular
bracts 1-3; rachis exceeding peduncle; rachis- and
rachilla-bracts distichous, tubular. Flower with
calyx stalklike basally, distally 3-lobed; petals 3,
basally connate; stamens 6, filaments distinct, an-
thers medifixed, oblong, latrorse; pollen sulcate,
exine reticulate, tectate. Fruit 1-3 seeded. Seed
subbasally attached, ovoid, laterally flattened, or
rounded, deeply scalloped, with lateral pit, em-
bryo lateral. Seven spp., W Africa and Congo
Basin.
42. Eremospatha (G. Mann & H. Wendl.)
H. Wendl.
Eremospatha (G. Mann & H. Wend!.) H. Wend!. in Kerchove,
Les Palmiers: 244 (1878).
Calamus subgenus Eremospatha G. Mann & H. Wend!. (1864).
Pleonanthic. Sheath unarmed; ocrea truncate;
petiole absent at maturity. Leaflets praemorse
or abruptly narrowed, linear to rhomboid, spiny
margined, proximal leaflets straplike, reflexed. In-
florescence with bracts inconspicuous through-
out; prophyll and peduncular bracts absent; rachis
bracts low, triangular; rachillae distichous. Flow-
ers with coriaceous, 3-lobed calyx, corolla coria-
ceous, with 3 short, lobes, remaining approximate;
stamens 6, in a fleshy, epipetalous ring, anthers
enclosed, medifixed, latrorse; pollen elliptic, sul-
cate, exine foveolate, tectate. Fruit 1-3-seeded.
Seed subbasally attached, embryo lateral. Twelve
spp. (but probably fewer), W Africa, the Congo
Basin, and eastward to Tanzania.
lb. Subtribe Eugeissoninae Becc. (1918).

Caespitose hapaxanthic, polygamous, densely

la. Subtribe Ancistrophyllinae Becc. (1918). spiny, acaulescent or tree palms; leaves pinnate;
inflorescence erect, compound, composed of axil-
Caespitose climbers, leaves cirrate, the cirrus lary inflorescences, each branched to 4 orders;
bearing acanthophylls, sheath tubular, ocrea pre- branches terminating in a cupule of 11-13 bracts,
sent, knee absent; inflorescence branched to 1 proximal 5 tubular, rest open enclosing a single
order; flowers paired, sometimes solitary, her- flower pair, consisting of a large staminate and
maphroditic; stigma pyramidal; mesocarp fleshy, lateral to it a large hermaphroditic flower, stami-
sweet, endocarp absent; .endosperm homoge- nate appearing first, then pushed out of cupule by
neous; germination adjacent-ligular; eophyll bifid. developing hermaphrodite; stamens numerous;
340
stigma pyramidal with 3 glandular angles; peri-
carp with numerous irregular scales; mesocarp
corky and fibrous at maturity, endocarp develop-
ing from a layer external to the locule wall, dark
brown to blackish, thick, with 3 + 3, or 3 + 3 + 6
flanges penetrating fruit cavity, forming sym-
metrical, incomplete partitions; germination
remote-ligular.
The single genus has no obvious close relatives.
The dyad of a hermaphroditic and a staminate
flower is found elsewhere only in Metroxylon but
there is almost nothing else to link the two genera.
Eugeissona is best regarded as having arisen prob-
ably rather early in the differentiation of the
Calamoideae. The flowers are among the largest in
the palms.
43. Eugeissona Griff.
Eugeissona Griff., Calc. J. Nat. Hist. 5: 101 (1844); J. Dransf.,
Studies in Malayan Palms Eugeissona and Johannes-
teijsmannia (1970).
Stem subterranean, erect, or borne on stilt roots.
Leaflets single-fold, entire, concolorous. Stami-
nate flowers symmetrical with short pedicel; calyx
tubular, 3-lobed; corolla tubular basally, distally
with 3 elongate, woody, valvate, spine-tipped
lobes; stamens 20-70, filaments short, anthers
elongate, basifixed, latrorse, deciduous; pollen
sulcate, with rugulate to reticulate, tectate exine;
pistillode minute. Hermaphroditic flowers with
asymmetrical tips, protandrous, sessile; ovary co-
lumnar. Fruit ovoid, beaked. Seed single, filling
fruit cavity; seed coat thin, dry; endosperm homo-
geneous; embryo basal; eophyll pinnate. Six spp.,
Malay Peninsula and Borneo.
lc. Subtribe Metroxylinae Bl. (1843).
Hapaxanthic or very rarely pleonanthic trees or
climbers; leaves pinnate; flowers solitary, her-
maphroditic, or borne in dyads of 1 staminate and
1 hermaphroditic, in spirally arranged pits on
cylindrical catkinlike rachillae; ovary with coni-
cal style with 3 stigmatic angles; germination
adjacent -ligular.
The two genera of this subtribe are very unlike
in habit but share similar rachilla form.
44. Metroxylon Rottb. Fig. 82E, 86A-V
Metroxylon Rottb., Nye Samling af det Kongelige Danske
Videnskabers Selskabs Skrifter 2: 527 ( 1783 ), nom. cons.;
Rauwerdinck, Principes 30: 165-180 (1986).
Palmae
Solitary or caespitose, hapaxanthic or pleon-
anthic, polygamous trees. Leaflets single-fold, lin-
ear, entire. Inflorescences branched to 2 orders;
rachilla bracts enclosing dyads of a staminate
and a similar hermaphroditic flower. Staminate
flowers opening before hermaphroditic; calyx tu-
bular, 3-lobed; corolla 3-lobed; stamens 6, anthers
medifixed, latrorse; pollen disulcate, exine tectate;
pistillode conical. Hermaphroditic flowers as
staminate. Fruit rounded, 1-seeded, mesocarp
dry, endocarp absent. Seed globose, invaginated
apically, sarcotesta thin, endosperm homoge-
neous; embryo basal. Germination adjacent-
ligular; eophyll bifid or pinnate. About eight
spp., native to E Malesia and W Pacific Islands;
one sp., M. sagu Rottb., thought to be native to
New Guinea and the Moluccas, now widespread
and naturalized throughout the SE Asian region.
45. Korthalsia Bl.
Korthalsia Bl., Rumphia 2: 166 ("1836") (1843); J. Dransf., Kew
Bull. 36: 163-194 (1981).
Hapaxanthic caespitose climbers. Leaves cirrate;
knee absent; ocrea present, various; leaflets single-
fold, linear to rhomboid, praemorse, often dis-
colorous. Inflorescences branching to 1-2 orders;
rachilla bracts subtending a single hermaphro-
ditic protandrous flower. Calyx tubular, 3-lobed;
corolla exceeding calyx, tubular basally, 3-lobed;
stamens 6-9; pollen sulcate, monoporate, me-
ridionosulcate, disulcate, or diporate, exine in-
tectate, spiny. Fruit 1-seeded, mesocarp fleshy,
endocarp absent. Seed with thin testa, endosperm
homogeneous or ruminate, with conspicuous pit;
embryo lateral. Germination adjacent; eophyll un-
divided or bifid, praemorse. About 26 spp., cen-
tered on the perhumid areas of the Sunda Shelf
with outliers north as far as Indochina, Burma,
and the Andaman Islands, and south-eastward to
Celebes and New Guinea.
ld. Subtribe Calaminae Meisner (1842).
Hapaxanthic (rarely) or pleonanthic, dioecious,
acaulescent or climbing; leaves pinnate, or entire-
bifid and pinnately ribbed; cirri, where present,
lacking acanthophylls; sheaths in climbing species
usually with a well-defined knee; inflorescences
axillary, frequently adnate to internode and
sheath of next leaf; staminate inflorescence with
paired or solitary flowers; pistillate inflorescence
bearing dyads of a sterile staminate flower and
 Palmae
Fig. 86A-V. Palmae-Calamoideae. A- D Metroxylon sagu. A
Compound suprafoliar inflorescence. B Single inflorescence,
branching to 2 orders, with basal2-lobed prophyll and tubular
peduncular and rachis bracts, a rachilla, b first-order branch.
C Juvenile plant of spiny form. D Fruit, showing reflexed peri-
carp scales. E- J M. vitiense. E Single inflorescence showing
basal prophyll. F Part of first-order branch with catkinlike
rachillae. G Detail of base of rachilla. H, I Seed. J Seed in
vertical section, showing embryo and large hollow in en-
dosperm. K-T M. warburgii. K Dyad of a staminate and a
341
hermaphroditic flower. L Staminate flower. M Staminate
flower in longitudinal section, showing united filaments and
pistillode. N Hermaphroditic flower in longitudinal section. 0
Stamen, ventral view. P Same, dorsal view. Q Pistil. R Same,
longitudinal section. S Same, cross-section at level of ovules.
T, U M. salomonense. T Habit, showing hapaxanthy and pen-
dulous rachillae. U Detail of leaf sheaths with grouped spines.
V M. amicarum, pleonanthic habit, inflorescences interfoliar.
(Takhtajan 1982)
342 Palmae
pistillate flower (except in Salacca sect. Leio-
salacca and Retispatha); ovary with large, fleshy
recurved stigmas; mesocarp thin; endocarp ab-
sent; sarcotesta well developed. Germination
adjacent-ligular.
The eight genera, all dioecious, have their center
of distribution in SE Asia, though Calamus itself is
very widely dispersed outside this area. Recent
work by Baker and Dransfield suggests that
Salacca and Eleiodoxa may be misplaced with the
other genera, and Calamus itself may need to be
divided into several different groups, probably at
the generic level.
46. Eleiodoxa (Becc.) Burr.
Eleiodoxa (Becc.) Burr., Notizbl. Bot. Gart. Mus. Bertin-
Dahlem 15: 733 (1942).
Salacca section Eleiodoxa Becc. (1918).
Acaulescent, caespitose, hapaxanthic. Leaves pin-
nate; leaflets single-fold, entire, concolorous. In-
florescences crowded into a head borne among
leaf bases, staminate and pistillate superficially
similar, branching to 1 order; rachillae cylindri-
cal catkinlike, bearing spirally arranged bracts.
Staminate flowers paired, embedded among hairs;
stamens 6, anthers elongate, introrse; pollen ellip-
tic, diporate, exine smooth, perforate; pistillode
absent. Pistillate flower larger than staminate;
staminodes 6. Fruit irregularly rounded; seed with
thick adherent sarcotesta; endosperm homoge-
neous, with pit at apex; embryo basal. Eophyll
bifid. Only one sp., E. conferta (Griff.) Burr.,
Sumatra, Malay Peninsula, S Thailand, and
Borneo.
47. Salacca Reinw. Fig. 80J
Salacca Reinw., Nov. Pl. Ind. Gen., in Sylloge Plantarum
Novarum 2: 3 (1826) ("1828").
Lophospatha Burr. (1942).
Acaulescent, caespitose, pleonanthic. Leaves pin-
nate or entire-bifid; leaflets single-fold, except for
compound terminal segments, concolorous or
discolorous, entire, acuminate. Inflorescences ax-
illary, emerging from a cleft in abaxial surface of
subtending petiole, spicate or branched to 1 or 2
orders; rachillae catkinlike. Staminate flowers
paired; stamens 6, epipetalous, anthers short, in-
trorse; pollen elliptic, meridionosulcate, exine
tectate, perforate with supratectal spinules. Pistil-
late flower larger than staminate. Fruit irregular;
epicarp scales smooth or with recurved spinelike
tips; endosperm homogeneous; embryo basal.
Eophyll bifid. About 20 spp., Burma and China to
Borneo and Java, several species cultivated for
their fruit.
48. Daemonorops Bl. Fig. 82I
Daemonorops Bl. in J.A. & J.H. Schultes, Syst. Veg. 7: 1333
(1830); Becc., Ann. Roy. Bot. Gard. Calcutta 12(1): 1-237
(1911).
Acaulescent or climbing, hapaxanthic or pleon-
anthic. Leaves usually cirrate, knee usually
present, leaflets single-fold, entire, concolorous.
Staminate inflorescence branched to 3 orders,
pistillate to 2, otherwise similar; prophyll con-
spicuous, 2-keeled, sometimes enclosing entire
inflorescence, splitting longitudinally; other pri-
mary bracts as prophyll, included in prophyll and
persistent (sect. Daemonorops), or exceeding
prophyll and caducous (sect. Piptospatha); flowers
distichous, staminate flowers solitary, calyx much
shorter than corolla; stamens 6, anthers elongate,
introrse; pollen disulcate or diporate, intectate or
tectate; pistillode short. Pistillate flower much
larger than staminate. Fruit 1-seeded. Endosperm
ruminate, embryo basal. Eophyll bifid or pinnate.
About 115 spp., India and S. China to New Guinea.
49. Calamus L. Fig. 80L
Calamus L., Sp. Pl. 325 (1753); Becc., Ann Roy. Bot. Gard.
Calcutta 11: 1-518 (1908) and Suppl. 11 (App): 1-142
(1913).
Acaulescent, erect or climbing, pleonanthic.
Leaves cirrate or ecirrate, knee usually present;
leaflets single-fold, linear to rhomboid, entire
or rarely praemorse, concolorous or discolorous.
Climbing whip (flagellum) sometimes present.
Staminate inflorescence branched to 3 orders, pis-
tillate to 2; prophyll strictly tubular, persistent;
other primary bracts as prophyll; flowers disti-
chous or rarely spiral. Staminate flowers solitary,
calyx much shorter than corolla; stamens 6 (rarely
12), anthers elongate, latrorse; pollen disulcate,
tectate, reticulate; pistillode short. Pistillate flower
much larger than staminate. Fruit 1-3-seeded.
Endosperm homogeneous or ruminate; embryo
basal or lateral. Eophyll bifid or pinnate. About
400 spp., one in equatorial Africa, the rest from
India and S China to Fiji and Australia.
50. Calospatha Becc.
Calospatha Becc., Ann. Roy. Bot. Gard. Calcutta 12(1): 232
(1911); Dransfield, Man. Ratt. Malay Peninsula (1979).
 Palmae 343

Short-climbing, pleonanthic. Leaves ecirrate, knee 53. Retispatha J. Dransf.

present; leaflets single-fold, linear, entire, con-
Retispatha J. Dransf., Kew Bull. 34: 529 (1979).
colorous. Staminate inflorescence branched to 3
orders, pistillate to 2; prophyll tubular at base,
Robust, short, pleonanthic climber. Leaves
with expanded 2-keeled limb and abruptly nar-
ecirrate; knee absent; leaflets single-fold, linear,
rowed tip, persistent; rachis bracts similar to
entire, concolorous. Staminate inflorescence
prophyll, distichous in a plane at right angles to it;
branched to 3 orders, pistillate to 2; prophyll tubu-
staminate flowers solitary, calyx much shorter
lar at base, with expanded limb, persistent, all pri-
than corolla; stamens 6, anthers elongate, latrorse;
mary and secondary bracts netlike; staminate
pollen elliptic, diporate, exine tectate; pistillode
flowers crowded, spiral, pistillate flowers disti-
short. Pistillate flower much larger than stami-
chous. Staminate flowers solitary, calyx much
nate. Fruit usually 3-seeded. Endosperm homo-
shorter than corolla; stamens 6, anthers short,
geneous, embryo basal. Eophyll pinnate. One
latrorse; pollen sulcate, exine tectate, reticulate
species, C. scortechinii Becc., Malay Peninsula.
with supratectal spines. Sterile staminate flower
absent. Pistillate flower much larger than stami-
51. Pogonotium J. Dransf. nate. Fruit 1-seeded. Endosperm homogenous,
embryo basal. Eophyll pinnate. Only one sp., R.
Pogonotium J. Dransf., Kew Bull. 34: 763 (1980).
dumetosa J. Dransf., Borneo.
Short, climbing, pleonanthic. Leaves ecirrate,
sheaths bearing a spiny auricle on either side of le. Subtribe Plectocomiinae J. Dransf. & N. Uhl
petiole; leaflets single-fold, linear, entire, con- (1986).
colorous. Inflorescences held between auricles,
staminate branched to 3 orders, pistillate to 2; Robust, caespitose, spiny, hapaxanthic, dioecious,
prophyll enclosing entire inflorescence, splitting climbing palms. Leaves pinnate, cirrate, leaflets
longitudinally, other primary bracts small; ra- lanceolate, entire, acanthophylls absent, knee ab-
chillae slender. Staminate flowers solitary, sent; flagellum absent, sheaths densely spiny;
calyx much shorter than corolla; stamens 6, an- rachillae very slender or highly reduced; staminate
thers elongate, latrorse; pollen elliptic, diporate, flowers paired or solitary; pistillate flowers soli-
exine tectate; pistillode short. Pistillate flower tary; fruit 1-seeded; endocarp not differentiated;
slightly larger than staminate. Fruit 1-seeded. En- seed attached near the base, sarcotesta thick but
dosperm homogeneous, embryo basal. Eophyll not juicy; endosperm homogeneous; embryo
bifid or pinnate. Three spp., Borneo and Malay basal; germination adjacent-ligular.
Peninsula. This subtribe includes three genera of massive
hapaxanthic climbing palms restricted to main-
land SE Asia and W Malesia. The group is distin-
52. Ceratolobus Blume Fig. 81W
guished by the solitary pistillate flowers, not
Ceratolobus Blume in J.A. & J.H. Schultes, Syst. Veg. 7: 1334 accompanied by sterile staminate flowers.
(1830); Dransfield, Kew. Bull. 34: 1-33 (1979).

Slender, caespitose, pleonanthic climbers. Leaves 54. Myrialepis Becc.

cirrate, knee present; leaflets single-fold, linear,
entire, or rhomboid and praemorse, concolorous
or discolorous. Staminate inflorescence branched
to 3 orders, pistillate to 2; prophyll flattened, en-
closing inflorescence, beaked, opening apically by
2 lateral pores; other primary bracts diminutive.
Staminate flowers solitary, calyx shorter than co-
rolla; stamens 6, anthers elongate, latrorse; pollen
elliptic, disulcate, exine tectate; pistillode short.
Pistillate flower larger than the staminate. Fruit 1-
seeded. Endosperm homogeneous or ruminate,
embryo basal. Eophyll with several crowded
leaflets. Six spp., Malay Peninsula, Sumatra,
Borneo, Java.
Myrialepis Becc. in J.D. Hook., Fl. Br. India 6: 480 (1893);
Dransfield, Kew. Bull. 37: 237-254 (1982).
Ocrea absent; leaflets concolorous. Inflorescences
branched to 3 orders; staminate rachillae bearing
groups of up to 8 distichous flowers, pistillate
rachillae bearing groups of up to 7 flowers. Stami-
nate flowers symmetrical; calyx membranous, tu-
bular, 3-lobed; corolla deeply 3-lobed; stamens 6,
filaments connate laterally, inflexed, anthers
oblong, introrse; pollen elliptic, disulcate, with
finely reticulate, tectate exine; pistillode minute.
Pistillate flowers larger than staminate. Epicarp
covered in minute, irregularly arranged scales,
344
mesocarp thin, fibrous; embryo basal. Only one
sp., M. paradoxa (Kurz) J. Dransf., widespread
in Indochina, Burma, Thailand, Malaya, and
Sumatra.
55. Plectocomiopsis Becc.
Plectocomiopsis Becc. in J.D. Hook., Fl. Br. India 6: 479 (1893);
Dransfield, Kew. Bull. 37: 237-254 (1982).
Ocrea present; leaflets concolorous. Inflores-
cences branched to 2(3) orders. Staminate flowers
in clusters of up to 32, each flower with rachilla
bract and a 2-keeled bracteole; calyx thick, 3-
lobed; corolla thick, split distally with 3 approxi-
mate lobes; stamens 6, epipetalous, forming a tube
with 6 short, reflexed, filaments bearing rounded,
medifixed, introrse anthers; pollen disulcate,
exine tectate; pistillode minute. Pistillate flowers
rarely solitary, or borne in groups of 2-4. Epicarp
scales somewhat irregular, smooth. Eophyll bifid.
Five spp., S Thailand, Malaya, Sumatra, Borneo.
56. Plectocomia Mart. ex Bl.
Plectocomia Mart. ex Bl. in J.A. & J.H. Schultes, Syst. Veg. 7:
1333 (1830); Madulid, Kalikasan 10: 1-94.
Ocrea absent; leaflets concolorous or discolorous.
Inflorescences branched to 2 orders; first-order
branches with conspicuous, distichous distal
bracts splitting longitudinally almost to the base,
enclosing rachillae; rachillae slender, in staminate
bearing 2-ca. 100 flowers borne in dyads, in pistil-
late (1)2-10 flowers borne singly. Staminate flower
with tubular 3-lobed calyx; corolla tubular, 3-
lobed; stamens 6(12), anthers elongate, latrorse;
pollen disulcate, exine tectate; pistillode absent.
Pistillate flowers often pedicellate. Epicarp scales
with tips often fringed, upward-pointing. Eophyll,
simple, lanceolate. About 16 spp., from the
Himalayas, S China and Hainan, south through
Burma and Indochina to the Sunda Shelf and the
Philippines; not known east of Wallace's Line.
If. Subtribe Pigafettinae J. Dransf. & N. Uhl
(1986).
Massive, single-stemmed, spiny, pleonanthic,
dioecious tree palms; leaves pinnate; rachillae
slender, bearing highly reduced bracts; staminate
flowers paired; pistillate flowers solitary; fruit 1-
seeded; mesocarp thin, endocarp lacking; seed
basally attached, laterally flattened, with thick
Palmae
sarcotesta, endosperm homogeneous, shallowly
pitted; embryo lateral, opposite pit; germination
adjacent -ligular.
The subtribe consists of a single genus without
obvious close relatives. The rachilla structure is,
however, reminiscent of that of Plectocomia.
57. Pigafetta (Bl.) Becc.
Pigafetta (Bl.) Becc., Malesia 1: 89 (1877), nom. cons.
("Pigafettia").
Sagus section Pigafetta Bl., Rumphia 2: 154 (1843); Dransfield,
J., Principes 42: 34-40 (1998).
Leaflets single-fold, concolorous. Inflorescences
interfoliar at anthesis, branching to 2 orders;
prophyll tightly sheathing; peduncular bracts ca.
8, tubular; flower-bearing part of rachilla exserted
on bare basal stalk. Staminate flowers symmetri-
cal; calyx 3-lobed; corolla exceeding calyx, 3-
lobed; stamens 6, borne at mouth of corolla tube,
filaments connate laterally in a low ring, anthers
elongate, latrorse; pollen inaperturate, with finely
reticulate, tectate exine; pistillode minute. Pistil-
late flowers globose. Fruit small, ovoid. Eophyll
bifid. Two spp., Sulawesi to New Guinea.
1g. Subtribe Raphiinae J.D. Hook. in Benth. &
J.D. Hook. (1883).
Massive hapaxanthic monoecious armed acaules-
cent or tree palms; leaves massive, pinnate;
flowers solitary, dimorphic; staminate distal on
the rachilla, unibracteolate, pistillate proximal,
bibracteolate; ovary with short conical 3-lobed
stigma; fruit 1-seeded, mesocarp thick, mealy, oil-
rich, endocarp not differentiated; seed with dry
seed coat; endosperm with few large ruminations;
embryo lateral; germination adjacent-ligular.
Raphia, the only genus in the subtribe, has
no obvious close relatives. Phylogenetic work in
progress (Baker and Dransfield, in prep.) may
shed new light on its relationships.
58. Raphia P. de Beauv.
Raphia P. de Beauv., Flore d'Oware et de Benin, en Afrique 1:
75 t. 44-46 (1806); Otedoh, J. Nig. Inst. Oil Palm Res. 6(22):
145-189 (1982).
Sagus J. Gaertn. (1788) (non Sagus Steck)
Solitary or caespitose; sheath disintegrating into
thin sheets or black fiber bundles; leaflets single-
fold, linear, entire. Inflorescences branched to 2
orders, either interfoliar and pendulous, or aggre-
 Palmae
gated into a massive, erect, suprafoliar, compound
inflorescence; rachillae bearing distichous or 4
ranked bracts subtending flowers. Staminate flow-
ers conspicuously exserted; stamens 6-30, anthers
elongate, latrorse; pollen elliptic, sulcate, with
scabrate, tectate exine. Pistillate flowers very dif-
ferent from staminate, partly exserted from the
rachilla bracts. Eophyll usually pinnate, rarely
bifid. About 28 spp., humid areas of Africa; one
sp. in Madagascar, possibly introduced; one sp.,
R. taedigera Mart., in tropical America.
Ih. Subtribe Oncocalaminae J. Dransf. & N. Uhl
(1986).
Caespitose, pleonanthic, monoecious climbers;
cirri bearing acanthophylls; rachilla bracts sub-
tending complex groups of flowers comprising a
central pistillate flower and 2 lateral cincinni with
0-2 basal pistillate flowers and distal staminate
flowers; endosperm homogeneous, laterally pen-
etrated by smooth-margined postament; embryo
lateral.
The strange flower cluster of Oncocalamus is
unique, not only in the subfamily, but indeed
within the whole family. Vegetatively, Onco-
calamus is certainly very similar to Eremospatha
and Laccosperma of the Ancistrophyllinae. We
have removed Oncocalamus from this group,
which includes other endemic African rattan
genera, because of the very different flower clus-
ters and unisexual flowers. Recent phylogenetic
studies (Baker and Dransfield, in prep.) suggest
that the three African rattan genera are, in fact,
more closely related than is suggested here.
59. Oncocalamus (G. Mann & H. Wendl.) G.
Mann & H. Wendl.
Oncocalamus (G. Mann & H. Wendl.) G. Mann & H. Wendl. ex
J.D. Hook. in Benth. & J.D. Hook., Gen. Pl. 3: 881, 936
(1883).
Calamus subgenus Oncocalamus G. Mann & H. Wendl. (1864).
Sheath bearing scattered, brittle spines; ocrea
truncate; knee absent; leaflets entire, acute, lan-
ceolate. Inflorescences branching to 1 order; first-
order branches pendulous, bracts distichous, each
enclosing a flower cluster. Staminate flowers sym-
metrical; calyx stalked basally, 3-lobed; corolla
valvate, only slightly exceeding calyx, 3-lobed;
stamens 6, filaments united, anthers rounded,
latrorse; pollen elliptic, sulcate, exine tectate;
pistillode conical. Pistillate flowers superficially
345
similar; gynoecium ellipsoid. Fruit spherical. Five
spp., described from equatorial west Africa and
the Congo Basin, but probably fewer.
2. Tribe Lepidocaryeae Mart. (1837).
Erect pleonanthic dioecious shrub or tree palms
with palmate or briefly costapalmate leaves, with
single-fold or compound segments.
This tribe of three New World genera is immedi-
ately distinguished from Calameae by the palmate
or costapalmate leaf.
60. Mauritia L. f. Fig. 81E,F
Mauritia L. f., Suppl. Plant. 70, 454 (1782); Henderson, Palms
of Amazon: 70-74 (1994).
Massive, solitary, unarmed. Staminate rachilla
catkinlike, bearing crowded, spirally inserted
bracts, each subtending a pair of flowers, each
with a bracteole; pistillate rachilla very short, not
catkinlike, bearing solitary pistillate flowers.
Staminate flowers with tubular, 3-lobed calyx;
petals 3, elongate; filaments free, anthers elongate,
latrorse; pollen sulcate or monoporate, exine
intectate with spines; pistillode minute. Pistillate
flowers larger than staminate. Fruit rounded,
large. Seed apically with a blunt beak; embryo
basal. Eophyll with 2 divergent leaflets. Two spp.,
wetter parts of tropical S America.
61. Mauritiella Burr.
Mauritiella Burr., Notizbl. Bot. Gart. Mus. Berlin-Dahlem 12:
609 (1935); Henderson, Palms of Amazon: 74-77 (1994).
Moderate, caespitose. Stem bearing spinelike ad-
ventitious roots. Staminate and pistillate rachillae
catkinlike, each bract subtending a single flower;
pistillate rachilla very short. Staminate flower
symmetrical; calyx 3-lobed; corolla 3-lobed, much
exceeding calyx; filaments distinct, anthers elon-
gate, latrorse; pollen sulcate, exine intectate with
spines; pistillode minute. Pistillate flowers larger
than staminate. Fruit rounded. Seed rounded to
ellipsoid, apically with an elongate knob; embryo
basal. Eophyll with 2 divergent leaflets. Three spp.,
northern South America.
62. Lepidocaryum Mart.
Lepidocaryum Mart., Hist. Nat. Palm. 2:49 (1824}; Henderson,
Palms of Amazon: 77-79 (1994}.
346
Slender, caespitose, unarmed, spreading slender
rhizomes. Staminate and pistillate rachillae short,
bracts subtending a solitary flower. Staminate
flowers symmetrical; calyx 3-lobed; petals exceed-
ing calyx, 3-lobed; filaments fleshy, anthers small,
basifixed, latrorse; pollen sulcate or monoporate,
exine intectate spiny; pistillode minute or short,
columnar. Pistillate flowers larger than staminate.
Fruit rounded or oblong. Embryo lateral. Eophyll
bifid. Only one sp. with three varieties, wetter
parts of Colombia, Peru, Venezuela, Guyana, and
Brazil.
III. Subfamily Nypoideae Griff. (1850).
Moneocious, pleonanthic; inflorescences solitary,
interfoliar, erect, terminating in a head of pistillate
flowers and below this bearing 7-9 spirally ar-
ranged, inflated, tubular bracts each subtending a
first-order branch; first-order branches adnate ca.
1/2 of their length above the subtending bracts,
each bearing and enclosed by a tubular prophyll in
bud; subsequent branches all bearing a closed
prophyll and ending in a short catkinlike rachilla,
bearing spirally arranged, solitary staminate flow-
ers, each subtended by a small bract. Staminate
flowers sessile; sepals 3, distinct; petals 3, distinct,
slightly imbricate, similar to the sepals but slightly
smaller, both loosely closed over the stamens in
bud; stamens 3, filaments and connectives con-
nate in a solid stalk, anthers elongate, extrorse;
pollen elliptic, meridionosulcate, exine tectate,
spiny; pistillode lacking. Pistillate flowers differ-
ent; sepals 3, distinct, petals 3; staminodes lacking;
carpels 3( -4), distinct, much longer than and ob-
scuring the perianth at maturity, ± obovoid, with
a + lateral, funnel-shaped stigmatic opening,
ovule anatropous. Fruiting head ± globose, 1-3
carpels per flower maturing a seed; fruit develop-
ing from 1 carpel, stigmatic remains terminal,
pyramidal; epicarp smooth, mesocarp fibrous,
endocarp thick, composed of interwoven fibrous
strands, with an adaxial internal longitudinal
ridge intruded into the seed. Seed broadly ovoid,
grooved adaxially, endosperm homogeneous; em-
bryo basal. Germination on the head with the plu-
mule exserted and pushing the fruit away; eophyll
bifid or with several leaflets.
The mangrove palm, N. fruticans, is remarkably
disjunct morphologically and has an extensive
fossil record. Characteristic Nypa pollen in the
Senonian of tropical West Africa and Borneo
represent some of the earliest records of palms
(Harley et al. 1991; Harley 1996). Cladistic analy-
Palmae
ses of morphological and molecular characters
have resolved Nypa as separate from all other
palms (Uhl et al. 1995). It appears to represent one
of the earliest lineages in the family. Its cupular,
somewhat ascidiform carpel may be different
from the follicular forms in other palms and
monocotyledons and is of special interest.
Fig. 81M, 82L, 870-X
63. Nypa Steck
Nypa Steck, Dissertatio Inauguralis Medica de Sagu, 15 (1757),
Uhland Dransf., Gen. Palm.: 285-288 (1987).
Nipa Thunb. (1782).
Large, creeping, unarmed. Stem stout, prostrate or
subterranean, branching dichotomously, curved
leaf scars evident above, roots borne along the
lower side. Leaves few, erect, reduplicately pin-
nate; sheath splitting, glabrous; petiole stout,
elongate, the base often persistent as a conical
stub; leaflets numerous, single-fold, regularly
arranged, acute, coriaceous; midrib bearing
shining, chestnut-colored, membranous ramenta
abaxially. Only one sp., N. fruticans Wurmb., from
Sri Lanka and the Ganges Delta to Australia, the
Solomon Is., and the Ryukyu Is.
IV. Subfamily Ceroxyloideae Drude (1881).
Very small to tall, unarmed, solitary or caespitose,
pleonanthic; leaves reduplicate, pinnate or entire
and pinnately ribbed; inflorescence with prophyll
and numerous (very rarely 1 only) peduncular
bracts; flowers solitary or, if grouped, not in dyads
or triads but in acervuli, when unisexual not strik-
ingly dimorphic.
The subfamily comprises three tribes. Recent
phylogenetic work (Uhl et al. 1995) suggests that
the subfamily may not be monophyletic; tribe
Hyophorbeae seems to be related to Arecoideae
while the position of the other two tribes seems
to be basal in a clade comprising most pinnate-
leaved palms apart from Nypa and Calamoideae.
As a new phylogeny is still not yet resolved, the
arrangement of genera follows that in Genera
Palmarum.
KEY TO THE GENERA OF CEROXYLOIDEAE
1. Flowers hermaphroditic 64. Pseudophoenix
- All flowers unisexual 2
2. Dioecious; crownshafts absent; flowers borne singly on
short stalks, bracteolate; flowers open from early in devel-
opment, stamens equalling or exceeding petals 3
 Palmae 347

(?(
N\_J

- Monoecious or dioecious; crownshafts present or absent; Fig. 87 A-X. Palmae-Nypoideae. Nypa fruticans. A Habit,
flowers sessile, usually ebracteolate at anthesis, borne singly showing infructescences. B Detail of apical portion of inflores-
or in acervuli of a pistillate and 2 to several staminate, cence showing central terminal pistillate head and lateral
or in lines of staminate or, rarely, pistillate flowers catkinlike staminate rachillae. C Pistillate flower head with 1
(Wendlandiella); flowers closed in bud; stamens included staminate rachilla. D Portion of staminate rachilla in cross-
6 section. E Staminate flower with distinct perianth segments. F
3. Stigmatic remains basal in fruit 4 Same in longitudinal section. G Sepal and H petal of staminate
- Stigmatic remains lateral to subapical in fruit 5 flower. I Connate stamens. JCross-section of anthers. K Pistil-
4. Petals basally united; stamens 6-15 or more. Andes of late flower. L Pistillate flower, carpels removed. M Carpel. N
S America 65. Ceroxylon Same in longitudinal section. 0 Infructescence. P Fruit. Q
- Petals free, valvate; stamens 6. Queensland Same in longitudinal and R cross-section, seed stippled. S, T
66. Oraniopsis Seed in two views. U Germinating fruit. V Same, longitudinal
5. Pistillate flower with staminodes bearing rudimentary section showing (a) first scale leaves and (b) enlarging cotyle-
anthers; staminate inflorescence often multiple; prophyll don. W Seedling. X Dichotomously branched rhizome.
incomplete. Madagascar 68. Ravenea (Takhtajan 1982)
348 Pal mae

- Pistillate flower with staminodes lacking rudimentary an- The tribe includes the single genus Pseudo-
thers; staminate inflorescence solitary; prophyll complete. phoenix, confined to the Caribbean region.
Juan Fernandez Is. 67. ]uania
6. Trees of medium size 7
- Understory palms 8 64. Pseudophoenix H. Wendl. ex Sargent
7. Crownshaft present; inflorescences infrafoliar in bud,
rachillae pendulous. Mascarene Islands 70. Hyophorbe Fig. 88A-E
- Crownshaft lacking; inflorescences interfoliar in bud, be- Pseudophoenix H. Wend!. ex Sargent, Bot. Gaz. 11: 314 (1886);
coming infrafoliar, often persistent, rachillae divaricate. Read, Gentes Herb. 10: 169-213 (1968).
Cuba, Puerto Rico, Mexico, Guatemala, Belize Cyclospathe O.F. Cook (1902).
69. Gaussia
8. Dioecious understory palms 9
- Monoecious, flowers in acervuli. Southern Mexico, Central Leaflets acute, single-fold, entire. Inflorescences
America, and northwestern S America interfoliar, branching to 5 orders; prophyll persis-
71. Synechanthus
9. Peduncular bracts several, flowers solitary or rarely in
curved lines, sepals and petals free or connate, endocarp
hard. Central Mexico to Brazil and Bolivia
72. Chamaedorea
- Peduncular bracts 1, both staminate and pistillate flowers
in acervuli, sepals and petals always connate, endocarp
membranous. Amazonia 73. Wendlandiella

1. Tribe Cyclospatheae O.F. Cook {1902).

Hermaphroditic, solitary; sheaths forming short

crownshafts; flowers borne singly, stalked, spirally
arranged; filament apex lying in a groove in
abaxial surface of anther to about midpoint of
connective, then bent sharply inward; gynoecium
trilocular, triovulate, with 3 glands at base-
opposite petals, stigmas sessile, short; ovules
campylotropous, inserted on adaxial side of
locule; seed with basal hilum; endosperm homo-
geneous; embryo subbasal; germination
remote-tubular.

Fig. 88A-B'. Palmae-Ceroxyloideae (A-L), Arecoideae (M-

B'). A-E Pseudophoenix sargentii. A Portion of rachilla with
staminate flowers. B Hermaphroditic flower. C Same, longitu-
dinal section. D Ovary in cross-section at level of ovules. E
Two-seeded fruit. F Ceroxylon alpinum, staminate flower
viewed from above. G Hyophorbe verschaffeltii, detail of base
of crown, showing base of crownshaft and infrafoliar inflores-
cences. H-L Synechanthus warscewiczianus. H portion of
rachilla showing acervuli, each with a basal pistillate flower
and distal staminate flowers. I Staminate flower in longitudi-
nal section. J Pistillate flower. K Fruit in longitudinal section
showing ruminate endosperm. L Seed. M-S Socratea
exorrhiza. M Habit, showing stilt roots, crownshaft, and
infrafoliar inflorescences. N Portion of rachilla at anthesis of
staminate flowers. 0 Staminate flower in longitudinal section.
P Pistil. Q Same in cross-section at level of ovules. R Fruit. S
Seed. T Praemorse leaflet. U Part of stilt root with prickly
pneumathodes. V, W Wettinia quinaria, ovary with 1 fertile
locule and basal style in lateral view and W in cross-section.
X-B' Podococcus barteri. X Habit. Y Portion of inflorescence
with flower buds. Z Pistillate flower just after fertilization. A'
2-lobed fruit, each lobe with a seed. B' Seed with adherent
fibres. (Takhtajan 1982)
 Palmae
tent; peduncular bracts 2, first similar to prophyll,
second collarlike. Flowers stalked; calyx 3-lobed;
petals 3, valvate, much longer than calyx, basally
connate; stamens 6, connate basally, anthers,
elongate, dorsifixed, latrorse; pollen elliptic, sul-
cate, exine tectate, reticulate. Fruit 1-3-seeded,
globose or 2-3-lobed, stigmatic remains near base;
epicarp smooth, mesocarp fleshy, with raphides,
lacking fibers, endocarp hard, smooth. Eophyll
narrow lanceolate. Four spp., Florida, the Bahama
Islands, Cuba, Hispaniola, and Dominica to
Mexico and Belize.
2. Tribe Ceroxyleae Satake (1962).
Often tall, solitary, unarmed, pleonanthic, dioe-
cious palms; sheaths not forming a crownshaft;
leaflets narrow, single-fold; inflorescences inter-
foliar, prophyll 2-keeled, flattened, open apically;
flowers open from early in development, borne
singly along the rachillae, pedicellate; gynoecium
trilocular, triovulate; ovules pendulous, hemiana-
tropous; endosperm homogeneous.
The four genera of the tribe are very similar in
many morphological aspects and the monophyly
of the tribe is well supported (Uhl et al. 1995). The
greatest variation in habit and morphology is to
be found in Ravenea, in which some species are
highly reduced (e.g., the slender R. hildebrandtii
Bouche), and some have condensed inflorescences
perhaps adapted to beetle pollination. The distri-
bution of the genera of the tribe, widely separated
in the Southern Hemisphere, is of great biogeo-
graphic interest.
65. Ceroxylon Bonpl. ex DC.
Ceroxylon Bonpl. ex DC., Bull. Sc. Soc. Phil. 3: 239 (1804);
Galeano, Caldasia 17 (82-85): 395-407 (1995).
Tall. Leaflets usually discolorous. Inflorescences
solitary, branched to 3-4 orders; prophyll incom-
plete; lower peduncular bracts short, upper 3-4
completely enclosing inflorescence in bud;
rachillae short, flexuous. Staminate flowers with
3-lobed calyx; petals 3, connate basally; stamens
6-15(-17), anthers basifixed; pollen sulcate, exine
tectate, reticulate; pistillode minute. Pistillate
flowers similar to staminate. Fruit 1-seeded, stig-
matic remains lateral near base. Seed globose;
embryo lateral near base. Germination adjacent-
ligular; eophyll lanceolate. Eleven spp., high
elevations in the Andes from Venezuela through
Colombia, Ecuador, and Peru to Bolivia.
349
66. Oraniopsis (Becc.) J. Dransf., Irvine &
N. Uhl.
Oraniopsis (Becc.) J. Dransf., Irvine & N. Uhl, Principes 29(2):
57 (1985).
Orania Zipp. subgenus Oraniopsis Becc. in Becc. & Picchi-
Sermolli (1955).
Leaflets discolorous. Inflorescences solitary,
branching to 4 orders; prophyll incomplete;
peduncular bracts 3-5, distal2-3 enclosing inflo-
rescence in bud. Staminate flowers with calyx 3-
lobed; petals 3, distinct; stamens 6, antepetalous
epipetalous, filaments tapering, anthers oblong,
versatile, latrorse; pollen elliptic, sulcate, exine
tectate; pistillode small. Pistillate flowers like
staminate. Fruit 1-seeded, rounded, stigmatic re-
mains basal. Seed rounded, embryo lateral to sub-
basal. Germination adjacent-ligular; eophyll bifid.
Only one sp., Oraniopsis appendiculata (F.M.
Bail.) J, Dransf., Irvine & N. Uhl, Queensland,
Australia.
67. ]uania Drude
juania Drude, Nachr. Kon. Ges. Wiss. 1878 (1): 40 (1878);
Moore, Gentes Herb. 10: 385-393 (1969).
Leaflets discolorous. Inflorescences solitary,
branched to 2 orders; prophyll complete; pedun-
cular bracts 3, similar to prophyll, second enclos-
ing inflorescence in bud. Staminate flowers with
calyx 3-lobed; petals 3, distinct, imbricate basally;
stamens 6, filaments distinct, anthers basifixed,
versatile, latrorse; pollen elliptic, sulcate, exine
tectate reticulate; pistillate flowers similiar to
staminate, staminodes lacking abortive anthers.
Fruit globose with subapical stigmatic remains.
Seed globose; embryo lateral near the base. Germi-
nation adjacent-ligular; eophyll lanceolate. Only
one sp., ]uania australis (Mart.) Drude ex Hook.f.,
on Juan Fernandez Is.
68. Ravenea Bouche ex H. Wendl.
Ravenea Bouche ex H. Wend!. in J.D. Hook., Bot. Mag. 110: t.
6776 (1884); Beentje, Kew Bull. 49: 623-671 (1994); Dransf.
& Beentje, Palms Madag.: 71-113 (1995).
Slender to robust. Leaflets usually concolorous.
Inflorescences 1 or several within a common in-
complete prophyll, branching to 1-2 orders; pe-
duncular bracts 3-5, basal2 short, distal enclosing
inflorescence in bud. Staminate flower with 3-
lobed calyx; petals 3; stamens 6, anthers latrorse;
pollen monoporate, exine tectate, reticulate or
350
spiny; pistillode short. Pistillate flowers similar
to staminate; staminodes 6, with empty anthers.
Fruit globose, 1-3-seeded, stigmatic remains
subbasal, lateral, or subapical. Seed globose or
angled; embryo basal. Germination adjacent-
ligular or remote-ligular; eophyll bifid or pinnate.
Seventeen spp., 15 in Madagascar, two in the
Comoro Islands.
3. Tribe Hyophorbeae Drude (1881).
Robust to slender, acaulescent or erect, rarely
climbing, unarmed; monoecious or dioecious;
flowers solitary or in acervuli with basal pistillate
and 2-several distal staminate flowers, or in lines
of staminate or pistillate flowers. Gynoecium
trilocular, triovulate. Germination adjacent-
ligular.
Monophyly of the tribe is well supported (Uhl et
al. 1995) but the relationships of the tribe remain
unresolved. The tribe includes stout tree palms
and some of the smallest of all palms. Most mem-
bers of the tribe are found in Central and nor-
thern South America; Hyophorbe, confined to the
Mascarene Is. in the Indian Ocean represents a
remarkably, disjunct outlier.
69. Gaussia H. Wendl.
Gaussia H. Wend!., Nachr. Konig. Ges. Wiss.: 327 (1865);
Quero & Read, Syst. Bot. 11: 145-154 (1986).
Solitary monoecious trees. Crownshaft absent.
Leaflets acute, single-fold. Inflorescences inter-
foliar or infrafoliar, branching to 2-3 orders,
protandrous; peduncular bracts 4-7; flowers
borne in acervuli. Staminate flowers with 3,
rounded, imbricate, sepals; petals 3, valvate;
stamens 6, filaments distinct, anthers dorsifixed;
pollen elliptic, sulcate, exine tectate. Pistillate
flowers with sepals as staminate; petals 3,
imbricate basally, valvate distally; staminodes 6,
toothlike; gynoecium with 3 recurved stigmas,
ovules lateral, campylotropous. Fruit with
basal stigmatic remains; endocarp absent. En-
dosperm homogeneous; embryo lateral. Five spp.,
Mexico, Guatemala, Belize, Cuba, and Puerto
Rico.
70. Hyophorbe J. Gaertn. Fig. 88G
Hyophorbe J. Gaertn., De Fructibus 2: 186 (1791); Moore,
Gentes Herb. 11: 212-245 (1978).
Palmae
Solitary, monoecious trees. Stem often swollen.
Crownshaft present; leaflets acute, single-fold. In-
florescences infrafoliar, solitary, branched to 3-4
orders, protandrous; peduncular bracts 5(4-9).
Flowers borne in acervuli. Staminate flowers sym-
metrical; sepals 3, distinct, or basally connate;
petals 3, valvate; stamens 6, anthers dorsifixed,
latrorse; pollen elliptic, sulcate, exine tectate;
pistillode conical. Pistillate flowers symmetrical;
sepals 3, distinct, imbricate; staminodes forming
6-lobed cupule; ovules lateral, hemianatropous.
Fruit 1-seeded, stigmatic remains basal. En-
dosperm homogeneous; embryo lateral to apical.
Eophyll bifid or pinnate. Five spp., endemic to the
Mascarene Is.
71. Synechanthus H. Wendl. Fig. 88H-L
Synechanthus H.A. Wend!., Bot. Zeit. 16: 145 (1858); Moore,
Principes 15: 10-19 (1971).
Slender solitary or clustered, monoecious.
Crownshaft absent; leaflets with 1-several folds,
or blade entire-bifid. Inflorescences branched to 1
or 2 orders, protandrous; peduncular bracts 4-5.
Flowers in acervuli. Staminate flowers with 3-
lobed calyx; petals 3, valvate; stamens 6, with
short filaments, or 3, with long filaments, anthers
basifixed, latrorse; pollen sulcate, exine tectate.
Pistillate flowers with 3-lobed calyx; petals 3, dis-
tinct, imbricate; ovules lateral, campylotropous.
Fruit with basal stigmatic remains. Endosperm
homogeneous or ruminate; embryo lateral to sub-
apical. Eophyll bifid. Two spp., Central and north-
western S America.
72. Chamaedorea Willd. Fig. 80F, 81Q
Chamaedorea Willd., Sp. Pl. 4(2): 638, 800 (1806), nom. cons.;
Hodel, The Chamaedorea Palms (1992); Henderson et a!.,
Field Guide: 79-104 (1995).
Morenia Ruiz & Pavon (1794), nom. rejec.
Small, erect, procumbent, climbing or acaules-
cent, solitary or clustered, dioecious. Leaves
entire, bifid or variously pinnate; crownshaft
sometimes present. Inflorescences solitary or
multiple, spicate or branched to 1(-2) orders; pe-
duncular bracts 2-several. Flowers sessile, solitary
or in acervuli. Staminate flowers with 3 sepals,
united basally or distinct; petals 3; stamens 6,
anthers dorsifixed; pollen sulcate, exine tectate;
pistillode various. Pistillate flower with sepals as
in staminate; petals 3, usually connate; ovule
campylotropous, lateral. Fruit with basal stigmatic
 Palmae 351

remains. Endosperm homogenous; embryo basal 3. Inflorescence rarely spicate, usually branched to 1-2 orders;
or lateral. Eophyll bifid or pinnate. Seventy-seven flowers sessile, not sunken in pits 2. Iriarteeae (p. 352)
- Inflorescence spicate; flowers sunken in pits and exserted at
to over 110 spp., depending on species concept, anthesis on elongate receptacles 3. Podococceae (p. 354)
from central Mexico to Brazil and Bolivia. 4. Flowers usually superficial, rarely enclosed in pits; petals of
staminate flowers free and valvate; pistillate petals imbri-
cate with minutely to conspicuously valvate apices, rarely
73. Wendlandiella Damm. basally connate or valvate, styles not elongate 5
- Flowers always sunken in pits in the rachillae; petals of both
Wendlandiella Damm., Bot. Jahrb. Syst. 36 (Beibl. 80): 31 staminate and pistillate flowers connate basally in a soft
(1905); Henderson, Palms of Amazon: 105-106 (1994). tube, the lobes valvate, styles elongate, conspicuous
6. Geonomeae (p. 384)
Dwarf, slender clustering, dioecious. Leaves pin- 5. Gynoecium usually pseudomonomerous, only very rarely
nate or entire bifid; crownshaft absent. Inflores- triovulate, when triovulate, the fruit rarely more than 1-
seeded, if so, then fruit lobed; fruit with a thin or rarely
cences interfoliar, branching to 1(-2) orders;
thick endocarp, sometimes with a basal operculum but
peduncular bract 1. Staminate flowers in acervuli lacking 3 or more pores 4. Areceae (p. 354)
of 2-6; sepals 3, connate; petals 3; stamens 6, - Gynoecium trilocular, triovulate, the fruit never lobed; fruit
filaments distinct, anthers short; pollen elliptic, almost always with a thick bony endocarp enclosing 1-3,
sulcate, exine tectate reticulate; pistillodes 3. Pis- rarely more, seeds and with 3, or rarely more, clearly
defined pores 5. Cocoeae (p. 377)
tillate flowers solitary or in vertical pairs; calyx
cupular, 3-lobed; petals 3, imbricate; staminodes
3, minute; gynoecium subglobose. Fruit with basal
stigmatic remains. Endosperm homogeneous; 1. Tribe Caryoteae Drude (1881).
embryo lateral. Only one sp. with three varieties,
W. gracilis Wendl., western Amazon region. Small to very robust, unarmed; leaves induplic-
ately imparipinnate or bipinnate or rarely entire
and induplicately ribbed; sheath extended beyond
V. Subf. Arecoideae Griff. (1845). the petiole to form a ligule, eventually disintegrat-
ing into a mass of black fibers; distal margins of
Monoecious; leaves pinnate, pinnately ribbed or leaflets praemorse; peduncle bearing prophyll and
bipinnate, induplicate or reduplicate; flowers several conspicuous, spirally arranged, peduncu-
borne in triads of a central pistillate flower and lar bracts, splitting adaxially; flowers superficial;
2 lateral staminate, or a grouping derived from a pistillode absent in staminate flower; gynoecium
triad. trilocular, triovulate, ovules inserted adaxially at
The Arecoideae include six diverse tribes. base, hemianatropous; fruit with apical stigmatic
Caryoteae are distinctive in having induplicate remains, epicarp .smooth, mesocarp fleshy, with
leaves and many members are hapaxanthic, al- abundant raphides, endocarp not differentiated;
though pleonanthy also occurs. The inclusion of seed coat black.
this tribe in the subfamily has been disputed and This Old World tribe, immediately recognizable
is in need of verification. All other tribes have by the induplicate nature of the leaves, includes
reduplicate leaves and pleonanthic flowering three easily recognized genera. Generic bound-
behavior. aries have recently become less clear with the
discovery of Caryota ophiopellis J. Dowe, that
KEY TO THE TRIBES OF ARECOIDEAE combines features of Caryota with some of
1. Palms frequently hapaxanthic with basipetal production Arenga. Basipetal hapaxanthy is a very character-
of inflorescences; leaves pinnate, or doubly pinnate, istic feature of the tribe; however, a few species of
induplicate, leaflets praemorse; inflorescences bisexual or Arenga are, in fact, acropetal pleonanthic. The
unisexual by reduction of triads I. Caryoteae three genera have often been regarded as consti-
- Palms never hapaxanthic; leaves pinnate or pinnately
ribbed, reduplicate, the leaflets various, sometimes
tuting a separate subfamily. They are included
praemorse; inflorescences rarely unisexual 2 here within Arecoideae because of the presence of
2. Leaflets praemorse at the tips and with several principal ribs triads or triad derivatives in the inflorescence and
divergent from the base, or leaflets sometimes divided lon- the manifest similarity in inflorescence form be-
gitudinally into 1-several ribbed parts; inflorescence with tween them and some members of Arecoideae
a prophyll and more than 2 large peduncular bracts 3
- Leaflets usually acute, or if praemorse or longitudinally di-
(such as Iriartea). The inclusion of the tribe in
vided, the inflorescence with a prophyll and (0)1(2) large Arecoideae is still questioned and needs to be
peduncular bracts 4 confirmed with new phylogenetic studies.
352 Palmae
KEY TO THE GENERA OF CARYOTEAE
1. Leaf bipinnate; inflorescences always bearing flowers of
both sexes; stamens numerous; endosperm homogeneous
or ruminate. SE Asia, Malesia, Australia, and Vanuatu
75. Caryota
- Leaf simply pinnate; inflorescences rarely bisexual, usually
unisexual; stamens 3 to numerous; endosperm homo-
geneous 2
2. Basipetally hapaxanthic palms; inflorescences always uni-
sexual; sepals of staminate flower connate in a tube; sta-
mens (3)6(9-15). India, China, Indochina southwards to S
Thailand 76. Wallichia
- Basipetally hapaxanthic or acropetal pleonanthic palms; in-
florescences sometimes bisexual; sepals of staminate flower
free, imbricate; stamens (6)-numerous. China, Ryukyu, SE
Asia, Malesia, Christmas Is. (Indian Ocean), Australia
74. Arenga
74. Arenga Labill.
Arenga Labill. in DC., Bull. Sc. Soc. Phil. 2: 162 (1800), nom.
cons.
Dwarf to large, solitary or clustered, acropetally
pleonanthic or basipetally hapaxanthic, monoe-
cious or dioecious. Leaves pinnate, leaflets usually
discolorous. Inflorescences bisexual, or unisexual,
sometimes multiple, spicate or branched to 1-2
orders. Staminate flowers with 3 imbricate,
distinct, or briefly connate sepals; corolla val-
vate; stamens 6, 9-numerous, anthers elongate,
latrorse; pollen sulcate, exine intectate spiny. Pis-
tillate flowers globose; sepals 3, distinct, imbricate;
petals 3, connate basally, valvate; staminodes 3 or
absent. Fruit 1-3-seeded. Endosperm homoge-
neous; embryo lateral. Eophyll flabellate or bifid.
About 17 spp., from India, South China, Ryukyus
and Taiwan, through SE Asia, Malesia including
Christmas Is. (Indian Ocean) to north Australia,
the greatest diversity occurring on the Sunda
Shelf.
75. Caryota L. Fig. 81T-V
Caryota L., Sp. Pl.: 1189 (1753); W.H. Hahn, Ph.D. Thesis,
Univ. of Wisconsin, Madison (1993).
Schunda-Pana Adanson.
Solitary or caespitose, basipetally hapaxanthic,
monoecious. Leaves bipinnate; leaflets wedge-
shaped, concolorous. Inflorescences bisexual,
solitary, branched to 1(2) orders, or spicate; triads
protandrous, spiral. Staminate flowers symmetri-
cal; sepals 3, distinct, imbricate; petals 3, valvate;
stamens 6-ca.lOO, anthers linear, latrorse; pollen
sulcate, exine intectate, spiny. Pistillate flower
with sepals 3, imbricate, basally connate; petals 3,
connate; staminodes 0-6. Fruit 1-2-seeded. Seeds
globose or hemispherical, endosperm homoge-
neous or ruminate; embryo lateral. Eophyll bifid
with rhomboid segments. About 12 spp., from
Sri Lanka, India, southern China, southwards
through SE Asia, Malesia to northern Australia,
Solomon Islands, and Vanuatu.
76. Wallichia Roxb.
Wallichia Roxb., Pl. Coromandel 3: 91 (1820).
Dwarf to large, solitary or clustered, hapaxanthic,
monoecious or ?dioecious. Leaves spiral or disti-
chous, pinnate, leaflets discolorous. Inflores-
cences solitary, unisexual, dimorphic, pistillate
distal, with inconspicuous bracts, staminate
proximal with conspicuous bracts. Staminate
flowers with tubular 3-lobed calyx; corolla exceed-
ing calyx, tubular basally; stamens 3-15, anthers
linear; pollen sulcate, exine intectate, spiny. Pistil-
late flowers solitary, sepals 3, imbricate, distinct;
petals 3 united basally; staminodes 0-3. Fruit 1-2,
rarely 3-seeded. Endosperm homogeneous; em-
bryo lateral. Eophyll ovate to elliptic. Seven spp.,
from the Indian Himalayas and upper Burma to
China and southwards to peninsular Thailand.
2. Tribe Iriarteeae Drude (1881).
Unarmed (Wettinia with spinelike trichomes),
pleonanthic, monoecious tree palms with stilt
roots. Leaves reduplicately pinnate, sheaths usu-
ally forming a crownshaft; leaflets praemorse,
often secondarily divided between ribs to give a
plumose appearance; inflorescences infrafoliar,
bisexual or unisexual, spicate or branched to 1
order; peduncle with prophyll and up to 9 pedun-
cular bracts completely sheathing at first, then
splitting apically to allow elongation of the pe-
duncle, all shed at anthesis; flowers superficial;
pistillode minute or absent; gynoecium tricar-
pellate, 1-3-ovulate; fruit 1-seeded or if more than
1-seeded, then fruit lobed; endosperm homoge-
neous; germination adjacent-ligular.
The tribe is characterized by the almost constant
presence of stilt roots. The leaflets are praemorse
and often longitudinally divided and displayed
in many planes. There is often a wide variety of
indumentum on leaves and bracts. In Genera
Palmarum two subtribes were recognized,
Iriarteinae and Wettiniinae. Socratea, included
in Iriarteinae, was almost certainly misplaced
(Henderson 1990). In this account, no subtribes
are recognized.
 Palmae
KEY TO THE GENERA OF THE IRIARTEEAE
1. Staminate and pistillate flowers borne on the same inflores-
cence in triads, or toward the tips of the rachillae the stami-
nate paired or solitary; inflorescences always solitary at a
node
- Staminate and pistillate flowers borne on separate inflores-
cences (pistillate flowers sometimes accompanied by abor-
tive lateral staminate flowers), these usually several at a
node, the central pistillate, the lateral staminate. Colombia,
Venezuela, Ecuador, Peru 81. Wettinia
2. Stigmatic residue apical or subapical in fruit; staminate
flowers with 9-100 or more stamens
- Stigmatic residue near or at the base in fruit; staminate
flowers with 6 stamens
3. Inflorescence terete and decurved in bud, the numerous {10
or more) bracts falling from the terete peduncle as the inflo-
rescence expands, the rachillae long, slender, laxly pendu-
lous; staminate flowers symmetrical, closed in bud, with
imbricate sepals and 9-20 stamens; seed with lateral em-
bryo; stilt roots mostly slender, sparsely prickly, forming a
dense cone obscuring the central stem. Costa Rica to Bolivia
79. Iriartea
- Inflorescences somewhat dorsiventrally compressed and
erect in bud, the 4-7 bracts splitting abaxially, subpersistent
and erect on the dorsiventrally compressed peduncle which
becomes reflexed at anthesis; rachillae short, stout, rather
stiffly spreading at anthesis, pendulous in fruit; staminate
flowers asymmetrical, or at least angled by close packing,
open in bud, sepals usually united basally at least briefly,
stamens 20-100 or more; seed with apical or eccentrically
apical embryo; stilt roots stout, usually densely prickly
forming an open supporting cone. Costa Rica to Bolivia
80. Socratea
4. Slender undergrowth palmlets of lowland forest. Leaflets
borne in one plane, undivided; inflorescence interfoliar, or
becoming infrafoliar in fruit, erect, branching to 1 order
only, long pedunculate, the bracts persistent; fruit small,
ellipsoid or obovoid; seed with apical or eccentrically apical
embryo. Guyana, Amazonia 78. Iriartella
- Robust canopy palms of upland or montane forest; leaflets,
at least in midleaf, divided into several segments displayed
in different planes; inflorescences infrafoliar erect or de-
curved in bud, branching to 2 orders; the bracts deciduous;
fruit mostly globose; seed with basal embryo. Panama and
northern S America, mostly Andean 77. Dictyocaryum
77. Dictyocaryum H. Wendl.
Dictyocaryum H. Wendl., Bonplandia 8: 106 {1860);
Henderson, Fl. Neotrop. 53 {1990).
Solitary, rarely clustered, discolorous. Leaves
plumose. Inflorescences solitary, branched to 2
orders, protandrous. Staminate flowers symme-
trical; sepals 3, distinct, imbricate; petals 3, ex-
ceeding sepals, basally connate, valvate; stamens
6, anthers elongate, basifixed, latrorse; pollen el-
liptic, sulcate, exine tectate. Pistillate flowers
smaller than the staminate; sepals 3, distinct, im-
bricate; petals exceeding sepals, imbricate;
staminodes 6, minute. Fruit 1-seeded, stigmatic
remains basal; epicarp smooth. Seed spherical;
353
embryo basal. Eophyll bifid. Three spp., north-
western S America.
2 78. Iriartella H. Wendl. Fig. 82F
Iriartella H. Wendl., Bonplandia 8: 103, 106 {1860);
Henderson, Fl. Neotrop. 53 {1990).
Caespitose, slender. Inflorescences solitary, bi-
sexual, branching to 1 order, protandrous. Stami-
3 nate flowers symmetrical; sepals 3, distinct,
4
imbricate or basally connate; petals 3, distinct,
valvate; stamens 6, anthers oblong, basifixed,
latrorse; pollen elliptic, sulcate, pontoperculate,
exine tectate. Pistillate flowers smaller than stami-
nate; sepals 3, distinct, imbricate or basally con-
nate; petals 3, imbricate basally; staminodes 6.
Fruit 1-seeded, stigmatic remains basal; epicarp
smooth. Embryo apical. Eophyll shallowly bifid.
Two spp., Guyana, Amazonia.
79. Iriartea Ruiz & Pavon
Iriartea Ruiz & Pavon, Fl. Peruv. Chil. Prodr.: 149 {1794);
Henderson, Fl. Neotrop. 53 {1990).
Solitary, robust. Stem often ventricose. Inflores-
cences solitary, infrafoliar, branching to 2 orders,
protandrous. Staminate flowers symmetrical;
sepals 3, distinct, imbricate; petals 3, exceeding
sepals, valvate; stamens 9-20, anthers elongate,
latrorse; pollen sulcate, exine intectate. Pistillate
flowers smaller than staminate; sepals 3, distinct,
imbricate; petals 3, distinct, imbricate except for
triangular valvate tips; staminodes to 12, tooth-
like. Fruit 1-seeded, stigmatic remains apical; epi-
carp smooth. Embryo lateral. Eophyll undivided.
Only one sp., Iriartea deltoidea R. & P., wide-
spread in Central and northern S America.
80. Socratea Karsten Fig. 81R,S, 88M-S
Socratea Karsten, Linnaea 28: 263 {1857) ("1856"); Henderson,
Fl. Neotrop. 53 (1990).
Solitary. Inflorescences solitary, branching to 1
order, protogynous. Staminate flowers open con-
genitally in bud; sepals 3, basally united; petals 3,
exceeding sepals, distinct, asymmetrical, imbri-
cate basally; stamens 20-100+, anthers linear,
basifixed, latrorse; pollen elliptic, sulcate, exine
intectate, spinose. Pistillate flowers symmetrical,
smaller than staminate; sepals 3, imbricate; petals
3, distinct, imbricate, tips valvate; staminodes
6, minute, toothlike. Fruit 1-seeded, stigmatic
354
remains eccentrically apical; epicarp minutely
roughened. Embryo eccentrically apical. Eophyll
bifid. Five spp., Central and northern S America.
81. Wettinia Poepp. 88V,W.
Wettinia Poepp. in End!., Gen. Pl.: 243 (1837); Bernal, Caldasia
17: 367-378 (1995).
Catoblastus H. Wend!. (1860).
Solitary or clustered, moderate to robust. Inflores-
cences unisexual, infrafoliar, 3-8( -15) per node,
or single, spicate or branched to 1 order. Stami-
nate flowers congenitally open; stamens 6-19, an-
thers basifixed, elongate, latrorse; pollen elliptic,
sulcate, exine intectate, spinose. Pistillate flowers:
sepals 3, imbricate, connate basally; petals 3, simi-
lar to sepals; gynoecium of 1(-2) fertile carpels
and (1- )2 abortive carpels, style basal, stylodia 3,
large, elongate. Fruits 1-2-seeded, stigmatic re-
mains basal; epicarp hairy, warty, or prickly. Em-
bryo basal. Eophyll undivided. Twenty-one spp.,
Panama, northwestern S America, and the western
part of the Amazon basin.
3. Tribe Podococceae J. Dransf. & N. Uhl (1986).
Unarmed, pleonanthic, monoecious; leaves pin-
nate, reduplicate, leaflets praemorse; inflores-
cence solitary, bisexual, rachis bracts spirally
arranged, forming lower margins of hair-filled pits
enclosing triads of flowers basally, paired or soli-
tary staminate flowers distally; flowers exserted
from pits by elongation of floral receptacle be-
tween calyx and corolla; pistillode short, briefly
3-lobed; gynoecium trilocular, triovulate, ovules
pendulous; seeds fusiform; endosperm homoge-
neous; germination adjacent-ligular.
The tribe contains the single monotypic genus
Podococcus. Its relationships are possibly with the
Iriarteeae, but this has yet to be confirmed by rig-
orous analysis. The floral pits, flowers exserted on
stalks, and curious horizontally displayed stalked
fruits are unusual features.
82. Podococcus G. Mann & H. Wendl.
Fig. 88X-B'
Podococcus G. Mann & H. Wend!., Trans. Linn. Soc. London
24: 426 (1864); Uhl & Dransf., Gen. Palm.: 329 (1987).
Small, spreading by stolons, prop roots present.
Leaflets rhomboid, single-fold, discolorous. Inflo-
rescences interfoliar, spicate, protandrous; pe-
duncular bracts 2-3. Staminate sepals 3, distinct,
Palmae
imbricate; petals 3, valvate; stamens 6, filaments
short, anthers short, dorsifixed, latrorse; pollen
sulcate, exine tectate reticulate. Pistillate flower
symmetrical; sepals 3, distinct, imbricate; petals
basally connate; staminodes 6, minute; stigmas 3,
recurved. Fruit stalked, stigmatic remains lateral;
epicarp smooth, mesocarp fleshy, endocarp crus-
taceous. Embryo lateral. Eophyll rhomboid,
praemorse. Only one sp., P. barteri G. Mann & H.
Wendl., equatorial western Africa, ranging from
the Niger delta almost to the Congo River.
4. Tribe Areceae Drude (1881).
Pleonanthic reduplicately pinnate or pinnately
ribbed. Inflorescence with a prophyll and a
single (very rarely 2) peduncular bracts; gyno-
ecium trilocular, triovulate or, more commonly,
pseudomonomerous; fruit with apical, lateral, or
basal stigmatic remains; endocarp without 3 well-
defined pores.
The relationships within this large tribe remain
poorly understood and much in need of a modern
cladistic analysis using both morphological and
molecular datasets.
KEY TO THE SUBTRIBES OF ARECEAE
1. Gynoecium triovulate 2
- Gynoecium pseudomonomerous, rarely two abortive
carpels present 5
2. Inflorescence at anthesis entirely enclosed by a netlike
prophyll and peduncular bract; fruit corky-warted; leaf
very large, undivided, or irregularly divided
4b. Manicariinae
- Inflorescence not enclosed by prophyll and peduncular
bract at anthesis; fruit smooth; leaf pinnate or, if entire,
rather small 3
3. Prophyll very much smaller than peduncular bract(s), usu-
ally obscured by the leaf sheaths; peduncular bracts large,
woody, usually beaked 4a. Oraniinae
- Prophyll and peduncular bract similar; peduncular bract
membranous or coriaceous but scarcely woody, not
conspicuously beaked 4
4. Inflorescence with numerous spreading branches of up to
the 4th order; staminate flowers rounded with 6 stamens;
fruit lenticular or ovoid, stigmatic remains basal; moder-
ate palms with stems obscured by sheath fibers
4c. Leopoldiniinae
- Inflorescence spicate, or with few more or less approxi-
mate branches of 1, rarely 2 orders; staminate flowers
pointed with 8-40 stamens; fruit ovoid to ellipsoidal, stig-
matic remains apical; diminutive to moderate palms with
slender stems not obscured by sheath fibers
4d. Malortieinae
5. Endocarp with well-defined operculum covering the
embryo 6
- Endocarp lacking well-defined operculum 7
 Palmae
6. Spines present, at least in juvenile stages
4o. Oncospermatinae
- Spines absent 4n. Iguanurinae
7. Massive litter-trapping palms without crownshafts, with
bi- or unisexual inflorescences; stigmatic remains apical;
endosperm homogeneous 4q. Masoalinae
- !flitter-trapping, then not massive; crownshafts present or
not; stigmatic remains various; endosperm homogeneous
or ruminate 8 zigzag, not irregularly angled; flowers borne spirally,
8. Staminate flowers with over 50 stamens; fruit corky-
warted; embryo apical 4f. Lemurophoenicinae
- Staminate flower if multistaminate, then fruit not corky-
warted, embryo basal 9 not, then only 1 bract in the inflorescence
9. Petals of pistillate flower connate; staminodes connate in a
cupule adnate to the petals 4h. Roystoneinae
- Petals of pistillate flower free, imbricate, very rarely con-
nate (Siphokentia); staminodes toothlike, not forming a
cupule 10
10. Inflorescence spicate, congested, bearing 1large complete
and 2-several small, incomplete peduncular bracts; stami-
nate sepals narrow, separated in bud; leaf large, bifid or
irregularly divided
4p. Sclerospermatinae
- Inflorescence usually not bearing incomplete peduncular
bracts, variously branched, congested or not; staminate
sepals imbricate, valvate, or connate, not widely separate;
leaves various 11
11. Flowers borne in pits or depressions in long pedunculate,
interfoliar spikes, spikes solitary or several together in
a leaf axil; staminate flowers symmetrical or nearly so;
sepals rounded, broadly imbricate; stigmatic remains
apical 4k. Linospadicinae
- Flowers superficial or, if in pits, infloresence branched,
infrafoliar; flowers symmetrical or asymmetrical; sepals
various; stigmatic remains various 12
12. Flowers borne in pits with prominent, rounded lips; inflo-
rescence infrafoliar, branched to 3 orders; peduncle very
short; basal branches sharply divaricate; crownshaft well
developed; staminate flowers symmetrical, rounded in
bud; sepals broad, rounded, imbricate; stamens 6-15; stig-
matic remains apical 4j. Cyrtostachydinae
- Flowers not borne in pits or, if so, then staminate flower
with acute sepals and/or fruit with lateral stigmatic re-
mains and the inflorescence branching to 1 order only 13
13. Staminate flowers strictly symmetrical, rounded, or bullet-
shaped 14
- Staminate flowers usually asymmetrical, if symmetrical,
then inflorescence bracts 1 only, or staminate flower not
rounded or bullet-shaped 15
14. Inflorescences usually interfoliar, spicate or branched up
to 4 orders; staminate flowers often very small, more or
less rounded apically in bud with sepals nearly half as long
as the petals in bud; stamens 3-6; gynoecium often
trilocular but uniovulate; stigmatic remains basal; leaflet
tips almost always entire 4e. Dypsidinae
- Inflorescence infrafoliar, branched to 3 orders; staminate
flowers moderate, bullet-shaped in bud with sepals less
than half as long as the petals; stamens numerous; gyno-
ecium unilocular; stigmatic remains apical; leaflet tips
praemorse 41. Ptychospermatinae
15. Inflorescence branched to 1 order, sometimes hip-
puriform, rarely spicate, peduncular bract slightly to
markedly exserted from the prophyll, if not exserted, then
stigmatic remains lateral, staminate flower usually with
filaments markedly inflexed; stigmatic remains apical,
basal, or lateral. New World 4g. Euterpeinae
355
- Inflorescence spicate or branched to 3 orders, peduncular
bract, if present, not exserted from the prophyll; staminate
flowers with filaments not markedly inflexed; stigmatic
remains always apical. Old World 16
16. Inflorescence branches bearing a prophyll only or, if also
bearing a peduncular bract, triads usually borne along the
entire length of the rachilla (except Loxococcus ), branches
usually not widely spreading, rachillae straight, scarcely
distichously, in verticils or on one side of the rachilla only;
staminate flowers usually more or less 3 times as long as
the pistillate or more when inflorescence bract( s) open, if
4m. Arecinae
- Inflorescence always bearing a prophyll and a pedancular
bract, branches spreading widely, triads borne almost to
the ends of the rachillae, rachillae usually distinctly zigzag,
irregularly angled, flowers spirally inserted; staminate
flowers only slightly larger than the pistillate when bracts
open 4i. Archontophoenicinae
4a. Subtribe Oraniinae J. Dransf. & N. Uhl
(1986).
Unarmed, pleonanthic, monoecious, reduplicately
pinnate, crownshaft not present, apices prae-
morse; inflorescences axillary, interfoliar, solitary,
bisexual, prophyll short, tubular, 2-keeled; pedun-
cular bract 1(2), conspicuous, almost woody, tu-
bular, completely enclosing the inflorescence in
bud, then splitting and falling; flowers superficial,
staminate and pistillate superficially similar;
pistillode absent or minute; gynoecium trilocular,
triovulate; fruit developing from 1, 2, or rarely 3
carpels; endocarp with a basal swelling; seed ba-
sally attached; endosperm homogeneous; germi-
nation remote-tubular.
A monotypic subtribe of triovulate, tricarpellate
palms. Its relationships are as yet obscure.
83. Orania Zipp.
Orania Zipp. in Bl., Algemene Konst- en Letter-bode 1: 297
(1829); Essig, Lyonia 1: 211-233 (1980).
Halmoorea J. Dransf. & N. Uhl (1984).
Solitary. Leaves distichous or spiral; leaflets
single-fold, discolorous. Inflorescences branching
to 1-3 orders, protandrous; triads subdistichous
or spiral. Staminate flowers asymmetrical; sepals
3, distinct or united; petals 3, distinct, valvate,
stamens 3-30, anthers linear-oblong, basifixed,
latrorse; pollen sulcate, exine tectate. Pistillate
flowers with sepals and petals similar to stami-
nate; staminodes 3-6; epicarp smooth. Embryo
subapical or lateral. Eophyll bifid or pinnate, with
praemorse apices. About 17 spp., south Thailand
to New Guinea, and three spp., Madagascar.
356 Palmae

4b. Subtribe Manicariinae J. Dransf. & N. Uhl circumscissile basally; peduncular bract 1, like
(1986). the prophyll, also early caducous; fruit 1-seeded,
stigmatic remains basal; seed attached
Unarmed, pleonanthic, monoecious, reduplicately laterally, hilum vertical; endosperm homoge-
pinnate; crownshaft not present; inflorescen- neous; embryo subbasal; germination adjacent-
ces solitary, interfoliar, bisexual, protandrous, ligular.
branched to 1-4 orders; prophyll tubular, beaked, The subtribe contains the genus Leopoldinia,
enclosing inflorescence, netlike; peduncular bract peculiar in several respects. Sheaths are variously
like the prophyll; flowers partially sunken in fibrous; inflorescences may tend to be unisexual
rachillae; pistillode lacking; gynoecium trilocular, or have unisexual rachillae, the gynoecium is pos-
triovulate, ovules lateral, anatropous; fruit large, sibly pseudomonmerous, but, if so, has relatively
rounded, 1-3-lobed, 1-3-seeded, stigmatic re- well-developed sterile carpels and the fruit is
mains subbasal; epicarp obsolescent at maturity, extraordinarily fibrous.
mesocarp developing into corky warts, endocarp
thin, smooth, with a basal buttonlike swelling;
seed basally attached; endosperm homogeneous, 85. Leopoldinia Mart.
hollow; embryo basal; germination adjacent- Leopoldinia Mart., Hist. Nat. Palm. 2: 58 (1824); Kubitzki,
ligular. Nord. J. Bot. 11: 429-432.
This subtribe includes the single genus
Manicaria. The gynoecium is trilocular and Solitary or clustered. Sheath with conspicuous fi-
triovulate, and the endocarp has a basal heart- bers, leaflets single-fold, acuminate, concolorous
shaped swelling also present in Orania, or discolorous. Inflorescences interfoliar, solitary,
Lemurophoenix, Sclerosperma, and the Phyteleph- branching to 4 orders, sometimes unisexual.
antoideae. The netlike inflorescence bracts that Staminate flowers very small; sepals 3, distinct,
enclose the inflorescence are highly distinctive. imbricate; petals 3, distinct, valvate; stamens 6,
Fruit is corky-warted. anthers rounded, latrorse; pollen elliptic, sulcate,
exine tectate; pistillode barrel-shaped. Pistillate
flowers larger than the staminate; otherwise simi-
84. Manicaria J. Gaertn. lar; staminodes 6. Fruit ovoid to disciform, epicarp
Manicaria J. Gaertn., Fruct. 2(3): 468 (1791); Henderson, smooth, mesocarp fibrous, endocarp thin. Seed
Palms of Amazon: 118 (1994). rounded or lenticular. Eophyll bifid, the segments
very slender. Three spp., confined to NW Brazil,
Solitary or caespitose. Leaves undivided or Amazonian Colombia, and southern Venezuela.
variously divided, sometimes with separated
leaflets with pointed tips, concolorous. Rachillae
crowded, bracts stiff, pointed, subtending sunken 4d. Subtribe Malortieinae J.D. Hook. (1883).
triads. Staminate flowers slightly asymmetrical;
sepals 3, united basally; petals 3, exceeding sepals, Unarmed, pleonanthic, monoecious, reduplicately
united basally; stamens 30-35, anthers elongate, pinnate, crownshaft absent, inflorescences soli-
dorsifixed, introrse; pollen sulcate, exine tectate. tary, interfoliar; prophyll tubular, membranous,
Pistillate flowers with ca. 15 staminodes, styles 2-keeled, included within leaf sheath; peduncular
conical. Eophyll bifid. Only one sp., M. saccifera J. bract 1, elongate, papery, disintegrating, rachilla
Gaertn., Central America to the lower Amazon bracts spiral to distichous, triads borne in shallow
River. depressions; pistillode lacking; gynoecium trilo-
culate, triovulate; fruit 1-seeded, stigmatic re-
mains apical; seed basally or laterally attached,
4c. Subtribe Leopoldiniinae J. Dransf. & N. Uhl endosperm homogeneous or ruminate; embryo
(1986). basal germination adjacent-ligular.
The single genus displays several unusual fea-
Unarmed, pleonanthic, monoecious, reduplicately tures and is morphologically quite diverse.
pinnate, crownshaft not present; inflorescence Sheaths are always fibrous, the leaves are some-
interfoliar, usually bisexual; prophyll borne times windowed (with incomplete split at the base
considerably above the base, tubular, 2-winged, of the folds at the leaf tip). There is a well-
membranous, splitting early in development, developed staminodal ring.
 Palmae 357

86. Reinhardtia Liebm. Very small to very large, acaulescent to erect,

rarely climbing, solitary or caespitose. Crownshaft
Reinhardtia Liebm. in Mart., Hist. Nat. Palm. 3: 311 (1849);
Gentes Herb. 8: 541-576 (1957). present or not; sheath sometimes disintegrating
into fibers; leaflets single or multifold. Inflores-
Solitary or caespitose. Leaves undivided or pin- cences protandrous; rachillae with superficial
nate, sometimes "windowed," leaflets with bifid triads proximally, staminate flowers distally.
tips, concolorous. Inflorescences protandrous, Stamens 3 (antesepalous or antepetalous) or 6,
spicate or branching to 2 orders. Staminate flow- where three, 3 staminodes sometimes present, fila-
ers with sepals 3, distinct, imbricate; petals 3, ments free or united, erect or inflexed, anthers
valvate, connate basally; stamens 8-40, anthers linear or didymous, rarely with divergent thecae;
basifixed, elongate, latrorse; pollen sulcate, exine pistillode present or absent. One hundred and
tectate. Pistillate flowers with sepals and petals forty spp., Madagascar, Comoro Islands, and
similar to staminate; staminodal ring with 6-15 Pemba Is. off the coast of Tanzania.
teeth; ovary with robust style and 3 stigmas. Epi-
carp smooth, mesocarp fleshy, endocarp fragile.
Eophyll simple or bifid. Six spp., Mexico to 4f. Subtribe Lemurophoenicinae J. Dransf.
Panama, one sp. northwest Colombia, another {1995).
Dominican Republic.
Staminate flowers symmetrical; sepals 3, imbri-
cate; petals 3, valvate, reflexed at anthesis; stamens
52-59, filaments irregular, connate or distinct, an-
4e. Subtribe Dypsidinae Becc. {1914). thers linear, medifixed, latrorse. Pistillate flower
with sepals 3, imbricate; petals 3, imbricate;
Unarmed, sometimes dichotomously branched, staminodes 10-12; gynoecium pseudomono-
monoecious, pleonanthic; leaves pinnate or pin- merous. Fruit warted, with basal stigmatic re-
nately ribbed, reduplicate; leaflets entire, rarely mains.
praemorse; inflorescences usually interfoliar, The subtribe includes a single monotypic genus
sometimes infrafoliar, bisexual, spicate or from northeastern Madagascar. The multis-
branched to 1-4 orders; peduncle :±: elongate, taminate flowers and corky-warted fruits are
basal branches not divaricate; peduncular bract unique among Madagascar arecoid palms. The
usually conspicuous, exserted and caducous, large fruit has well-developed endocarp with a
prophyll often adnate to peduncle; staminate basal heart-shaped button, reminiscent of that of
flowers symmetrical; petals of pistillate flowers Orania and Manicaria, while the endosperm is
imbricate; gynoecium sometimes trilocular but shallowly ruminate and the embryo apical.
uniovulate; fruit with basal stigmatic remains and
smooth epicarp; germination adjacent-ligular;
eophyll bifid. 88. Lemurophoenix J. Dransf.
The subtribe contains the single genus Dypsis Lemurophoenix J. Dransf, Kew Bull. 46: 61 (1991).
with over 140 spp., confined to Madagascar,
Comoro Islands, and Pemba. Many genera have Massive, solitary tree. Crownshaft present; leaflets
been recognized in the past, but recently collected single-fold, acuminate. Inflorescences infrafoliar,
material has clouded the generic boundaries. This branching to 3 orders, protandrous; prophyll
is one of the largest and most diverse of palm and peduncular bract similar, inserted close
genera. together, caducous; rachillae bearing spiral
triads basally, staminate only distally. Pollen
sulcate, exine tectate, perforate; pistillode colum-
87. Dypsis Noronha ex Mart. nar. Epicarp split into low corky warts; mesocarp
Dypsis Noronha ex Mart., Hist. Nat. Palm. 3: 180 (1837); corky-spongy, easily separating from endocarp;
Dransfield & Beentje, Palms Madag.: 122-413 (1995). endocarp smooth, with a basal heart-shaped
Chrysalidocarpus H. Wendl. (1878). button; seed basally attached; endosperm
Phloga Noronha ex J.D. Hook. (1883). shallowly ruminate; embryo apical. Germination
Neophloga Baillon (1894).
Neodypsis Baillon (1894). adjacent-ligular; eophyll bifid. Only one sp., L.
Vonitra Becc. (1906). halleuxii J. Dransf., restricted to northeastern
Antongilia Jum. (1928). Madagascar.
358 Palmae

4g. Subtribe Euterpeinae J. Dransf. & N.W. Uhl - Crownshaft usually not clearly defined except in a few spe-
(1986). cies, but then leaf sheaths tubular but partly open; petiole
usually long; inflorescence branches glabrous to lightly
hairy or scaly, stiffly ascending or divaricate in all direc-
Small to large, unarmed; crownshafts present tions, often markedly bulbous at the base; prophyll and
or not well differentiated; leaves pinnate or pin- peduncular bract markedly unequal, the peduncular bract
nately ribbed; leaflets entire, regularly or irregu- longer, terete, and beaked, usually inserted some distance
larly arranged; inflorescences infrafoliar, rarely above the prophyll; flowers superficial 90. Prestoea
interfoliar, branched to 1 order or rarely bifurcate
or spicate; prophyll not basally adnate to the pe- 89. Euterpe Mart.
duncle; peduncular bracts slightly to markedly
exserted from the prophyll before anthesis; stami- Euterpe Mart., Hist. Nat. Palm. 2: 28.1823; emended 3:165
(1837); 3: 230 (ed. 2) (1845), nom. cons.; Henderson &
nate flowers usually asymmetrical, somewhat Galeano, Fl. Neotrop. 72: 1-89 (1996).
acute; filaments usually inflexed; petals of pistil-
late flowers imbricate; gynoecium pseudomono- Tall, solitary or clustered. Crownshaft usually
merous; stigmatic remains various. prominent. Inflorescences branched to 1 order;
This subtribe of six genera includes all the peduncle dorsiventrally compressed; peduncular
pseudomonomerous members of the Areceae in bract and prophyll equal or subequal; rachillae
the New World except Roystonea. They inhabit pendulous, hairy; staminate flowers paired or
lowland or montane rainforest from the West single distally; staminate petals 3, valvate, stamens
Indies south through Central America to Peru, 6, pollen elliptic, sulcate or sometimes tricho-
Paraguay, and Argentina. The genera share pin- tomosulcate, exine tectate, finely reticulate or
nate or pinnately veined leaves, inflorescences foveolate; pistillode 3-lobed. Ovule hemiana-
branched to 1 order or rarely spicate, a unilocular tropous. Stigmatic remains lateral to subapical.
gynoecium, and rather thin fruit walls. The lack of Endosperm ruminate or homogeneous; embryo
a crownshaft (as in some of these taxa), superficial subbasal. Germination adjacent-ligular; eophyll
triads of flowers present throughout the rachillae, bifid or pinnate. Seven spp. Central through tropi-
the simple fruit wall, and chromosome numbers cal S America extending into Trinidad and the
of n = 18 are characters that appear to be Lesser Antilles.
unspecialized in the pseudomonomerous mem-
bers of the Areceae.
90. Prestoea J.D. Hook.
Prestoea J.D. Hook. in Benth. & Hook., Gen. Pl. 3: 875, 899
KEY TO THE GENERA OF THE EUTERPEINAE
(1883), nom. cons.; Henderson & Galeano, Fl. Neotrop. 72:
1. Inflorescences hippuriform (resembling a horse's tail); 1-89 (1996).
adaxial surface of the rachis lacking branches, rachillae all
lateral or abaxial, curved and pendulous; prophyll much Small to moderate, clustered, rarely solitary.
shorter than the peduncular bract, the latter terete, beaked; Crownshaft lacking or partial. Inflorescences
embryo large, about 2/3 as long as the seed 92. Oenocarpus
- Inflorescences not hippuriform; the rachis bearing spirally
branched to 1 order, rarely spicate; peduncle elon-
inserted branches on all sides, or spicate; prophyll and pe- gate, usually terete; prophyll shorter than the pe-
duncular bracts nearly equal or unequal; embryo small, duncular bract; rachillae white, red in fruit, lightly
much less than 1/2 as long as the seed 2 tomentose; staminate flowers paired or single
2. Inner stamens markedly adnate to the pistillode in stami- distally. Staminate petals 3, valvate, stamens 6,
nate flowers; sepals of pistillate flowers connate basally;
stigmatic remains basal in fruit 93. Hyospathe
pollen elliptic, sulcate, exine tectate, finely
- Stamens not adnate to the pistillode in staminate flowers; reticulate or foveolate; pistillode 3-lobed. Stig-
sepals of pistillate flowers imbricate; stigmatic remains matic remains subapical. Endosperm ruminate
apical or lateral 3 or rarely homogeneous; embryo subbasal.
3. Acaulescent or stem very short; inflorescence spicate; Germination adjacent-ligular; eophyll bifid or
stigmatic remains apical 91. Neonicholsonia
- Stems short, moderate, or tall; inflorescence branched to pinnate. Ten spp., Central America, the Antilles,
1 order; stigmatic remains lateral 4 and the Andean and Guayana highlands in S
4. Crownshaft clearly defined; petiole short; inflorescence America.
branches densely white to dark brown-tomentose, spread-
ing, pendulous from a somewhat dorsiventrally flattened
rachis, usually not markedly enlarged at the base; prophyll 91. Neonicholsonia Damm.
and peduncular bract about equal or unequal, peduncular
bract beaked, inserted close to prophyll; flowers often borne Neonicholsonia Damm., Gard. Chron. III, 30: 178 (1901);
in pits 89. Euterpe Henderson & Galeano, Fl. Neotrop. 72: 1-89 (1996).
 Palmae
Small, solitary, acaulescent or short. Crownshaft
lacking. Inflorescences spicate; peduncle elongate,
usually terete; peduncular bract much longer than
the prophyll; staminate flowers paired or single
distally. Staminate petals 3, valvate, stamens 6,
pollen elliptic, sulcate or rarely trichotomosulcate,
exine tectate, coarsely reticulate; pistillode trifid.
Stigmatic remains apical to subapical. Embryo
basal to subbasal, endosperm ruminate peripher-
ally; embryo subbasal. Germination adjacent-
ligular; eophyll bifid, tips divided. Only one
variable sp., N. watsonii Damm., Panama and
Nicaragua.
92. Oenocarpus Mart.
Oenocarpus Mart., Hist. Nat. Palm. 2: 21 (1823); Balick, Adv.
Econ. Bot. 3: 1-140 (1986); Henderson et a!., Field Guide:
129 (1995).
]essenia Karsten (1857).
Solitary or clustered. Crownshaft partial. Inflores-
cences becoming infrafoliar, branched to 1 order
laterally and abaxially (hippuriform), adaxial
branches absent, rarely bifurcate or spicate; pe-
duncle flattened; peduncular bract longer than
prophyll; staminate flowers paired or single dis-
tally. Staminate petals 3, valvate, stamens 6(7-20);
pollen elliptic, sulcate, exine tectate, scabrate or
reticulate; pistillode bifid or trifid. Stigmatic re-
mains apical to eccentric. Endosperm homo-
geneous; embryo very large. Germination
adjacent-ligular; eophyll bifid. Nine spp., from
Costa Rica and Panama to the Amazon and
Orinoco Valleys in S America.
93. Hyospathe Mart.
Hyospathe Mart., Hist. Nat. Palm. 2:1 (1823); Skov & Balslev,
Nord. J. Bot. 9: 189-202 (1989).
Small, rarely moderate, solitary or clustered.
Crownshaft short to long. Leaves pinnate, bifid,
bi- or trijugate, concolorous or discolorous. Inflo-
rescences branched to 1 order; prophyll shorter
than the peduncular bracts; staminate flowers
paired or single distally. Staminate petals 3, ovate,
asymmetrical; stamens 6, antesepalous free or
joined with pistillode, antepetalous longer adnate
to pistillode; pollen elliptic, sulcate, or rarely
trichotomosulcate, exine tectate scabrate; pistil-
lode 2-lobed, ovoid. Stigmatic remains basal.
Endosperm homogeneous, embryo large, basal.
Germination adjacent-ligular; eophyll bifid. Two
spp., one widespread in Central America and
359
northern S America, and one endemic on the east-
ern Andean slopes in Ecuador.
4h. Subtribe Roystoneinae J. Dransf. & N. Uhl
(1986).
Robust unarmed, monoecious palms with bi-
sexual inflorecences; leaves pinnate with entire
leaflets; inflorescences infrafoliar, branched to 4
orders; staminate flowers symmetrical; sepals 3,
distinct, imbricate; petals 3, distinct, valvate;
pistillate flower with petals connate, staminodes
connate in a ring adnate to petals; gynoecium
pseudomonomerous; ovule laterally attached;
fruit with basal stigmatic remains; endosperm ho-
mogeneous, embryo basal; germination adjacent-
ligular; eophyll entire, lanceolate.
The subtribe contains the single genus
Roystonea. It is distinct from other pseudomo-
nomerous subtribes in the connate petals with
valvate distinct lobes, an adnate staminodal ring
in the pistillate flower and the undivided eophyll.
Its relationships are obscure.
94. Roystonea O.F. Cook
Roystonea O.F. Cook, Science II, 12: 479 (1900); Zona, Fl.
Neotrop. 71: 1-35 (1996).
Solitary. Leaflets singlefold, entire. Inflorescences
branched to 3( -4) orders; prophyll tubular, elon-
gate, strongly 2-keeled; peduncular bract 2-3
times as long as prophyll, caducous with prophyll;
rachillae bearing spiral, superficial triads proxi-
mally, staminate flowers distally. Stamens 6-12,
anthers elongate, versatile, dorsifixed, latrorse;
pollen sulcate, exine tectate, foveolate or fossulate;
pistillode subglobose or trifid. Pistillate flowers
with distinct sepals. Epicarp smooth, mesocarp
thin fleshy with thin, flat, anastomosing fibers next
to endocarp, endocarp fragile. Hilum large, circu-
lar, lateral. Ten spp., throughout the islands of the
Caribbean and bordering continental areas.
360 Palmae
4i. Subtribe Archontophoenicinae J. Dransf. &
N. Uhl (1986).
Moderate to robust or tall, solitary, unarmed;
leaves pinnate, leaflets entire; crownshaft always
well developed; inflorescences branched to 1-3
orders; prophyll and peduncular bract similar;
staminate flowers symmetrical or asymmetrical;
filaments erect or inflexed in bud; petals of pistil-
late flower imbricate; gynoecium pseudomo-
nomerous; fruit with apical stigmatic remains;
endocarp lacking an operculum; embryo basal.
Germination adjacent-ligular; eophyll bifid.
A rather homogeneous subtribe of six genera,
one occurring in Australia, one on Lord Howe Is.,
one in New Zealand and neighboring islands, and
three in New Caledonia. These are medium-sized
palms showing less diversity in habit and leaf form
than is seen in the other subtribes of Areceae. Leaf
anatomy of the three New Caledonian genera is
distinctive and is thought to indicate that they
may have originated from a single ancestor (N.W.
Uhl, pers. observ.) This may be an ancient group;
its distribution pattern has been considered com-
mon to rainforest taxa that radiated throughout
Australasia in the Late Cretaceous and Early Ter-
tiary before the development in the Miocene of
extensive archipelagic links with Malesia. Its rela-
tionship to other palms is not clear, and is under
study by ].-C. Pintaud at Toulouse University.
KEY TO THE GENERA OF THE
ARCHONTOPHOENICINAE (J.-C. PINTAUD, PERS.
COMM.)
1. Pinnae with prominent marginal ribs, stigmas large, fleshy,
trifid 2 Hedyscepe H. Wendl. & Drude, Linnaea 39: 178, 203 (1875).
- Pinnae lacking marginal ribs, stigmas small, trifid 3
2. Trunk slender, leaves 3-5, outermost (1st) bracteole large
100. Actinokentia
- Trunk robust, leaves 7-12, outermost bracteole small 4
3. Staminate sepals lanceolate, stamens 6-12, endosperm ho-
mogeneous 5 divaricate, erect or pendulous, protandrous, with
- Staminate sepals short, broadly ovate, stamens usually
more than 12, endosperm ruminate 95. Archontophoenix
4. Fruits small (1.4-2.2cm long), pistillode of various shapes
present 99. Kentiopsis
- Fruits medium to large (2.5-5.5 em long), pistillode most
often absent 96. Chambeyronia
5. Crownshaft waxy, staminate sepals connate basally, fruits
large (Scm long) 97. Hedyscepe
- Crownshaft not waxy, staminate sepals free, fruits small ( 1-
1.5 em) 98. Rhopalostylis
95. Archontophoenix H. Wendl. & Drude
Archontophoenix H. Wendl. & Drude, Linnaea 39: 182, 211
(1875); Dowe & Hodel, Austrobaileya 4: 227-244 (1994).
Stem rather slender, swollen basally. Petiole short.
Rachillae somewhat divaricate, erect or pendu-
lous, protandrous, bearing triads with a few paired
or solitary staminate flowers distally. Staminate
flowers asymmetrical, lateral to pistillate, petals
longer than sepals, valvate; stamens 9-35, fila-
ments curved or deflexed; pollen elliptic, sulcate,
exine tectate, finely reticulate; pistillode cylindri-
cal, equal to stamens, apically lobed. Pistillate
flowers smaller, symmetric. Endosperm ruminate.
Six spp., eastern Australia from the southern coast
of New South Wales to the northern coast of
Queensland.
96. Chambeyronia Vieill.
Chambeyronia Vieill., Bull. Soc. Linn. Normandie II, 6: 229
(1873); Moore & Uhl, Allertonia 3: 313-402 (1984).
Stem enlarged basally. Leaves initially red; petiole
short. Rachillae stout, divaricate, tapering, sinu-
ous; triads distant, a few paired or solitary
staminate flowers distally. Staminate flowers
asymmetrical; stamens 19-55, filaments inflexed;
pollen elliptic, sulcate, rarely trichotomosulcate,
exine tectate, reticulate-foveolate; pistillode lack-
ing. Endosperm homogeneous. Two spp., New
Caledonia.
97. Hedyscepe H. Wendl. & Drude
Stem rather stout, leaf scars whitish. Leaves arch-
ing, stiff; petiole short, stout; inflorescences
branched to 1(-3) order; rachillae stout,
triads nearly throughout; staminate flowers asym-
metrical, stamens 9-10( -12), antesepalous soli-
tary, antepetalous paired, filaments inflexed;
pollen elliptic, sulcate, with finely reticulate,
tectate exine; pistillode with rounded ovarian part
and long, terete style, equalling stamens, En-
dosperm homogeneous. Only one sp., H. canter-
buryana (C. Moore & F. v. Muell.) H. Wendl. &
Drude, endemic to Lord Howe Is.
 Palmae
98. Rhopalostylis H. Wendl. & Drude
Rhopalostylis H. Wend!. & Drude, Linnaea 39: 180,234 (1875);
Bailey, Gentes Herb. 3(8): 410-437 (1935).
Stem green to gray, leaf scars prominent.
Crownshaft short, bulbous. Leaves ascending,
somewhat arched, often twisted; petiole short,
stout. Inflorescences branched to 3 orders;
rachillae stout, divaricate, with or without a short
bare portion; with triads basally, paired to solitary
staminate flower distally. Staminate flowers asym-
metrical, stamens 6, filaments markedly inflexed,
united basally; pollen elliptic, sulcate, with
foveolate to fossulate, tectate exine; pistillode
slender, cylindrical, about equaling stamens. En-
dosperm homogeneous. Variously considered as
three or two spp., one with two varieties, New
Zealand, Chatham Is., Norfolk Is., and Raoul
(Sunday) Is.
99. Kentiopsis Brongn.
Kentiopsis Brongn., Compt. Rend. Hebd. Seances Acad. Sci.
77:398 (1873); Pintaud & Hodel, Principes 42:32-33,41-53
(1998).
Mackeea H.E. Moore (1978).
Very tall. Stems gray, basally thick. Leaves spread-
ing or ascending, often twisted; petiole short,
stout; inflorescences branched to 1-3 orders;
rachillae rather slender, of equal length, with tri-
ads nearly throughout. Staminate flowers asym-
metrical, stamens 32-38, filaments erect tectate,
finely pitted or reticulate; pistillode as long as or
shorter than the stamens. Endosperm homoge-
neous. Four spp., New Caledonia.
100. Actinokentia Damm.
Actinokentia Damm., Bot. Jahrb. Syst. 39: 20 (1906); Moore &
Uhl, Allertonia 3: 313-402 (1984).
Small to moderate. Stems slender, sometimes with
prickly roots. Petiole short or elongate. Inflores-
cences divaricate, branched to 1-3 orders,
protandrous; rachillae rather slender, of equal
length, with triads basally, paired and solitary
flowers distally; bracteoles surrounding the pistil-
late flower sepallike, finely reticulate; pistillode
tapered as long as the stamens. Endosperm homo-
geneous. Two spp., New Caledonia.
361
4j. Subtribe Cyrtostachydinae J. Dransf. &
N. Uhl (1986).
Unarmed, pleonanthic monoecious palms; leaves
pinnate, leaflets entire; crownshaft well developed;
inflorescences infrafoliar, bisexual, branched to 3
orders; peduncle very short, basal branches
divaricate; prophyll and peduncular bract similar,
both caducous; triads borne in pits with rounded
lips; staminate flowers symmetrical, rounded
in bud, petals connate in basal 1/3; filaments
basally connate, inflexed; petals of pistillate
flower imbricate; gynoecium pseudomono-
merous; fruit with apical stigmatic remains; em-
bryo basal; germination adjacent-ligular; eophyll
bifid.
The subtribe contains the single genus,
Cyrtostachys. It is distinguished by the strongly
divaricate inflorescence branches, rachilla pits
and connate petals and filaments in the staminate
flowers. Its relationships are obscure.
101. Cyrtostachys Blume
Cyrtostachys Blume, Bull. Sc. Phys. Nat. Neerl. 1: 66 (1838).
Solitary or clustered, erect. Inflorescence protan-
drous; rachillae elongate, cylindrical, bearing tri-
ads throughout much of their length. Stamens
9-15, filaments terete, connate basally, inflexed,
anthers dorsifixed, latrorse; pollen sulcate, exine
tectate variously ornamented; pistillode almost as
long as filaments. Staminodes joined in a ring;
gynoecium with 3 short recurved stigmas, ovule
pendulous from the apex of the locule. Eight spp.,
C. renda Blume in West Malesia, the others in New
Guinea and Melanesia.
4k. Subtribe Linospadicinae J.D. Hook. (1883).
Slender to moderately robust, unarmed, monoe-
cious palms; crownshaft absent; sheaths fibrous;
leaves pinnate or pinnately ribbed, leaflets entire
or irregularly lobed; inflorescences bisexual,
interfoliar, sometimes multiple, always spicate;
prophyll persistent, inserted at base of peduncle;
peduncular bract inserted near base of peduncle
and persistent, or inserted distally and caducous;
staminate flowers symmetrical; petals of pistillate
flowers imbricate; gynoecium pseudomono-
merous; fruit with apical stigmatic remains; ovule
hemianatropous, attached laterally above the
middle of the locule; embryo basal; germination
adjacent-ligular; eophyll bifid.
362
The subtribe consists of four closely related
genera occurring in Papuasia, Australia, and
Lord Howe Is. The group is separated on the basis
of the spicate inflorescences with triads borne in
deep pits and the lack of a crownshaft. The
rachillae are similar to those of members of
Geonomeae, but the structure of the flowers is
very different.
KEY TO THE GENERA OF LINOSP ADICINAE
1. Peduncular bract inserted near the base of the peduncle, 2:
flattened, opening apically, shorter than and not enclosing
the spike before anthesis, 2: persistent and marcescent;
filaments mostly inflexed; anthers dorsifixed; endocarp
adherent to the seed 102. Calyptrocalyx
- Peduncular bract inserted at the tip of the peduncle, tubu-
lar, enclosing the spike in bud, marcescent or caducous,
leaving a ruffled scar on the peduncle; filaments not in-
flexed, anthers basifixed; endocarp adherent or free from
the seed
2. Endocarp thick, cartilaginous, not adhering to the seed;
endosperm homogeneous; staminate flower with 30-70
stamens; stems solitary, stout; leaflets always 1-ribbed
105. Howea
- Endocarp thin, adhering to the seed; endosperm homoge-
neous or ruminate; staminate flower with 6-15 stamens;
stems usually clustered, usually slender; leaflets 1-many
ribbed
3. Seed with ruminate endosperm, raphe extending its length,
the branches reticulate; leaflets 1-ribbed
104. Laccospadix
- Seed with homogeneous endosperm, raphe extending 1/3
its length or less, the branches free or anatomosing; leaflets
1-several-ribbed or the blade undivided 103. Linospadix
102. Calyptrocalyx Blume
Calyptrocalyx Blume, Bull. Sc. Phys. Nat. Neerl. 1: 66 1838;
Burret, Notizbl. Bot. Gart. Mus. Berlin-Dahlem 12: 1331
{1935).
Solitary or caespitose, erect. Leaves undivided
or variously pinnate. Inflorescences solitary or
multiple; peduncular bract basally inserted, per-
sistent. Stamens 6-120, filaments inflexed, anthers
dorsifixed, latrorse; pollen sulcate, exine tectate
reticulate; pistillode slender, club-shaped, or
absent. Staminodes (2-)3-6. Epicarp smooth,
mesocarp fleshy or dry, endocarp thin. Seed sub-
basally to laterally attached, hilum short to elon-
gate, endosperm homogeneous or ruminate.
About 38 spp., probably fewer, one widespread in
the Moluccas, the remainder in New Guinea.
Palmae
103. Linospadix H. Wendl.
Linospadix H. Wend!. in H. Wend!. & Drude, Linnaea 39: 177,
198 {1875); Burret, Notizbl. Bot. Gart. Mus. Berlin-Dahlem
12: 331 (1935).
Solitary or caespitose, erect. Leaves undivided
or variously pinnate. Inflorescences solitary; pe-
duncular bract inserted distally, caducous. Sta-
mens 6-12, anthers dorsifixed, latrorse; pollen
sulcate, exine tectate reticulate; pistillode minute.
Staminodes 3-6. Epicarp smooth, mesocarp fleshy
with fibers adhering to endocarp, endocarp thin.
Seed subbasally attached, hilum elongate, en-
dosperm homogeneous. Eleven spp., five in New
Guinea, the rest in Australia.
104. Laccospadix Drude & H. Wendl.
2 Laccospadix Drude & H. Wend!., Nachr. Kon. Gesellsch. Wiss.
Georg-Augusts-Universitat 1875: 59 {1875).
Solitary or caespitose, erect. Leaves pinnate,
leaflets single-fold, entire. Inflorescences solitary
or multiple; peduncular bract inserted near tip of
peduncle, caducous. Stamens 9-12, filaments very
3
short, anthers basifixed, latrorse; pollen sulcate,
exine tectate reticulate; pistillode minute, trifid.
Staminodes 3. Epicarp smooth, mesocarp fleshy,
with stout longitudinal fibers next to endocarp,
endocarp thin. Seed laterally attached, hilum
short, oblong; endosperm ruminate. Only one sp.,
L. australasica H. W endl. & Drude, northeastern
Queensland, Australia.
105. Howea Becc.
Howea Becc., Malesia 1: 66 (1877); Bailey, Gentes Herb 4: 188-
198 (1939).
Solitary, erect. Leaves pinnate; leaflets numerous,
single-fold, acute. Inflorescences solitary or
multiple; peduncular bract inserted some distance
from base, caducous. Stamens 30-70, filaments
elongate, variously connate, anthers basifixed,
elongate, connective sometimes prolonged,
latrorse; pollen sulcate, exine tectate scabrate;
pistillode absent. Staminodes 3-6. Epicarp
smooth, mesocarp thinly fleshy with abundant
longitudinal fibers, endocarp thin. Seed laterally
attached; endosperm homogeneous. Two spp.,
Lord Howe Is.
 Palmae
41. Subtribe Ptychospermatinae J.D. Hook.
(1883).
Unarmed, pleonanthic, monoecious palms; leaves
pinnate, crownshaft always present; leaflets
single-fold, but sometimes longitudinally split-
ting, praemorse. Inflorescences infrafoliar, pro-
tandrous; peduncular bract tubular, slender,
much longer than the prophyll and emerging
through its tip; rachillae bearing spirally arranged
triads of flowers basally, and staminate flowers
distally; floral bracteoles low, rounded or truncate;
staminate flowers with sepals 3, distinct, imbri-
cate, rounded; petals boat-shaped, distinct, val-
vate; pistillode usually well-developed; pistillate
flower with distinct imbricate sepals and
petals; gynoecium pseudomonomerous, stigmas
3, recurved; ovule hemianatropous, attached
laterally or pendulous from the top of locule;
fruit with apical stigmatic remains; embryo basal;
germination adjacent-ligular; eophyll bifid, or
entire.
The subtribe comprises nine genera occurring
in the western Pacific region. The genera are
closely related and form a group distinct in the
form of the prophyll and the peduncular bract, the
staminate and pistillate flowers, and in the fruit. A
phylogenetic study of the Ptychospermatinae is
currently being undertaken by S. Zona; prelimi-
nary results (Zona, pers. comm.) suggest that
there will be major changes to the generic delimi-
tation presented here.
KEY TO THE GENERA OF THE
PTYCHOSPERMA TIN AE
1. Leaflets of juvenile leaves entire, cuneate, several-nerved;
leaflets of mature leaves divided longitudinally into 7-17
linear segments; peduncular bract similar to and included
within the prophyll, prophyll and peduncular bract
caducous
- Leaflets in both juvenile and mature leaves cuneate, trun-
cate acute, acuminate or oblique, undivided; peduncular
bract similar to or longer than and inserted well above and
protruding from the prophyll, prophyll and peduncular
bract marcescent
2. Stem slightly bottle-shaped; primary leaflets regularly ar-
ranged, divided into 11-17 segments, lacking white woolly
scales abaxially; outer endocarp with large, conspicuously
branched, tough black fibers; endosperm homogeneous
111. Wodyetia
- Stem moderate to slender, not bottle-shaped; primary
leaflets clustered, divided into 7-9 segments, bearing white
woolly scales abaxially; outer endocarp with sparsely
branched, thin, terete, straw-colored fibers; endosperm
ruminate 110. Normanbya
3. Seed rounded in cross-section
- Seed angled or grooved in cross-section
363
4. Leaflets broadly to narrowly wedge-shaped or broadly lan-
ceolate to obovate; peduncular bract terete and promi-
nently beaked, inserted well above and becoming exserted
from the prophyll 106. Drymophloeus
- Leaflets lanceolate, medium to narrow; peduncular bract
similar to and included within the prophyll 5
5. Leaflets wide in the middle or throughout, moderate to
rather short, truncate, oblique, acute, or acuminate, chaffy
ramenta present abaxially near the base of the midribs;
meso carp with several series of fibrous bundles, ± rounded
in section, next to the endocarp 108. Veitchia
- Leaflets narrow, long, usually with 2-4 rather short jagged
points, lacking chaffy ramenta on midribs; meso carp with a
single series oflarge, flat fibrous bundles closely adherent to
the endocarp 107. Carpentaria
6. Peduncle elongate; peduncular bract inserted well above
and exserted from the prophyll; fruit often angled and ta-
pered at both ends; seed irregularly angled in cross-section,
acute 109. Balaka
- Peduncle short, peduncular bract similar to and included in
the prophyll; fruit ovoid; seed 3-5-furrowed in cross-
section, acute and truncate or pointed at both ends 7
7. Leaflets acute, oblique, truncate, or 2-pronged and concave
apically; anthers dorsifixed, sagittate basally; fruit rounded
or irregularly ridged when dry; seed with 5 rounded or
irregular ridges 8
- Leaflets 3-pronged at the apex, the center prong the longest
anthers basifixed, not versatile; fruit very strongly 5-angled
when dry; the seed with 5 acute ridges 114. Brassiophoenix
8. Seed rounded apically and with 5 rounded ridges; endocarp
thin, fibrous or, if somewhat bony, then not elaborated and
staminate flowers with short, conic-ovoid pistillode.
Malesia to Micronesia and NE Australia112. Ptychosperma
- Seed pointed apically, with irregular ridges; endocarp very
thick, hard; pistillode always bottle-shaped
113. Ptychococcus
106. Drymophloeus Zipp.
Drymophloeus Zipp., Algemene Konst- en Letter-bode 1829
(19): 297 (1829); Zona, Blumea (in press).
Solitary, stilt roots sometimes present. Inflores-
cence branched to 1 to 2-3 orders. Stamens 25 to
more than 200, anthers elongate, sagittate, un-
2 even, medifixed, versatile, latrorse; pollen sulcate
or trichotomosulcate, exine tectate, punctate;
pistillode with 3 short, rounded, 3-angled lobes
or ovoid-attenuate, to 2/3 as long as stamens.
3 Staminodes 3, shortly joined basally. Fruit with
epicarp thin, smooth, mesocarp fleshy, fibrous,
with raphides, fibers adherent to thin, smooth
endocarp. Seed ovoid, hilum apical, endosperm
homogeneous or ruminate. About 15 spp., from
Halmahera, Ceram, Ambon to New Guinea, the
Solomons, to Fiji and Samoa.
107. Carpentaria Becc.
4
6 Carpentaria Becc., Ann. Jard. Bot. Buitenzorg 2: 128 (1885).
364 Palmae
Solitary. Inflorescences branched to 3 orders. Sta-
mens ca. 33, filaments short, anthers elongate, bi-
fid basally, dorsifixed, latrorse, connective broad,
tanniniferous; pollen sulcate, exine tectate, finely
reticulate; pistillode bottle-shaped, as long as sta-
mens in bud. Staminodes 3, toothlike, bifid
apically. Epicarp smooth, mesocarp fleshy over a
layer of broad, flat fibers, anastomosing distally,
closely appressed to endocarp, endocarp thin,
glasslike. Seed attached laterally, round in cross-
section, hilum elongate, raphe branches anasto-
mosing, endosperm homogeneous. Only one sp.,
C. acuminata (H. Wendl. & Drude) Becc., North-
ern Territory, Australia.
108. Veitchia H. Wendl.
Veitchia H. Wendl. in Seemann, Flora Vitiensis: 270 (1868);
H.E. Moore, Gentes Herb. 8: 483-536 (1957).
Solitary. Inflorescence branched to 3 or 4 orders
basally. Stamens to 100 or more, filaments slender,
united basally, not inflexed, anthers linear,
basifixed or dorsifixed, sagittate, latrorse; pollen
sulcate, exine tectate; pistillode attenuate from a
bulbous base, equalling stamens. Staminodes 3-6,
toothlike, variously connate. Epicarp smooth, me-
socarp, thin-fleshy, with 2-severallayers of fibers,
endocarp thin to thick, fragile. Seed attached later-
ally, hilum elongate; endosperm homogenous or
ruminate [V. merrillii (Becc.) H.E. Moore]. About
18 spp., Vanuatu, Fiji, and Philippine Is., ten spp.
endemic to Fiji.
109. Balaka Becc.
Balaka Becc., Ann. Jard. Bot. Buitenzorg 2: 91 (1885).
Solitary. Leaflets single-fold. Inflorescences
branched to 2 orders. Stamens 24-50, filaments
erect, anthers elongate, dorsifixed, sagittate, bifid
apically, latrorse; pollen sulcate, exine tectate;
pistillode bottle-shaped with long neck. Sta-
minodes usually 6, small, united. Fruit tapered
distally or at both ends, often angled; epicarp
smooth, mesocarp with outer sclereid layer, inner
fleshy layer, and single series of vascular bundles
next to thin endocarp. Seed elongate, pointed,
ridged, 4-5-angled in cross-section, hilum elon-
gate, lateral. About seven spp., five endemic to Fiji
and two in Samoa.
110. Normanbya F. Muell. ex Becc.
Normanbya F. Muell. ex Becc., Ann. Jard. Bot. Buitenzorg 2:
91, 170, 171 (1885).
Solitary. Leaflets singlefold, longitudinally split
into numerous segments held in different planes.
Inflorescences branched to 2(? more) orders. Sta-
mens 24-40, filaments short, anthers elongate,
shortly bifid apically, dorsifixed, introrse, connec-
tive elongate, tanniniferous; pollen sulcate, exine
tectate, scabrate; pistillode flask-shaped with long
narrow neck, slightly longer than stamens.
Staminodes 3. Epicarp smooth, mesocarp thin,
with longitudinal, branched, straw-colored fibers
adherent to smooth endocarp. Seed laterally at-
tached, endosperm ruminate. Only one spp., N.
normanbyi (W. Hill) L.H. Bailey, rainforests of
northern Queensland, Australia.
111. Wodyetia A.K. Irvine
Wodyetia A.K. Irvine, Principes 27: 161 (1983).
Solitary, erect. Leaflets single-fold, divided into
segments, held in different planes. Inflorescences
branched to 4 orders. Stamens 60-71, filaments
slender, short, anthers elongate, narrow, sagittate,
bifid apically, dorsifixed, latrorse, connective
tanniniferous; pollen sulcate, exine tectate, punc-
tate; pistillode bottle-shaped with long neck.
Staminodes 6. Epicarp smooth; mesocarp thin
with longitudinal fibers, some forked, endocarp
complex with outer thick, flat, branched fibers
and inner layer of horizontal fibers. Endosperm
homogeneous. Only one sp., W. bifurcata A.K.
Irvine, northeastern Queensland, Australia.
112. Ptychosperma Labill.
Ptychosperma Labill., Mem. Classe Sci. Math. Phys. lost. Nat.
France 1808 (2): 252 (1809); Essig, Allertonia 1: 415-478
(1977).
Solitary or clustered. Leaflets single-fold. Inflores-
cences branched to 2, 3, or 4 orders. Stamens 9-
100, filaments short, not inflexed, anthers
linear-lanceolate, sagittate, bifid apically, dor-
sifixed, versatile, latrorse, connective tannni-
ferous; pollen sulcate, exine tectate, reticulate;
pistillode bottle-shaped with a long neck, or coni-
cal. Staminodes toothlike, linear or united. Epi-
carp smooth; mesocarp fleshy, mucilaginous or
tanniniferous, sometimes with raphides, endocarp
fibrous. Seed longitudinally 5-grooved or angled,
rarely 3-grooved or rounded in cross-section, hi-
lum lateral, endosperm homogeneous or rumi-
nate. Twenty eight spp., centered in New Guinea
and the D'Entrecasteaux and Louisiade archipela-
gos but extending west to east from the Moluccas
 Palmae
to the Solomon Islands and south from the
Caroline Islands to northeastern Australia.
113. Ptychococcus Becc.
Ptychococcus Becc., Ann. Jard. Bot. Buitenzorg 2: 100 (1885).
Solitary. Leaflets single-fold. Stamens numerous
(-100), filaments short, anthers elongate, notched
apically, bifid basally, dorsifixed, introrse, connec-
tive tanniniferous; pollen elliptic, sulcate, exine
tectate, reticulate; pistillode bottle-shaped equal-
ling stamens. Staminodes 3, united in a cupule.
Epicarp smooth, mesocarp fleshy with tannini-
ferous cells and fibers, endocarp thick with large
keel, deeply grooved between 3 lateral and 2 ven-
tral ridges. Seed 5-lobed, hilum circular, apical,
endosperm with shallow ruminations and deep
intrusion in the rapheal lobes. Seven spp. de-
scribed, all rather poorly known; six in New
Guinea and one in the Solomon Is.
114. Brassiophoenix Burret
Brassiophoenix Burret, Notizbl. Bot. Gart. Mus. Bertin-
Dahlem 12: 345 (1935); Essig, Principes 19: 100-103 (1975).
Solitary, erect; leaflets single-fold, wedge-shaped,
apically trilobed, the center lobe much prolonged.
Inflorescences branched to 2( -3) orders. Stamens
numerous (ca. 100-230), inserted on a conical re-
ceptacle, filaments long, slender, anthers elongate,
basifixed, latrorse, connective tanniniferous, pro-
longed basally between the anthers; pollen sulcate,
exine tectate, finely reticulate; pistillode small,
conical. Staminodes forming a shallow 3-lobed
ring. Epicarp with short, single, oblique fibrous
bundles and interspersed brachysclereids, meso-
carp fleshy, endocarp hard, thick, ridged. Seed lat-
erally attached, irregular with 5 ridges, pointed
distally, hilum elongate, endosperm homoge-
neous. Two species, New Guinea.
4m. Subtribe Arecinae J.D. Hook. (1883).
Diminutive to large, unarmed; crownshaft usually
present; leaves pinnate or pinnately-ribbed;
leaflets entire or praemorse; inflorescence usually
infrafoliar; prophyll and peduncular bract (where
present) inserted close together, enclosing the in-
florescence in bud, splitting and falling at anthesis;
staminate flower usually asymmetrical; petals
usually pointed; pistillate flower smaller than the
staminate, except in Areca, where usually larger;
365
gynoecium pseudomonomerous; fruit with apical
stigmatic remains; embryo basal; germination ad-
jacent ligular.
The eight genera of Arecinae display consider-
able morphological diversity. Areca, Pinanga, and
Nenga are unusual in lacking a large sheathing
peduncular bract. These three genera are pre-
dominantly West Malesian, although Areca has a
minor radiation in Papuasia. Gronophyllum,
Siphokentia, Hydriastele, and Gulubia are closely
related to each other and rather poorly delimited.
They are much in need of a modern reappraisal.
Pollen structure is extraordinarily diverse in
terms of aperture in Areca and in terms of exine
ornamentation in Pinanga.
KEY TO THE GENERA OF ARECINAE
1. Inflorescence bearing 2 large bracts, the prophyll and a
similar peduncular bract, the peduncle later bearing 2 cir-
cular scars at the base 2
- Inflorescence bearing 1 large bract, the prophyll alone, the
peduncle later bearing a single circular scar 6
2. Rachillae bearing spirally arranged triads in the proximal
1/2-3/4, distally bearing paired or solitary staminate
flowers; peduncular bract distinctly shorter than the
prophyll, scarcely sheathing the inflorescence
115. Loxococcus
- Rachillae bearing opposite and decussate or whorled, very
rarely spirally arranged triads ± throughout their length;
peduncular bract :±: as large as the prophyll 3
3. Pistillate flower bearing connate sepals and petals
117. Siphokentia
- Pistillate flower bearing free sepals and petals 4
4. Inflorescence protandrous; petals of pistillate flowers more
than twice as long as the sepals, with conspicuous long,
valvate tips 116. Gronophyllum
- Inflorescence protogynous; petals of pistillate flower less
than twice as long as the sepals with very short valvate tips
5
5. Robust solitary palms; leaves large; leaflets acute or irregu-
larly bilobed at the tip 119. Gulubia
- Moderate, usually clustered palms; leaves short; leaflets
conspicuously praemorse at the tip 118. Hydriastele
6. Inflorescence protogynous, spicate or branched to 1 order
only; rachillae bearing distichous or, more rarely, spirally
arranged triads throughout their length 121. Pinanga
- Inflorescence protandrous (in a few Areca spp. recorded as
protogynous but perhaps erroneously so), spicate or
branched to 1-3 orders; rachillae bearing triads proximally
and paired or solitary, spirally arranged, distichous or
unilateral staminate flowers distally 7
7. Triads usually very few in number at the base of each
rachilla, rarely numerous; seed basally attached with circu-
lar hilum 122. Areca
- Triads numerous, often occupying 3/4 of the rachilla length;
seed laterally attached with narrow longitudinal hilum
120. Nenga
115. Loxococcus H. Wendl. & Drude
Loxococcus H. Wend!. & Drude, Linnaea 39: 185 (1875).
366
Solitary. Crownshaft present. Leaflets single-fold,
linear, praemorse. Inflorescence infrafoliar,
branching to 2 orders, protandrous; peduncular
bract borne just above prophyll, not completely
sheathing. Rachillae bearing spirally arranged tri-
ads in basal 1/2-3/4, solitary or paired staminate
flowers distally. Staminate flowers asymmetrical;
stamens 12, anthers elongate, latrorse; pollen sul-
cate, exine tectate; pistillode conical. Pistillate
flowers with distinct sepals and petals. Seed glo-
bose, basally and laterally attached, hilum lateral
throughout length of seed; endosperm deeply
ruminate. Only one sp., L. rupicola (Thwaites)
H. Wendl. & Drude, Sri Lanka.
116. Gronophyllum Scheff.
Gronophyllum Scheff., Ann. Jard. Bot. Buitenzorg 1: 135
(1876); Essig & Young, Principes 29: 129-137 (1985).
Solitary or clustered. Crownshaft present. Lamina
entire or variously pinnate, leaflets acute, bifid or
praemorse. Inflorescence infrafoliar, spicate or
branching to 1-3 orders, often hippuriform, pro-
tandrous; peduncular bract similar to prophyll;
rachillae bearing opposite and decussate, or alter-
nating whorls of 3 triads throughout. Staminate
flowers asymmetrical; stamens 6-9, anthers
basifixed, elongate, latrorse; pollen sulcate, exine
tectate, reticulate; pistillode minute. Pistillate
flowers with distinct sepals and petals. Seed later-
ally attached; endosperm homogeneous or rumi-
nate. Eophyll bifid. About 33 spp., Sulawesi
through the Moluccas to New Guinea, the
Bismarck Archipelago, and Australia.
ll7. Siphokentia Burret
Siphokentia Burret, Notizb. Bot. Gart. Mus. Berlin- Dahlem 10:
198 (1927); Moore, Gentes Herb. 8: 295-315 (1953).
Solitary. Crownshaft present. Lamina entire or
variously pinnate, leaflets praemorse. Inflores-
cence infrafoliar, branching to 1 order, pro-
tandrous; peduncular bract similar to prophyll;
rachillae bearing opposite and decussate triads
throughout. Staminate flowers asymmetrical; sta-
mens 9-10, anthers basifixed, elongate, latrorse;
pollen sulcate, exine, tectate, coarsely reticulate;
pistillode minute. Pistillate flowers with sepals
and petals connate in basal 1/2, distally 3-lobed.
Seed basally attached; endosperm ruminate.
Two spp., Halmahera, Obi, and Kahatola, in the
northwest Moluccas; perhaps also in New
Guinea.
Palmae
ll8. Hydriastele H. Wendl. & Drude
Hydriastele H. Wendl. & Drude, Linnaea 39: 180, 208 (1875).
Caespitose. Crownshaft present. Leaflets mostly
single-fold, praemorse. Inflorescences infrafoliar,
branching to 1-2 orders, protogynous; peduncu-
lar bract similar to the prophyll; rachillae bearing
opposite and decussate pairs of triads throughout.
Staminate flowers asymmetrical; stamens 6, an-
thers elongate, basifixed, latrorse; pollen, sulcate,
exine tectate, scabrate or reticulate; pistillode ab-
sent. Pistillate flowers with distinct sepals and pet-
als. Seed laterally attached with elongate hilum;
endosperm homogeneous or ruminate. Eophyll
bifid. About eight spp., New Guinea, Bismarck
Archipelago, and northern Australia.
119. Gulubia Becc.
Gulubia Becc., Ann. Jard. Bot. Buitenzorg 2: 128, 131, 134
(1885); Essig, Principes 26: 159-173 ( 1982).
Solitary. Crownshaft present. Leaflets mostly
single-fold, acute or bifid. Inflorescences infra-
foliar, branching to 1-3 orders, protogynous; pe-
duncular bract similar to the prophyll; rachillae
bearing opposite and decussate pairs of triads
throughout. Staminate flowers asymmetrical; sta-
mens 6-24, anthers elongate, basifixed, latrorse;
pollen sulcate, exine tectate, reticulate, granulate;
pistillode minute or absent. Pistillate flowers with
distinct sepals and petals. Seed laterally attached
with elongate hilum; endosperm homogeneous or
ruminate. Eophyll bifid. Nine spp., Moluccas to
New Guinea, Bismarck Archipelago, Solomon
Islands, Australia, Vanuatu, Fiji, and Palau.
120. Nenga H. Wendl. & Drude
Nenga H. Wend!. & Drude, Linnaea 39: 182 (1975); Fernando,
Principes 27: 55-70 (1983).
Solitary or caespitose, acaulescent or erect, crown-
shaft present or not, leaflets with !-several folds,
acuminate. Inflorescence infrafoliar or interfoliar,
spicate or branching to 1 or 2 orders, protandrous;
peduncular bract absent; rachillae bearing spiral
flowers, triads proximally, solitary or paired
staminate flowers distally. Stamens 6, anthers ob-
long to linear, latrorse; pollen sulcate, exine
tectate coarsely reticulate, pistillode absent. Pistil-
late flowers with distinct sepals and petals. Seed
laterally attached, hilum longitudinal, endosperm
deeply ruminate; embryo basal. Eophyll bifid. Five
 Palmae 367

sp., from Vietnam and Burma to Sumatra, the 3, distinct, broadly imbricate; petals 3, distinct,
Malay Peninsula, Borneo, and Java. broadly imbricate except for the minute, triangu-
lar, valvate tips; gynoecium pseudomonomerous;
ovule hemianatropous (? always), laterally at-
121. Pinanga Blume Fig. 82C
tached; endocarp with operculum; embryo almost
Pinanga Blume, Bull. Sc. Phys. Nat. Neerl. 1: 65 (1838). always basal (lateral in Alloschmidia); germina-
tion adjacent-ligular.
Solitary or caespitose, acaulescent or erect, A perplexingly complex subtribe of genera
crownshaft present or not, blade entire or vari- found mostly in the western Pacific region, but
ously pinnate, leaflets acuminate. Inflorescence with outliers in the Indian Ocean. The subtribe is
infrafoliar or interfoliar, spicate or branching to 1 distinctive in the presence of an operculum on the
order, protogynous; prophyll rarely persistent; endocarp (not always easy to see). The group
peduncular bracts absent; rachillae bearing spiral seems most closely related to Oncospermatinae,
or distichous triads throughout. Staminate flowers that differs in the single character of the presence
asymmetrical, stamens (6) 12-30, anthers linear, of epidermal emergence spines. Relationships of
latrorse; pollen sulcate, exine intectate or tectate, the two subtribes and their contained genera are
varied in ornamentation; pistillode absent. Pistil- in need of further phylogenetic study, which is
late flowers with distinct sepals and petals or being undertaken by Carl Lewis at Cornell
petals connate. Seed basally attached, hilum circu- University.
lar; endosperm homogeneous or ruminate; em-
bryo basal. Eophyll bifid. About 120 spp. ranging KEY TO THE GENERA OF THE IGUANURINAE
from the Himalayas and south China to New
Guinea. 1. Prophyll completely encircling the peduncle at insertion,
leaving a circular scar when caducous; stamens 6 or more
2
122. Areca L. Fig. 82K - Prophyll incompletely encircling the peduncle at inser-
tion, opening abaxially, leaving an incomplete scar upon
Areca L., Sp. Pl.: 1189 (1753). falling; stamens always 6 21
2. Seed irregularly ridged, furrowed, and sculptured with
adherent fibers 136. Alsmithia
Solitary or caespitose, acaulescent or erect; crown- - Seed small, not ridged or sculptured 3
shaft present or not; blade entire or variously pin- 3. Staminate flowers borne in vertically oriented pairs
nate, leaflets acuminate. Inflorescence infrafoliar sunken in distinct depressions distally, smaller than and
or interfoliar, spicate or branching to up to 3 or- lateral to the pistillate flowers proximally on the rachillae;
ders, protandrous or protogynous; peduncular fruit large, with apical stigmatic remains 123. Neoveitchia
- Staminate flowers borne in horizontally oriented pairs
bracts absent; rachillae bearing spiral or disti- distally, lateral to pistillate flowers proximally on the
chous triads basally, paired or solitary staminate rachillae; fruit moderate, rarely large with apical, lateral,
distally. Staminate flowers ± symmetrical, sta- or basal stigmatic remains 4
mens 3-30, anthers latrorse or poricidal; pollen 4. Inflorescence interfoliar; fruit covered with prominent
sulcate, monoporate or triporate, exine tectate or corky warts; stigmatic remains basal in fruit 5
- Inflorescence interfoliar, or infrafoliar, fruit smooth or
intectate. Pistillate flowers with distinct sepals and merely pebbled to granulose when dry; stigmatic remains
petals. Seed basally attached, hilum circular; en- various 6
dosperm ruminate; embryo basal. Eophyll bifid. 5. Peduncular bract inserted near the base of the peduncle;
About 60 spp., from India and south China fruit more than 2.5 em in diameter 124. Pelagodoxa
- Peduncular bract inserted at the apex of the peduncle; fruit
through Malesia to New Guinea and the Solomon 1.5 em in diameter or less 129. Sommieria
Islands. 6. Flowers borne in laterally compressed pits, the staminate
on long, hairy pedicels; fruit with stigmatic remains lateral
in lower 1/4; seeds ridged and grooved 130. Bentinckia
4n. Subtribe Iguanurinae J.D. Hook. (1883). - Flowers sessile or impressed in the rachillae but neither in
laterally compressed pits nor the staminate with hair-
covered pedicels; stigmatic remains apical to basal; seeds
Unarmed, pleonanthic, monoecious palms with ~o~h 7
bisexual inflorescences; prophyll tubular, com- 7. Leaflets several-ribbed with praemorse apices or leaf
pletely sheathing the peduncle, or incompletely blades, when not divided with toothed margins; inflores-
tubular and only partially sheathing; flowers cences usually interfoliar; triads shallowly to deeply
sunken in depressions in the rachillae 125. Iguanura
sometimes borne in pits; protandrous; pollen el- - Leaflets 1-ribbed with acute or acuminate apices; inflores-
liptic, sulcate, exine tectate; pistillate flowers cences various; triads superficial 8
much larger than the staminate, spherical; sepals 8. Seed with ruminate endosperm 9
368 Palmae
- Seed with homogeneous endosperm 12
9. Leaf sheaths splitting opposite the petiole, not forming a
prominent crownshaft; inflorescences interfoliar, at least
in bud, sometimes infrafoliar at anthesis or in fruit; the
peduncle elongate, prominent, usually as long as the rachis
or longer 128. Heterospathe
- Leaf sheaths tubular, forming a prominent crownshaft;
inflorescence infrafoliar; peduncle usually much shorter
than the rachis 10
10. Inflorescence lacking branches adaxially except at the
apex, branched to 1 order only and the lower branches
ascending, not divaricate from the rachis at a 90° angle;
fruit black at maturity 139. Dictyosperma
- Inflorescence with branches spirally arranged, the lower
branches abruptly divaricate at about a 90° angle from the
rachis and again once- or twice-branched; fruit yellow,
orange, or red 11
11. Stamens 6-9; pistillode prominent 138. Rhopaloblaste
- Stamens 15-30 or more; pistillode minute or lacking
140. Actinorhytis
12. Staminate flowers mostly larger than the pistillate; stamen
filaments inflexed at the apex in bud; anthers dorsifixed,
with elongate connective, not didymous 13
- Staminate flowers mostly smaller than the pistillate;
stamen filaments erect in bud; anthers didymous 19
13. Stamens 12; fruit with basal stigmatic remains, lacking a
shell of sclereids 132. Cyphokentia
- Stamens 6; fruit various 14
14. Endocarp minutely pitted; seed with lateral embryo
142. Alloschmidia
- Endocarp not pitted; seed with basal embryo 15
15. Leaf sheaths split opposite the petiole; inflorescence
interfoliar 127. Lepidorrhachis
- Leaf sheaths forming a prominent crownshaft; inflores-
cence infrafoliar 16
16. Inflorescence densely tomentose; fruit with apical stig-
matic residue 137. Satakentia
- Inflorescence glabrous or at most minutely hairy; fruit
with subapical to nearly basal stigmatic remains 17
17. Complete peduncular bracts 2; endocarp hard, moderately
thick, with longitudinal ridges 135. Carpoxylon
- Complete peduncular bracts 1; endocarp thin or thick and
prominently sculptured 18
18. Stilt roots usually developed; staminate flowers markedly
asymmetrical, with short, trifid pistillode and acute sepals
and petals; fruit lacking sclereids but with prominent,
often greatly thickened fibers 134. Clinostigma
- Stilt roots not developed; staminate flowers symmetrical,
with pistillode as long as stamens in bud and rounded
sepals and petals; fruit with a layer of short sclereids
beneath the exocarp 133. Moratia
19. Fruit ellipsoidal with basal stigmatic remains; sclereids
lacking in mesocarp but tannin cells present
126. Brongniartikentia
- Fruit globose or nearly so with lateral stigmatic remains;
mesocarp with a shell of short sclereids beneath the
exocarp 20
20. Leaf sheaths with minute scales, split opposite the petiole
and not forming a crownshaft; peduncle short; fruit small,
1.4-1.6cm in diameter, with tannin cells interior to
sclereid layer 131. Clinosperma
- Leaf sheaths densely scaly, tubular and forming a promi-
nent crownshaft; peduncle elongate; fruit large, ca. 3.2 em
in diameter, lacking tannin cells 141. Lavoixia
21. Seed terete or 2-lobed in cross-section, ovoid, ellipsoidal,
globose, or rarely kidney-shaped in outline 22
- Seed irregular in cross-section, externally angled or in-
tricately ridged, furrowed, and sculptured 24
22. Fruit with apical stigmatic remains 23
- Fruit with lateral stigmatic remains 145. Basselinia
23. Stilt roots prominent and stout; pistillode of staminate
flower shorter than stamens; fruit often curved apically
144. Campecarpus
- Stilt roots not prominently developed; pistillode of stami-
nate flower longer than stamens, columnar; fruit straight
143. Cyphophoenix
24. Leaf sheaths split opposite the petiole in bud, not forming
a prominent crownshaft; inflorescence among the leaves in
bud, becoming infrafoliar in fruit; peduncle elongate,
much exceeding the rachis; prophyll and peduncular bract
:±:: persistent, at length marcescent; inflorescence branches
with long bare basal portions, prominently swollen at the
insertion, stiffly and divaricately branched to 1 order or
the distal undivided 146. Cyphosperma
- Leaf sheaths forming a prominent crownshaft; inflores-
cence infrafoliar; peduncle shorter than the rachis;
prophyll and peduncular bracts caducous; inflorescence
branches without a long basal bare portion, nor swollen at
the insertion 25
25. Staminate flowers symmetrical; pistillode thick, columnar,
longer than the stamens in bud, expanded into a broadly
capitate apex; fruit sub globose with stigmatic remains !at-
era! in upper third, the surface minutely granular-papillate
147. Veillonia
- Staminate flowers slightly to markedly asymmetrical, the
pistillode elongate-conic to angled-columnar, shorter than
the stamens in bud, not broadly capitate; fruit smooth or
drying pebbled but not granular-papillose 26
26. Bracteoles surrounding the pistillate flower sepal-like; an-
thers with locules not continuous but interrupted by ster-
ile connectivelike areas; fruit drying densely pebbled and
shouldered; mesocarp not readily separable from the
stony, intricately sculptured, 4-angled endocarp with a
dorsal groove flanked by 2 ridges 148. Burretiokentia
- Bracteoles surrounding the pistillate flower very narrow,
rarely (P. dennisii H.E. Moore) with a slender limb but
never sepal-like; locules of anthers continuous; fruit glo-
bose or subglobose or collapsing and drying wrinkled but
not pebbled; mesocarp with a shining inner layer adjacent
to and readily separated from the endocarp, endocarp
sharply 4-angled to variously ridged and sculptured but
always with a dorsal ridge 149. Physokentia
123. Neoveitchia Becc.
Neoveitchia Becc., Le Palme della Nuova Caledonia: 9 (1920};
and Webbia 5: 77 (1921}; Dowe & Cabalion, Austr. Syst. Bot.
9: 1-60 (1996}.
Solitary, erect. Crownshaft absent. Leaflets single-
fold, acute. Inflorescences infrafoliar, branching
to 3 orders, protandrous; prophyll tubular, de-
ciduous, peduncular bract protruding through
prophyll, deciduous. Rachillae with triads basally,
staminate distally, partially sunken in pits. Stami-
nate flowers asymmetrical, in vertical pairs, sepals
variously connate; stamens 6, filaments distinct,
inflexed; anthers medifixed, latrorse; pistillode
columnar, equalling stamens. Staminodes 3,
 Palmae
toothlike; ovule laterally attached. Stigmatic re-
mains apical; endocarp smooth, operculum cir-
cular. Seed laterally attached, hilum elongate;
endosperm homogeneous. Two spp., Viti Levu,
Fiji, and Vanuatu.
124. Pelagodoxa Becc.
Pelagodoxa Becc. in Bois, Rev. Hort. II, 15: 302 (1917).
Solitary. Crownshaft absent. Lamina entire,
margins praemorse. Inflorescences interfoliar,
branching to 2 orders, protandrous; peduncular
bract longer than prophyll. Rachillae with triads in
pits, staminate flowers distally. Staminate sepals 3,
distinct; petals 3, basally connate; stamens 6, fila-
ments united basally to pistillode, anthers medi-
fixed, sagittate, latrorse, connective prolonged;
pistillode pyramidal. Staminodes 3, triangular,
small. Stigmatic remains basal; epicarp corky-
warted; endocarp smooth. Seed basal, hilum
rounded; endosperm homogeneous. Eophyll bifid.
Only one sp., P. henryana Becc., Marquesas.
125. Iguanura Blume
Iguanura Blume, Bull. Sc. Phys. Nat. Neerl. 1:66 (1838); Kiew,
Gard. Bull. Singapore 28: 191-226 (1976); Lim, Gard. Bull.
Singapore 48: 1-64 ( 1998).
Solitary or clustered, acaulescent or erect. Leaves
undivided or variously pinnate, praemorse,
crownshaft present or not. Inflorescences inter-
foliar or infrafoliar, spicate or branching to 1 or 2
orders; rachillae with spiral triads throughout,
superficial or sunken in pits. Staminate flowers
sessile, symmetrical, sepals 3, distinct; petals 3,
basally connate; stamens 6, filaments distinct, in-
flexed, anthers oblong, latrorse; pistillode colum-
nar, equalling petals. Staminodes 6. Stigmatic
remains basal; epicarp smooth, mesocarp fleshy,
endocarp smooth or variously ridged, operculum
basal, circular. Seed basally attached, hilum cir-
cular, endosperm homogeneous or ruminate.
Eophyll entire. About 25 spp., Sumatra, Malay
Peninsula (including south Thailand), and
Borneo.
126. Brongniartikentia Becc.
Brongniartikentia Becc., Le Palme della Nuova Caledonia: 48
(1920); Moore & Uhl, Allertonia 3: 313-402 (1984).
Solitary, erect. Crownshaft absent. Leaflets single-
fold, acute. Inflorescence interfoliar, becoming
369
infrafoliar, branched to 1 or 2( -3) orders; pedun-
cular bract inserted near and exceeding prophyll;
rachillae bearing spiral triads basally, staminate
distally. Staminate flowers symmetrical, sepals
and petals distinct; stamens 6, filaments short, not
inflexed, anthers didymous with separated locules
introrse; pistillode longer than stamens. Sta-
minodes 3. Stigmatic remains subbasal; epicarp
smooth, endocarp thin, vitreous, fragile, oper-
culate basally. Seed with basal circular hilum
endosperm homogeneous. Two spp., New
Caledonia.
127 Lepidorrhachis (H. Wendl. & Drude)
O.F. Cook
Lepidorrhachis (H.A. Wendl. & Drude) O.F. Cook, J. Hered.
18: 408 (1927).
Clinostigma section Lepidorrhachis H. Wendl. & Drude,
Linnaea 39: 218 (1875).
Solitary, erect. Crownshaft absent. Leaflets single-
fold, acute. Inflorescences interfoliar becoming
infrafoliar, branched to 3 orders; rachillae bearing
superficial triads nearly throughout. Staminate
flowers asymmetrical; sepals and petals distinct.
Stamens 6, filaments distinct, inflexed, anthers
oblong, latrorse; pistillode cylindrical, slightly
longer than stamens in bud. Staminodes 3. Stig-
matic remains lateral in upper 1/3-1/4; epicarp
smooth, mesocarp fleshy, thin, with included lon-
gitudinal fibers, endocarp fragile. Seed with
elongate, apical hilum, endosperm homogeneous;
embryo subbasal. Only one sp., L. mooreana
(F. Muell.) O.F. Cook, Lord Howe Is.
128. Heterospathe Scheff.
Heterospathe Scheff., Ann. Jard. Bot. Buitenzorg 1: 141 (1876).
Solitary or clustered, acaulescent or erect.
Crownshaft absent. Leaflets single-fold, acute. In-
florescences interfoliar or infrafoliar, branched to
1-4 orders; prophyll complete, peduncular bract 1
or 2, exceeding prophyll; rachillae bearing spiral
triads throughout, or with staminate distally.
Staminate flower asymmetrical; sepals and petals
distinct; stamens 6-36, filaments inflexed, anthers
dorsifixed, versatile, latrorse; pistillode present.
Staminodes 3. Stigmatic remains eccentrically api-
cal or lateral; epicarp smooth, endocarp thin. Seed
attached apically and laterally, hilum elongate,
endosperm homogeneous or ruminate. Eophyll
bifid. About 32 spp. the Philippines and Micro-
nesia to eastern Indonesia, New Guinea, Solomon
Islands, Vanuatu, and Fiji.
370
129. Sommieria Becc.
Sommieria Becc., Malesia 1: 66 (1877).
Solitary, acaulescent or erect. Crownshaft absent.
Leaves entire, or irregularly pinnate, segments
acute, discolorous. Inflorescences interfoliar,
branching to 1 order; prophyll short; peduncular
bract borne at tip of peduncle; rachillae few, bear-
ing partially sunken triads throughout. Staminate
flowers symmetrical, sepals and petals distinct;
stamens 6, filaments connate basally, inflexed, an-
thers medifixed, versatile, latrorse; pistillode co-
lumnar. Staminodes 3-6. Stigmatic remains basal;
mesocarp corky-warted, endocarp thin. Seed ba-
sally attached, hilum circular; endosperm homo-
geneous; embryo subbasal. Eophyll bifid. Three
spp., New Guinea and Waigeo.
130. Bentinckia A. Berry
Bentinckia A. Berry in Roxb., Flora Indica (2nd edn.) 3: 621
(1832).
Solitary, erect. Crownshaft present. Leaflets
singlefold, acute. Inflorescences infrafoliar,
branched to 3 orders; prophyll and peduncular
bract similar, enclosing inflorescence in bud;
rachillae bearing spiral triads in laterally com-
pressed pits. Staminate flowers asymmetrical,
sepals and petals distinct; stamens 6, filaments
distinct, inflexed, anthers elliptic, basifixed,
latrorse; pistillode as long as stamens. Staminodes
3-6. Stigmatic remains lateral near base; epicarp
smooth, mesocarp fibrous, endocarp grooved.
Seed grooved, hilum rounded, basal; endosperm
homogeneous. Eophyll bifid. Two spp., Western
Ghats, India, and Nicobar Is.
131. Clinosperma Becc.
Clinosperma Becc., Le Palme della Nuova Caledonia: 51
(1920); Moore & Uhl, Allertonia 3: 313-402 (1984).
Solitary, erect. Crownshaft absent. Leaflets single-
fold, acute. Inflorescences infrafoliar, branched to
3 orders; prophyll complete, enclosing peduncular
bract, both caducous; rachillae bearing spiral
triads basally, staminate flowers distally. Stami-
nate flowers symmetrical; sepals and petals dis-
tinct; stamens 6, distinct, filaments not inflexed,
anthers didymous; pistillode equalling stamens.
Staminodes 3. Stigmatic remains lateral; epicarp
smooth, mesocarp with inner layer of fibers, en-
docarp vitreous, operculum basal, rounded. Seed
Palmae
with short basal hilum; endosperm homogeneous.
Only one sp., C. bracteale (Brongn.) Becc., New
Caledonia.
132. Cyphokentia Brongn.
Cyphokentia Brongn., Comptes Rendus Acad. Sc. 77: 399
(1873); Moore & Uhl., Allertonia 3: 313-402 (1984).
Solitary or caespitose, erect. Crownshaft present.
Leaflets single-fold, acute. Inflorescences infra-
foliar, branched to 2( -3) orders; prophyll and
peduncular bract completely enclosing inflores-
cence, caducous; rachillae bearing spiral superfi-
cial triads basally and staminate flowers distally.
Staminate symmetrical; sepals and petals distinct;
stamens 12, filaments inflexed, anthers dorsifixed,
latrorse; pistillode equalling stamens. Staminodes
S-6. Stigmatic remains lateral; epicarp smooth,
mesocarp fibrous; endocarp vitreous, operculum
basal, round, beaked. Seed with round basal
hilum; endosperm homogeneous. Only one sp., C.
macrostachya Brongn., New Caledonia.
133. Moratia H.E. Moore
Moratia H.E. Moore, Gentes Herb. 12: 18 (1980); Moore &
Uhl., Allertonia 3: 313-402 (1984).
Solitary, erect. Crownshaft present. Leaflets
single-fold, acute. Inflorescences infrafoliar,
branched to 2-3 orders; prophyll and peduncular
bract entirely enclosing inflorescence in bud, ca-
ducous; rachillae bearing superficial spiral triads
proximally, distally bearing staminate flowers.
Staminate flowers symmetrical; sepals and petals
distinct; stamens 6, filaments inflexed, anthers
dorsifixed, oblong; pistillode equalling stamens.
Staminodes 3. Stigmatic remains lateral; epicarp
roughened, mesocarp with few fibers, endocarp
vitreous, faintly reticulate, operculum spathulate.
Seed with basal hilum. Endosperm homogeneous.
Eophyll bifid. Only one sp., M. cerifera H.E.
Moore, New Caledonia.
134. Clinostigma H. Wendl.
Clinostigma H. Wend!., Bonplandia 10: 196 (1862).
Solitary, erect. Crownshaft present. Leaflets
single-fold, acute. Inflorescences infrafoliar,
branched to 3 orders; prophyll and peduncular
bract enclosing inflorescence in bud, caducous;
rachillae bearing superficial spiral triads through-
out, or proximally only with staminate flowers
 Palmae
distally. Staminate flowers asymmetrical; sepals
and petals distinct; stamens 6, filaments inflexed,
anthers versatile, latrorse; pistillode conical.
Staminodes (5- )6. Stigmatic remains eccentrically
apical, lateral, or basal; epicarp smooth, mesocarp
fibrous, endocarp smooth, operculum elliptic.
Seed with basal or lateral, rounded to elongate
hilum; endosperm homogeneous. Eophyll bifid or
entire. About 13 spp., Bonin and Caroline Is. to
Samoa, Fiji Is., Vanuatu, Solomon Is., and New
Ireland.
135. Carpoxylon H. Wendl. & Drude
Carpoxylon H. Wend!. & Drude, Linnaea 39: 177 (1875); Dowe
& Uhl, Principes 33(2): 68-73 (1989).
Solitary, erect. Crownshaft present. Leaflets
single-fold, acute. Inflorescences infrafoliar,
branching to 3 orders; prophyll completely
sheathing; peduncular bracts 2, similar to
prophyll, deciduous; rachillae bearing spiral
superficial triads proximally, staminate flowers
distally. Staminate flowers asymmetrical; sepals
and petals distinct; stamens 6, filaments inflexed,
anthers oblong, dorsifixed, latrorse; pistillode
longer than stamens in bud. Staminodes 5 connate
in ring. Stigmatic remains eccentrically apical;
epicarp smooth, mesocarp thick, fibrous, en-
docarp thick, longitudinally ridged, operculum
basal, circular. Hilum elongate, endosperm
homogeneous, hollow, embryo basal. Only one
sp., C. macrospermum H. Wendl. & Drude,
Vanuatu.
136. Alsmithia H.E. Moore
Alsmithia H.E. Moore, Principes 26: 122 (1982).
Solitary, erect. Crownshaft absent. Leaflets single-
fold, acute. Inflorescences interfoliar, branching
to 2 orders; prophyll complete, short; peduncular
bract much exceeding prophyll, deciduous;
rachillae bearing spiral superficial triads proxi-
mally, staminate flowers distally. Staminate
flowers symmetrical; sepals and petals distinct;
stamens 6, filaments inflexed, anthers oblong,
dorsifixed, latrorse; pistillode shorter than
stamens. Staminodes 3. Stigmatic remains apical;
epicarp smooth, mesocarp fibrous, endocarp
fragile, with thickened adnate fibers, irregularly
sculptured, beaked, operculum basal, rounded.
Seed angled in cross-section, hilum elongate, en-
dosperm homogeneous, embryo basal. Only one
sp., A. longipes H.E. Moore, Fiji Islands.
371
137. Satakentia H.E. Moore
Satakentia H.E. Moore, Principes 13: 5 (1969).
Solitary, erect. Crownshaft present. Leaflets
single-fold, acute. Inflorescences infrafoliar,
branching to 2 orders; prophyll and peduncular
bract enclosing inflorescence in bud, deciduous;
rachillae bearing spiral superficial triads proxi-
mally, staminate flowers distally. Staminate
flowers asymmetrical; sepals and petals distinct;
stamens 6, filaments inflexed, anthers oblong,
dorsifixed, latrorse; pistillode equalling stamens.
Staminodes 3. Stigmatic remains eccentrically
apical; epicarp smooth, mesocarp fibrous,
endocarp smooth, operculum elongate. Hilum
elongate, endosperm homogeneous, embryo
basal. Eophyll bifid. Only one sp., S. liukiuensis
(Hatusima) H.E. Moore, Ryukyu Islands.
138. Rhopaloblaste Scheff.
Rhopaloblaste Scheff., Ann. Jard. Bot. Buitenzorg 1: 137
(1876).
Solitary or clustered, erect. Crownshaft present
or absent. Leaflets single-fold, slender, acute. In-
florescences infrafoliar, branched to 3 orders;
prophyll and peduncular bract enclosing inflores-
cence in bud, caducous. Staminate flowers sym-
metrical; sepals and petals distinct; stamens 6-9,
the filaments connate basally, inflexed, anthers
elongate, medifixed, latrorse; pistillode conical to
columnar. Staminodes 6. Stigmatic remains
apical; epicarp smooth, endocarp fragile, opercu-
lum rounded, basal. Hilum lateral; endosperm ru-
minate. Eophyll pinnate. Six spp., Nicobar Islands,
Malay Peninsula, and Singapore, the Moluccas,
New Guinea, and Solomon Is.
139. Dictyosperma H. Wendl. & Drude
Dictyosperma H. Wend!. & Drude, Linnaea 39: 181 (1875).
Solitary, erect. Crownshaft present. Leaflets
single-fold, acute. Inflorescences infrafoliar,
branched to 1 order, prophyll and peduncular
bract enclosing inflorescence in bud, deciduous;
rachillae bearing superficial triads proximally,
staminate flowers distally. Staminate flowers
asymmetrical; sepals and petals distinct; stamens
6, filaments, inflexed, anthers elongate, dorsifixed,
versatile, latrorse; pistillode equalling stamens.
Staminodes 3. Stigmatic remains apical; epicarp
smooth, mesocarp fibrous, endocarp smooth,
372
operculum basal, circular. Hilum elongate, lateral;
endosperm ruminate. Eophyll bifid. Only one sp.,
D. album (Bory) H. Wendl. & Drude, Mascarene
Is.
140. Actinorhytis H. Wendl. & Drude
Actinorhytis H. Wendl. & Drude, Linnaea 39: 184 (1875).
Solitary, erect. Crownshaft present. Leaflets
single-fold, acuminate or bifid. Inflorescences
infrafoliar, branching to 3 orders; prophyll and
peduncular bract enclosing inflorescence in bud,
deciduous; rachillae bearing spiral superficial
triads proximally, staminate flowers distally.
Staminate flowers asymmetrical; sepals and petals
distinct; stamens 24-33, filaments distinct, in-
flexed, anthers elongate, medifixed, versatile,
latrorse; pistillode columnar, equalling stamens in
bud. Staminodes 3. Stigmatic remains apical;
epicarp smooth, mesocarp fibrous, endocarp thin,
operculum circular. Hilum lateral, elongate, en-
dosperm ruminate, hollow. Eophyll bifid. Two
spp., New Guinea and Solomon Is.
141. Lavoixia H.E. Moore
Lavoixia H.E. Moore, Gentes Herb. 11: 296 (1978); Moore &
Uhl, Allertonia 3: 313-402 (1984).
Solitary, erect. Crownshaft present. Leaflets
single-fold, acute. Inflorescences infrafoliar,
branched to 2 orders; prophyll complete, short;
peduncular bract exceeding prophyll, caducous;
rachillae bearing superficial triads proximally,
staminate flowers distally. Staminate flowers sym-
metrical; sepals and petals distinct; stamens 6, fila-
ments distinct, not inflexed, anthers didymous;
pistillode columnar, slightly exceeding stamens.
Staminodes 3. Stigmatic remains lateral; epicarp
smooth, mesocarp thick, not fibrous, endocarp
thin. Seed laterally compressed; hilum basal;
endosperm homogeneous. Only one sp., L.
macrocarpa H.E. Moore, New Caledonia.
142. Alloschmidia H.E. Moore
Alloschmidia H.E. Moore, Gentes Herb. 11: 293 (1978); Moore
& Uhl, Allertonia 3: 313-402 (1984).
Solitary, erect. Crownshaft present. Leaflets
single-fold, acute. Inflorescences interfoliar,
branched to 3 orders; prophyll short, enclosing
peduncular bract; triads in pits throughout.
Staminate flowers symmetrical; sepals and petals
Palmae
distinct; stamens 6, filaments inflexed, anthers
dorsifixed, oblong, latrorse; pistillode columnar
exceeding stamens. Staminodes 3. Stigmatic
remains apical; epicarp smooth, mesocarp with
thin fibers, endocarp fragile, minutely pitted,
operculum elongate. Hilum elongate, endosperm
homogeneous; embryo lateral. Eophyll bifid. Only
one sp., A. glabrata (Becc.) H.E. Moore, northeast-
ern New Caledonia.
143. Cyphophoenix H. Wendl. ex J.D. Hook.
Cyphophoenix H.A. Wendl. ex J.D. Hook. in Benth. & Hook.,
Gen. Pl. 3: 893 (1883); Moore & Uhl, Allertonia 3: 313-402
(1984).
Solitary, erect. Crownshaft present. Leaflets
single-fold, acuminate or bifid. Inflorescences
infrafoliar, branched to 1-2(-3) orders; prophyll
complete, caducous; peduncular bract exceeding
prophyll, enclosing inflorescence in bud, cadu-
cous; rachillae bearing slightly sunken triads
proximally, staminate flowers distally. Staminate
flowers symmetrical, briefly pedicellate or sessile;
sepals and petals distinct; stamens 6, filaments
distinct, inflexed, anthers versatile, basifixed;
pistillode columnar. Staminodes 3. Stigmatic
remains apical; epicarp smooth, mesocarp
fibrous, endocarp crustaceous, smooth, opercu-
lum circular, basal. Hilum obovoid, apical;
endosperm homogeneous. Eophyll bifid. Two
spp., New Caledonia and Loyalty Is.
144. Campecarpus H. Wendl.
Campecarpus H. Wendl. ex Becc., Le Palme della Nuova
Caledonia: 28 (1920); Moore & Uhl, Allertonia 3: 313-402
(1984).
Solitary, erect. Crownshaft present; leaflets single-
fold, acute. Inflorescences infrafoliar, branched to
2 orders; prophyll incomplete, caducous; pedun-
cular bract exceeding prophyll, enclosing inflores-
cences in bud, caducous; triads superficial
proximally, staminate flowers distally. Staminate
flowers symmetrical; sepals and petals distinct;
stamens 6, filaments inflexed; anthers oblong,
dorsifixed, versatile; pistillode shorter than sta-
mens in bud. Staminodes 3. Stigmatic remains ec-
centrically apical; epicarp smooth, mesocarp
fleshy, with fibers, endocarp vitreous, with longi-
tudinal adaxial ridge, operculum rounded, basal.
Hilum elongate, endosperm homogeneous. Only
one sp., C. fulcitus (Brongn.) H. Wendl., New
Caledonia.
 Palmae
145. Basselinia Vieill.
Basselinia Vieill., Bull. Soc. Linn. Normandie II, 6: 230 (1873);
Moore & Uhl, Allertonia 3: 313-402 (1984).
Solitary or clustered, erect. Leaves entire bifid or
variously pinnate; leaflets single-multi-fold,
acute. Inflorescences infrafoliar, branched to 1 or
2 orders; prophyll incomplete; peduncular bract
exceeding prophyll; spiral triads superficial proxi-
mally, staminate flowers distally. Staminate
flowers symmetrical; sepals and petals distinct;
stamens 6, filaments inflexed, anthers dorsifixed,
latrorse; pistillode equalling stamens in bud.
Staminodes 3. Stigmatic remains lateral to subapi-
cal; epicarp smooth, mesocarp with sparse fibers,
endocarp vitreous, operculum rounded, basal.
Hilum basal, rounded, endosperm homogeneous.
Eophyll bifid. Eleven spp., New Caledonia.
146. Cyphosperma H. Wendl.
Cyphosperma H. Wend!. ex J.D. Hook. in Benth. &Hook., Gen.
Pl. 3: 895 (1883); Moore & Uhl, Allertonia 3: 313-402
(1984).
Solitary, erect. Crownshaft absent. Leaves entire-
bifid or variously pinnate; leaflet single- or multi-
fold, acute. Inflorescences interfoliar, branched to
2 orders; prophyll incomplete, short; peduncular
bract tubular, much longer than prophyll, mar-
cescent; spiral triads almost throughout, sunken
in pits. Staminate flowers symmetrical; sepals and
petals distinct; stamens 6, filaments inflexed,
anthers oblong, dorsifixed; pistillode columnar
overtopping stamens in bud. Staminodes 3. Stig-
matic remains lateral; epicarp smooth, mesocarp
fleshy, endocarp vitreous, irregularly sculptured,
ridged, grooved. Seed laterally attached, hilum
linear, endosperm homogeneous. Eophyll bifid.
Four spp., two in Fiji, one in New Caledonia, one
in Vanuatu.
147. Veillonia H.E. Moore
Veillonia H.E. Moore, Gentes Herb. 11: 299 (1978); Moore &
Uhl, Allertonia 3: 313-402 (1984).
Solitary, erect. Crownshaft present. Leaflets
single-fold, acute. Inflorescences infrafoliar,
branched to 1-2 orders; prophyll incomplete, ca-
ducous; peduncular bract tubular, much exceed-
ing prophyll; rachillae bearing spiral triads
proximally, staminate flowers distally. Staminate
flowers symmetrical; sepals and petals distinct;
stamens 6, filaments inflexed, anthers oblong,
373
dorsifixed, latrorse; pistillode columnar. Sta-
minodes 3. Stigmatic remains subapical; epicarp
papillate, mesocarp sparsely fibrous, endocarp
fragile, sculptured with adaxial ridge and 2 lateral
and 2 abaxial irregular crests and a dorsal groove,
operculum basal. Hilum elongate, endosperm ho-
mogeneous. Only one sp., V. alba H.E. Moore,
endemic to New Caledonia.
148. Burretiokentia Pic.Serm.
Burretiokentia Pic.Serm. in Becc. & Pic.Serm., Webbia 11: 122
(1955); Moore & Uhl, Allertonia 3: 313-402 (1984); Pintaud
& Hodel, Principes 42 (1998) in press.
Solitary, erect. Crownshaft present. Leaflets
single-fold, acute. Inflorescences infrafoliar,
branched to 1(-2) orders; prophyll short, incom-
plete, caducous; peduncular bract close to and
slightly exceeding prophyll, enclosing inflores-
cence in bud, caducous; spiral, superficial triads
proximally, staminate flowers distally. Staminate
flowers usually asymmetrical; sepals and petals
distinct; stamens 6, filaments inflexed, anthers
dorsifixed, pistillode columnar. Staminodes 3.
Stigmatic remains apical; epicarp smooth, meso-
carp sparsely fibrous, endocarp irregularly sculp-
tured, operculum basal, rounded. Hilum linear,
lateral, endosperm homogeneous. Eophyll bifid.
Five spp., New Caledonia.
149. Physokentia Becc.
Physokentia Becc. in Martelli, Atti Soc. Tosc. Sc. Nat. Pisa,
Mem. 44: 152 (1934); Moore, Principes 13: 120-136 (1969).
Goniocladus Burret, Notizbl. Bot. Gart. Mus. Berlin-Dahlem
15: 86 (1940).
Solitary, erect, often stilt-rooted. Crownshaft
present. Lamina variously pinnate, leaflets acute.
Inflorescences infrafoliar, branching to 2(3)
orders; prophyll incomplete, peduncular bract ex-
ceeding prophyll, enclosing inflorescence in bud,
caducous with prophyll; rachillae with superficial
triads proximally, staminate flowers distally.
Staminate flowers briefly pedicellate, asymmetri-
cal; sepals distinct; petals distinct or briefly
connate; stamens 6, filaments inflexed, anthers
versatile, latrorse; pistillode conspicuous. Sta-
minodes 3. Stigmatic remains apical; endocarp
angled or ridged, sculptured, with adaxial keel and
abaxial ridge, operculum rounded. Hilum narrow,
lateral, endosperm homogeneous or ruminate.
Eophyll bifid. Seven spp., Fiji Is., New Hebrides,
Solomon Islands, and New Britain.
374
40. Subtribe Oncospermatinae J.D. Hook. {1883).
Small to robust, spiny (at least when juvenile);
crownshaft present or not; leaves pinnate or pin-
nately ribbed; leaflets entire or praemorse; inflo-
rescences interfoliar or infrafoliar; triads rarely
borne in pits, usually superficial; staminate
flowers symmetrical or asymmetrical, sometimes
congenitally open; pistillate flowers with imbri-
cate petals with triangular valvate tips; gynoe-
cium pseudomonomerous; ovule hemianatropous
(?always); endocarp with a well-defined opercu-
lum covering embryo; embryo basal; germination
adjacent-ligular.
The Oncospermatinae include eight genera, five
of them found only in the Seychelles, two in the
Mascarenes and the eighth, Oncosperma, wide-
spread from Sri Lanka to Malesia. There is re-
markable diversity in floral and fruit morphology.
All genera are spiny, at least when they are juve-
niles, and it is this character alone that separates
the subtribe from the other subtribe with opercu-
late endocarps, the Iguanurinae.
KEY TO THE GENERA OF ONCOSPERMATINAE
1. Leaves with leaflets all 1-ribbed, acute or acuminate;
sheaths usually forming a conspicuous crownshaft; inflo-
rescence usually infrafoliar; peduncular bract enclosed by
the prophyll, both inserted close together, splitting and usu-
ally caducous at an thesis; peduncle short, usually ± equal-
ling the rachis; staminate flowers asymmetrical, sepals
acute
- Leaves with blades undivided, bifid, or irregularly to regu-
larly divided into leaflets, these usually with more than 1
rib, acute, bifid or praemorse; sheaths not forming a con-
spicuous crownshaft; inflorescences interfoliar or becom-
ing infrafoliar in age; peduncular bract caducous, inserted
some distance from the prophyll and exceeding it; prophyll
persistent; peduncle elongate, usually much longer than the
rachis; sepals rounded
2. Fruit ovoid with apical or eccentrically apical stigmatic
remains; perianth ± 1/2 as high as the fruit; seed with ho-
mogeneous endosperm; triads borne in 6 vertical rows, each
triad subtended by a saucerlike bract, the bracts obscured
by the staminate flowers, but becoming conspicuous when
the flowers have fallen 153. Tectiphiala
- Fruit ellipsoid to globose, with lateral to basal stigmatic
remains; perianth less than 1/2 as high as the fruit; en-
dosperm homogeneous or ruminate; rachilla bracts not
saucerlike
3. Fruit ellipsoid with basal stigmatic remains; staminate
flower open early in development with ca. 9 stamens,
scarcely exceeding the petals and a conspicuous, slender,
elongate, trifid pistillode 150. Deckenia
- Fruit globose to ellipsoid with lateral stigmatic remains;
staminate flower closed in bud, stamens 6-12; pistillode
shorter than stamens and petals
4. Stamens exserted at anthesis; pistillode minutely trifid; seed
with homogeneous endosperm 151. Acanthophoenix
- Stamens included at anthesis; pistillode deeply trifid; seed
with ruminate endosperm 152. Oncosperma
Palmae
5. Staminate flowers with petals about twice as long as the
sepals; stamens 6; pistillode large, truncate, 3-angled and
lobed, about as high as the petals; leaflets or blade
praemorse 6
- Staminate flower with petals about 4 times as long as the
sepals; stamens 18 or more; pistillode not truncate, angled
or lobed, leaflets or blade lobes acute or acuminate 7
6. Leaf blade almost appearing undivided, the leaflets scarcely
separated; fruit large, 2-2.5cm diam., the endocarp con-
spicuously ridged and crested; seed ± ridged, raphe
branches anastomosing 154. Verschaffeltia
- Leaflets clearly separated; fruit small, ca. 5 mm diam. or
less; endocarp not ridged; seed not ridged, the raphe
branches few, ascending 155. Roscheria
7. Leaf blade usually undivided though lobed marginally, the
lobes bifid; inflorescence branching to 2 orders; staminate
flower asymmetrical; stamens ca. 18, the filaments tapered,
anthers ± versatile; pistillode small, slender; fruit ovoid
156. Phoenicophorium
- Leaf pinnate, with rather distant, mostly 2- to 3-ribbed
acute or acuminate leaflets; inflorescence branching to 1
order; staminate flower symmetrical; stamens 40-50, the
filaments expanded distally, anthers not versatile; pistillode
ovoid, minutely trifid; fruit subglobose 157. Nephrosperma
150. Deckenia H. Wendl. ex Seem.
Deckenia H. Wend!. ex Seem., Gard. Chron. 1870: 561 (1870).
Solitary, spiny when juvenile, becoming unarmed
at maturity; crownshaft present; leaflets single-
fold, acuminate. Inflorescences infrafoliar,
branching to 2 orders protandrous; prophyll and
peduncular bract similar, inserted close together,
caducous; rachillae coiled in bud, bearing spiral
2
triads throughout. Staminate flowers asymmetri-
cal, congenitally open; stamens 6-9, anthers short,
medifixed, latrorse; pollen sulcate, exine intectate,
clavate; pistillode columnar, longer than petals.
Staminodes 6, minute. Fruit narrow-ovoid, stig-
matic remains basal; operculum circular. Seed
5 basally attached, hilum circular; endosperm ho-
mogeneous; embryo basal. Eophyll bifid. Only one
sp., D. nobilis H. W endl., Seychelles Is.
151. Acanthophoenix H. Wendl.
Acanthophoenix H. Wend!., Fl. Serres Jardins de !'Europe 16:
181 (1867).
3 Solitary, sparsely spiny at maturity; crownshaft
present; leaflets single-fold, acuminate. Inflores-
cences infrafoliar, branching to 2 orders,
protandrous; prophyll and peduncular bract simi-
lar, inserted close together, caducous; rachillae
sinuous, bearing spiral triads throughout. Stami-
4 nate flowers asymmetrical, closed in bud; stamens
6-12, anthers linear, dorsifixed, latrorse; pollen
sulcate, exine tectate, reticulate; pistillode ovoid,
short. Staminodes 6-9, minute. Fruit ellipsoid,
 Palmae
stigmatic remains lateral to subapical; operculum
circular. Seed laterally attached, hilum circular;
endosperm homogeneous; embryo basal. Eophyll
bifid or pinnate. Only one sp., A. rubra H. Wendl.,
Mauritius and Reunion.
152. Oncosperma Blume
Oncosperma Blume, Bull. Sc. Phys. Nat. Neerl. 1: 64 (1838).
Caespitose, spiny throughout; crownshaft present;
leaflets single-fold, acuminate. Inflorescences
infrafoliar, branching to 2 orders, protandrous;
prophyll and peduncular bract similar, inserted
close together, caducous; rachillae sinuous,
bearing spiral triads ± throughout. Staminate
flowers asymmetrical, closed in bud; stamens 6-9,
anthers linear, dorsifixed, latrorse; pollen sulcate,
exine tectate with supratectal processes; pistillode
short, trifid. Staminodes 6, minute. Fruit globose,
stigmatic remains lateral to subapical; operculum
circular. Seed laterally attached, hilum elongate;
endosperm ruminate; embryo subbasal. Eophyll
bifid. Five spp., one endemic to Sri Lanka, two
endemic to the Philippines, and two widespread in
SE Asia and W Malesia, reaching Sulawesi and
Philippines.
153. Tectiphiala H.E. Moore
Tectiphiala H.E. Moore, Gentes Herb. 11(4): 285 (1978).
Solitary or caespitose, spiny throughout; crown-
shaft present; leaflets single-fold, acuminate.
Inflorescences infrafoliar, branching to 1 order,
protandrous; prophyll and peduncular bract
similar, inserted close together, caducous;
rachillae few, straight, bearing conspicuous,
spiral, saucerlike bracts and triads ± throughout.
Staminate flowers asymmetrical, closed in bud;
stamens 6-9, filaments coiled; anthers linear,
dorsifixed, latrorse; pollen monosulcate, exine
tectate, reticulate; pistillode short, trifid.
Staminodes 6, minute. Fruit ovoid, stigmatic re-
mains apical; operculum circular. Seed laterally
attached, hilum elongate; endosperm homoge-
neous; embryo subbasal. Only one sp., T. ferox
H.E. Moore, endemic to Mauritius.
154. Verschaffeltia H. Wendl.
Verschaffeltia H. Wend!., Illus. Hortic. 12 (Misc.): 5 (1865).
Solitary, stilt-rooted. Crownshaft absent. Lamina
undivided, lobed at margins, praemorse, irregu-
375
larly tattering with age. Inflorescences interfoliar,
branching to 2-3 orders, protandrous; prophyll
inserted above base, persistent; peduncular
bract similar; rachillae sinuous with spiral triads
± throughout. Staminate flowers symmetrical,
closed in bud; stamens 6, anthers rounded,
medifixed, latrorse; pollen sulcate, exine tectate,
reticulate; pistillode large, truncate. Fruit spheri-
cal, stigmatic remains basal; endocarp flanged
and ridged, operculum rounded, basal. Seed
basally attached, hilum rounded, endosperm
ruminate; embryo basal. Eophyll bifid. Only
one sp., V. splendida H. Wendl., endemic to
Seychelles.
155. Roscheria H. Wendl. ex l.B. Balf.
Roscheria H. Wend!. ex LB. Balf. in ].G. Baker, Fl. Maur.
Seych.: 386 (1877).
Solitary. Crownshaft present. Leaves irregularly
pinnate, segments acuminate or praemorse. In-
florescences interfoliar, becoming infrafoliar,
branching to 3(4) orders, prophyll inserted above
base, persistent, peduncular bract exceeding
prophyll, caducous; rachillae flexuous, bearing
spirally arranged triads proximally, staminate
flowers distally. Staminate flowers symmetrical,
closed in bud; stamens 6, anthers rounded,
medifixed, latrorse; pollen sulcate, exine tectate,
reticulate; pistillode large, truncate. Staminodes 6.
Fruit globose, stigmatic remains subbasal; en-
docarp smooth, operculum basal. Seed basally
attached, hilum rounded, endosperm ruminate;
embryo basal. Only one sp., R. melanochaetes (H.
Wendl.) H. Wendl. ex l.B. Balf., confined to Mahe
and Silhouette Is. in the Seychelles.
156. Phoenicophorium H. Wendl.
Phoenicophorium H. Wend!., Illus. Hortic. 12 (Misc.): 5 (1865).
Solitary, ± unarmed at maturity. Crownshaft
absent. Lamina entire-bifid, marginally lobed. In-
florescences interfoliar, branching to 1-2 orders,
protandrous; prophyll inserted above base, persis-
tent, peduncular bract included within prophyll;
rachillae flexuous, bearing spiral triads, except
distally staminate. Staminate flowers asymmetri-
cal, closed in bud; stamens 15-18, anthers elon-
gate, medifixed, latrorse; pollen sulcate, exine
tectate, spiny; pistillode absent. Staminodes 6.
Fruit ellipsoid; endocarp thin, operculum round,
basal. Seed basally attached, hilum rounded; en-
dosperm ruminate; embryo basal. Eophyll entire.
376
Only one sp., P. borsigianum (K. Koch) Stuntz,
widespread in the Seychelles Is.
157. Nephrosperma LB. Balf.
Nephrosperma I.B. Balf. in J.G. Baker, Fl. Maur. Seych.: 386
(1877).
Solitary, ± unarmed at maturity. Crownshaft
absent. Leaflets 2-3-fold, acute. Inflorescences
interfoliar, branching to 1 order, protandrous;
prophyll near base, persistent, peduncular bract
inserted near prophyll, deciduous; rachillae
bearing spiral triads, except near tips, where
staminate. Staminate flowers symmetrical, closed
in bud; stamens 40-50, anthers rounded,
basifixed, latrorse; pollen sulcate, exine tectate,
reticulate, pistillode ovoid. Staminodes 6. Fruit
globose; operculum round, basal. Seed globose to
kidney-shaped, laterally attached near base, hilum
oblong; endosperm ruminate; embryo basal.
Eophyll bifid. Only one sp., N. vanhoutteanum (H.
Wendl. ex Van Houtte) LB. Balf., confined to the
Seychelles Is.
4p. Subtribe Sclerospermatinae J. Dransf. &
N. Uhl {1986).
Acaulescent, unarmed, pleonanthic, monoecious;
crownshaft absent; leaves large, irregularly
divided or entire-bifid, discolorous, margins prae-
morse; inflorescence interfoliar, spicate; prophyll
short; peduncular bract disintegrating into a
fibrous net, partly covering flowers; flowers in
triads near base of spike, paired staminate
flowers distally. Staminate flowers with 3 imbri-
cate sepals and 3 valvate petals; stamens ca. 60;
pistillate flower with connate sepals and imbricate
petals; gynoecium pseudomonomerous. Fruit
with apical stigmatic remains. Endosperm
homogeneous, hollow; germination remote-
tubular.
As originally circumscribed, this subtribe also
included Marojejya, but now that the latter and
Masoala are better known, it seems very unlikely
that Marojejya is related to Sclerosperma. As de-
limited here, the subtribe contains the single
genus, Sclerosperma, a most remarkable, aberrant
palm of West Africa.
158. Sclerosperma G. Mann & H. Wendl.
Fig. 82J
Sclerosperma G. Mann & H. Wend!., Trans. Linn. Soc. London
24: 427 (1864).
Palmae
Clustering. Petiole elongate. Inflorescences con-
cealed among the leaf bases; peduncular bract
longer than the prophyll. Staminate flowers asym-
metrical, sessile or stalked; stamens with short
filaments, anthers elongate, basifixed, latrorse; pol-
len triangular, triporate, exine tectate, reticulate;
pistillode lacking. Staminodes 6. Stigma 3-angled;
ovule pendulous, probably campylotropous. Fruit
globose, beaked; endo carp bony, irregularly pitted
externally, with basal pore region. Seed globose,
hilum elongate, embryo basal. Eophyll bifid.
Three(?) spp., rainforest in equatorial Africa.
4q. Subtribe Masoalinae Beentje & J. Dransf.
{1996).
Robust, solitary, unarmed, pleonanthic, monoe-
cious, short-stemmed, litter-trapping palms;
leaves large; crownshaft not developed; inflores-
cences interfoliar, staminate, pistillate, or bi-
sexual; prophyll hidden by leaf bases; peduncular
bract similar to prophyll, numerous incomplete
peduncular bracts also present; stamens 6, ovary
pseudomonomerous, with 3 large triangular
stigmas; fruit beaked, with apical or lateral stig-
matic residue; endosperm homogeneous; germi-
nation adjacent-ligular; eophyll bifid.
The genera of the Masoalinae comprise massive
short-stemmed tree palms of the understory of
rainforest in Madagascar. The petioles of the
massive leaves are short or absent, the leaf blades
acting as funnels that trap litter in the crown of the
palm. The relationships of the two genera with
other arecoid palms are as yet obscure. The over-
all structure of the unisexual inflorescences of
Marojejya forms a remarkable parallel with that
of Elaeis in Tribe Cocoeae.
KEY TO THE GENERA OF MASOALINAE
1. Inflorescences bisexual, branching to 2 orders, elongate,
lax, long exserted from leaf sheath; peduncle elongate;
rachis longer than peduncle; staminate petals free;
pistillode as large as stamens; staminate sepals broadly im-
bricate; fruit symmetrical with apical stigmatic remains
159. Masoala
- Inflorescence unisexual, congested, ± hidden among leaf
bases, branching to 1 order; peduncle short; rachis shorter
than peduncle; staminate flowers with petals connate in
basal1/3 and adnate to receptacle; pistillode minute; sepals
narrow in staminate flower; fruit asymmetrical, with lateral
stigmatic remains 160. Marojejya
159. Masoala Jum.
Masoala Jum., Ann.lnst. Bot. Geol. Col. Marseille V, 1(1): 8
(1933); J. Dransf. & Beentje, Palms Madag.: 20-27 (1995).
 Palmae
Lamina irregularly or regularly divided, leaflets
single-fold, acuminate. Inflorescences branching
to 1-2 orders; peduncle elongate; prophyll in-
serted above base, peduncular bract similar to
prophyll, caducous; rachis longer than peduncle;
rachillae bearing spirally arranged, sunken triads,
subtended by coriaceous bracts, through most of
rachilla length. Staminate flowers symmetrical;
anthers elongate, basifixed, latrorse; pollen
sulcate, exine tectate, rugulate; pistillode colum-
nar. Staminodes absent. Fruit ovoid. Seed smooth.
Two spp., eastern Madagascar.
160. Marojejya Humb.
Marojejya Humb., Mem. Inst. Sc. Madagascar B. Biol. Veg. 6:
92 (1955); J. Dransf. & Beentje, Palms Madag.: 28-37 (1995).
Lamina regularly divided, leaflets single-fold,
acuminate, or entire-bifid to irregularly divided.
Inflorescences unisexual, branching to 1 order;
peduncle short; prophyll near base, peduncular
bract similar to prophyll; rachis shorter than pe-
duncle; rachillae bearing spirally arranged bracts
forming pits, with solitary pistillate or paired
staminate flowers. Staminate flowers ± asym-
metrical; anthers elliptic, medifixed, versatile,
latrorse; pollen sulcate, exine tectate, foveolate;
pistillode small. Staminodes 6. Fruit asymmetri-
cally globular; endocarp thick. Seed irregularly
shaped, smooth or grooved. Two spp., eastern
Madagascar.
5. Tribe Cocoeae Mart. (1837).
Pleonanthic, monoecious palms with reduplic-
ately pinnate leaves; crownshaft absent; inflores-
cences solitary, interfoliar, usually branched to 1
order only; prophyll short, usually fibrous; pedun-
cular bract much exceeding the prophyll, usually
woody and persistent, enclosing entire inflores-
cence in bud, splitting at anthesis; staminodes al-
most always connate in a short irregularly toothed
ring; gynoecium with 3 or more locules and 3 or
more ovules; endocarp well developed with 3 or
more pores.
KEY TO THE GENERA OF COCOEAE
1. Unarmed palms except for the sometimes sharply toothed
petiole margin; corolla of pistillate flowers always of
broadly imbricate petals; endocarp with pores above, at, or
below the middle 2 the terminal portion with dense staminate flowers;
- Prickly palms, armed in some or all parts with soft to
mostly stout spines, or when rarely unarmed (Bactris
377
spp.), then the perianth of pistillate flowers with connate
petals; endocarp with pores at or above the middle except
rarely in Acrocomia, the pores more or less deeply im-
pressed in the endocarp, usually covered by or plugged
with fibers adherent to the endocarp, lacking a clearly de-
marcated and visible operculum; fruit always (?) 1-seeded;
endosperm always homogeneous. 15
2. Pistillate flowers not or only slightly sunken in the
rachillae; endocarp with pores at or below the middle;
peduncular bract woody. 3
- Pistillate flowers deeply sunken in the rachillae; endocarp
with pores at or above the middle; peduncular bract
fibrous 14
3. Peduncular bract borne at the tip of the peduncle, circum-
scissile and deciduous at anthesis; stamens 18-21, the fila-
ments connate in a torus. Madagascar
161. Beccariophoenix
- Peduncular bract borne near the prophyll insertion or
higher, but not at the tip, not circumscissile, splitting
abaxially but not deciduous at anthesis, persistent or
marcescent; stamens 3-many, filaments variously free
or connate 4
4. Peduncular bract nearly smooth or finely striate but not
deeply grooved; pores at or below the middle, not im-
pressed in the endocarp, each with a clearly demarcated
thin operculum 5
- Peduncular bract shallowly to deeply grooved; pores
mostly near the base of and usually impressed in the en-
docarp, often covered by or plugged with fibers, some-
times with a clearly demarcated operculum 7
5. Staminate flowers with 6 stamens; petioles usually con-
spicuously toothed on the margin toward the base; fruit
1-3-seeded. Brazil to Paraguay 162. Butia
- Staminate flowers with more than 6 stamens; petioles not
toothed along the margin 6
6. Staminate flowers stalked, the sepals not imbricate; sta-
mens 15-30; leaflets clustered; trunks always solitary, mas-
sive, up to 1m in diameter or more. Coastal Central Chile
163./ubaea
- Staminate flowers not stalked; sepals imbricate, stamens
8-16; trunks slender and often multiple. SE Africa
164. ]ubaeopsis
7. Inflorescences not normally differentiated into androgy-
nous and staminate but instead the pistillate flowers borne
at the base of, in the lower part of, or nearly throughout the
rachillae or the spike, the staminate flowers lateral to the
pistillate and usually pedicellate in the triads, sessile and
paired or solitary in the upper part of the rachilla or spike;
pistillate flowers always trilocular and triovulate 8
- Inflorescences normally of more than one kind on the
same plant, androgynous and staminate, or sometimes
also pistillate; androgynous inflorescences with the pistil-
late flowers large and few at the base of short rachillae, the
pistillate flowers accompanied by lateral or abortive stami-
nate flowers and usually with some paired or solitary
staminate flowers on the distal portion of the rachillae;
staminate inflorescences with longer rachillae bearing
staminate flowers only; pistillate flowers sometimes with
more than 3 carpels, ovules, and stigmas. Central and S
America 172. Attalea
8. Triads separated on the inflorescence; sepals of staminate
flowers rarely as long as the petals 9
- Triads densely arranged on the lower portion of a spike,
bracteoles prominent; sepals of staminate flowers linear,
acute to acuminate, more than 1/2 as long as the petals, the
378 Palmae
flowers rather markedly asymmetrical, lateral to the
densely packed pistillate flowers on the lower portion, and
densely crowded on the terminal portion of the spike;
stamens 6-120 13
9. Stamens more than 6 10
- Stamens 6 11
10. Stamens ca. 15; endocarp externally irregularly sculptured
and roughened and with 3 prominent crests at the apex.
Ecuador and Bolivia 168. Parajubaea
- Stamens 12; endocarp externally deeply grooved, inter-
nally with irregular swellings penetrating the seed. Mada-
gascar 169. Voanioala
11. Pistillate flowers very large, globose-ovoid, the sepals and
petals, rounded, without valvate apices, imbricate; stami-
nate flowers with distinct sepals and 6 stamens; fruit very
large, up to 25 em long or more at maturity, with a very
thick, fibrous mesocarp and thick, bony endocarp; seed
normally 1, with liquid endosperm when young and a
large, hollow interior when completely mature. Old World
tropics, now pantropical 165. Cocos
- Pistillate flowers ovoid or conic-ovoid, the sepals acute
and more or less hooded, the petals usually with conspicu-
ously valvate apices 12
12. Leaflets densely white or pale brown tomentose abaxially,
very narrow, close, and regularly arranged; epicarp and
mesocarp splitting regularly and longitudinally from the
apex to the base, into 3 sections at maturity, exposing the
thin endocarp, anthers versatile, medifixed, filaments in-
flexed at tip 167. Lytocaryum
- Leaflets lacking a dense covering of tomentum abaxially
(very rarely thin indument present but then leaflets
grouped); epicarp and mesocarp not splitting at maturity;
endocarp very thick, variously beaked, ridged, minutely
pitted or invaginated; anthers only rarely versatile. S
America and Lesser Antilles 166. Syagrus
13. Plants usually without emergent stem; stamens 6-19; seed
with homogeneous endosperm. Brazil, Paraguay
170. Allagoptera
- Plants with prominent trunk; stamens 60-120; seed with
ruminate endosperm. Brazil 171. Polyandrococos
14. Flowers of both sexes consistently on the same inflores-
cence, the pistillate near the bases of rachillae; inflores-
cence long-peduncled, laxly branched. NE Brazil
173. Barcella
- Flowers normally in separate inflorescences on the same
plant; inflorescence short-peduncled, densely branched.
Costa Rica to Brazil (E. oleifera), W Africa (E. guineensis)
174. Elaeis
15. Petals of pistillate flowers distinct and broadly imbricate,
or if sometimes partially connate basally and with an
adnate staminodal tube, then at least the margins free and
imbricate; stamen-filaments inflexed at the apex in bud,
the anthers dorsifixed and versatile; fruit with abundant
short fibers in the mesocarp, these strongly adherent to the
smooth or only very shallowly pitted endocarp. Cuba and
Mexico to Argentina 175. Acrocomia
- Petals of pistillate flowers connate 1/3-1/2 of their length
in a campanulate tube with prominent spreading or erect,
valvate lobes, or more than 112 of their length in an ur-
ceolate, briefly 3-lobed, 3-toothed, or even truncate tube;
fruit lacking abundant short fibers adherent to the en-
docarp 16
16. Pistillate petals connate 1/3-1/2 of their length in a cam-
panulate tube with prominent valvate lobes; staminodes
connate and adnate to the corolla tube basally, but distinct
or continued in a free, 3-6-lobed or -toothed or truncate
tube sometimes nearly equalling the stigmas above; the
basifixed anthers often sagittate basally; pistillode evident
17
- Pistillate petals connate beyond the middle or completely
connate in a 3-lobed, 3-toothed, or truncate, urceolate or
tubular corolla, the lobes, when developed, not spreading;
staminodes distinct or united in a short tube, but not
adnate to the corolla; pistillode usually lacking 18
17. Sepals of pistillate flowers distinct and imbricate; sepals of
staminate flowers distinct (except where united to the flo-
ral receptacle) and separated or imbricate; lobes of pistil-
late corolla spreading at an thesis; pinnae variously oblique
or truncate, broad, and strongly toothed at the apex;
trunks essentially uniform in diameter. Lesser Antilles, S
America from Brazil to Peru and Bolivia 177. Aiphanes
- Sepals of both pistillate and staminate flowers connate in
cupular calyces; lobes of pistillate corolla erect; pinnae
acute or acuminate; trunk markedly ventricose. Cuba
176. Gastrococos
18. Staminate flowers often associated with the pistillate in
triads basally or along the upper part of the rachilla but
above these triads, then paired or generally solitary and
densely aggregated in a distinct terminal portion of the
rachilla, each pair of flowers or each flower subtended by a
prominent bractlet adnate to or coherent with adjacent
bractlets to form a cupule sometimes as high as the flow-
ers; stamen filaments inflexed at the apex in bud; anthers
dorsifixed, versatile. Mexico to Brazil 180. Astrocaryum
- Staminate flowers not densely aggregated in a distinct ter-
minal portion of the rachilla but associated with the pistil-
late in triads or irregularly interspersed among the triads
and subtended by short, distinct bracteoles 19
19. Erect plants; upper pinnae not modified into reflexed
climbing organs; stamen filaments inflexed at the apex or
from nearly the middle in bud; anthers mostly dorsifixed,
versatile; peduncular bract inserted near the prophyll at
the base of the peduncle. Cuba and Mexico to Brazil
178. Bactris
- Climbing plants; upper pinnae usually very distant and
modified into reflexed hooklike climbing organs
(acanthophylls); stamen filaments erect in bud, short; an-
thers basifixed, erect, sagittate basally; peduncular bract
often inserted above the middle of the peduncle. Mexico to
Brazil 179. Desmoncus
Sa. Subtribe Beccariophoenicinae J. Dransf. &
N. Uhl (1986).
Peduncular bract borne at the tip of the peduncle
(always), very thick and woody, beaked, grooved,
circumscissile and deciduous at anthesis leav-
ing collar-like scar; rachis very short; rachillae
with basal pulvinus; stamens 18-21; gynoecium
trilocular, triovulate; fruit 1-seeded; endocarp
with 3 basal pores; endosperm ruminate; embryo
lateral near the base; germination adjacent-
ligular; eophyll entire.
The sole member of the subtribe is a remarkable
palm from Madagascar. Most distinctive is the
peduncular bract, which is very thick and woody,
borne at the base of the inflorescence rachis and
circumscissile at anthesis, leaving a broad collar-
like scar.
 Palmae
161. Beccariophoenix Jum. & H. Perrier
Beccariophoenix Jum. & H. Perrier, Ann. Fac. Sc. Marseille
23(2): 34 (1915); Dransf. & Beentje, Palms Madag.: 438-443
(1995).
Robust, solitary, tree. Leaflets single-fold. Pro-
phyll fibrous, persistent; rachillae spirally ar-
ranged, spreading, bearing distichous triads
throughout except near tips, where staminate.
Staminate flowers subsymmetrical; sepals 3, dis-
tinct; petals 3, distinct; anthers elongate, basifixed;
pollen sulcate, exine tectate; pistillode absent.
Pistillate flowers slightly larger than staminate;
sepals 3, distinct, imbricate; petals 3, distinct,
imbricate, with short valvate tips. Fruit ovoid
with pyramidal beak, epicarp smooth, mesocarp
easily separated from endocarp. Seed attached
near the base, hilum broad. Only one sp., B.
madagascariensis Jum. & H. Perrier, eastern
Madagascar.
Sb. Subtribe Butiinae Saakov {1954).
Monoecious, unarmed or armed with fiber spines
on petiole margins, epidermal emergence spines
lacking; leaf sheath distintegrating into an inter-
woven mass of fibers; inflorescences solitary,
interfoliar, protandrous; prophyll short; pedun-
cular bract inserted near the prophyll, much
longer, tubular, enclosing the inflorescence in
bud; rachillae or spikes bearing spirally arranged
triads proximally, paired or solitary staminate
flowers distally; gynoecium trilocular, triovulate,
ovules hemianatropous or anatropous, laterally
attached; seed basally attached.
This subtribe includes the familiar coconut.
Members are predominantly South American but
there is one genus in Madagascar, one in South
Africa, and the coconut itself is probably western
Pacific in origin. Until about 500 years ago another
probable member of the subtribe, Paschalococos
J. Dransf., known from subfossil endocarps,
occurred on Easter Island (Dransfield 1991b). Fos-
sil cocoid fruit, but not assignable to subtribe,
have been recorded for New Zealand and India.
The sub tribe is distinguished by the bisexual inflo-
rescences, the persistent peduncular bract, and the
absence of epidermal emergence spines.
162. Butia (Becc.) Becc. Fig. 81N
Butia (Becc.) Becc., Agric. Col. 10: 489 (1916); Henderson et
al., Field Guide: 136-138 (1995).
Cocos subgenus Butia Becc. (1887).
379
Solitary, acaulescent or erect. Petiole usually
armed with coarse fiber spines; leaflets variously
lobed or acute. Inflorescence branched to 1 order;
peduncular bract, smooth or striate. Staminate
flowers sessile or pedicellate, asymmetrical; sepals
3, distinct or basally connate; petals 3, distinct;
stamens 6, filaments inflexed, anthers elongate,
medifixed, introrse; pollen sulcate or trichotomo-
sulcate, exine tectate. Staminodal ring collarlike.
Fruit 1-3-seeded, beaked; epicarp smooth, en-
docarp smooth, thick, with 1-3 developed cavities,
pores lateral below equator or subbasal. En-
dosperm homogeneous, solid. Eight spp., cooler,
drier areas of S America, in southern Brazil, Para-
guay, Uruguay, and Argentina.
163. ]ubaea Kunth
jubaea Kunth in H.B.K., Nov. Gen. Sp. Pl. 1: quarto edition
308; folio edition 247 (1816) ("1815"); Uhl & Dransf., Gen.
Palm.: 487-489 (1987).
Solitary, massive tree. Petiole unarmed; leaflets
single-fold, tips acute or hooked. Inflorescence
branched to 1 order; peduncular bract, smooth.
Staminate flowers stalked, asymmetrical; sepals 3,
basally connate; petals 3, distinct; stamens 18, fila-
ments inflexed, anthers rectangular, medifixed,
latrorse; pollen sulcate, exine tectate, foveolate.
Staminodal ring forming a low collar. Fruit
1-seeded, beaked; epicarp smooth, endocarp
smooth, thick, with 3 low crests, pores lateral be-
low equator. Endosperm homogeneous, hollow.
Germination adjacent-ligular; eophyll entire. Only
one sp., f. chilensis (Molina) Baill., Central Chile.
164. ]ubaeopsis Becc.
jubaeopsis Becc., Webbia 4: 171 (1913); Uhl & J. Dransf., Gen.
Palm.: 489-491 (1987).
Caespitose, stems also branching aerially by
forking. Petiole unarmed; leaflets single-fold,
tips 2-lobed. Inflorescence branched to 1 order;
peduncular bract striate but not grooved. Stami-
nate flowers sessile, asymmetrical; sepals 3, dis-
tinct; petals 3, distinct; stamens 7-16, filaments
inflexed, anthers linear, dorsifixed, introrse; pol-
len sulcate or trichotomosulcate, exine tectate.
Staminodal ring forming a low collar. Fruit 1-
seeded, beaked; epicarp smooth, endocarp thick,
with 3 vertical grooves, pores lateral below equa-
tor. Endosperm homogeneous, hollow. Germina-
tion remote-tubular; eophyll entire. Only one sp.,
]. caffra Becc., S. Africa.
380
165. Cocos L.
Cocos L., Sp. Pl.: 1188 (1753); Harries, Bot. Rev. 44: 265-319
(1978).
Solitary tree. Petiole unarmed; leaflet tips asym-
metrical. Inflorescence branched to 1 order;
peduncular bract shallowly grooved. Staminate
flowers sessile, asymmetrical; sepals 3, distinct;
petals 3, distinct; stamens 6, filaments not in-
flexed, anthers linear, medifixed, latrorse; pollen
sulcate or trichotomosulcate, exine tectate, reticu-
late. Pistillate flower very large; staminodal ring
not lobed. Fruit 1-seeded, not beaked; epicarp
smooth, mesocarp very thick, fibrous, endocarp
with 3 longitudinal ridges, pores basal. En-
dosperm homogeneous, hollow, partially filled
with fluid. Germination adjacent-ligular; eophyll
entire. Only one sp., C. nucifera L., western Pacific,
widely cultivated throughout the tropics.
166. Syagrus Mart.
Syagrus Mart., Palmarum Familia: 18 (1824); Glassman, Illi-
nois Biol. Monographs: 56 (1987).
Solitary or clustered, acaulescent to trees. Petiole
rarely armed; leaflet tips various. Inflorescence
branched to 1 order, rarely spicate; peduncular
bract grooved. Staminate flowers sessile, rarely
stalked, asymmetrical; sepals 3, distinct or con-
nate; stamens 6, filaments not inflexed, anthers
linear, latrorse; pollen sulcate or trichotomo-
sulcate, exine tectate. Staminodal ring 6-lobed.
Fruit 1-seeded, sometimes beaked; epicarp
smooth or striate; endocarp smooth or variously
beaked, ridged, rarely with internal swellings,
pores basal or sub-basal. Endosperm homoge-
neous or ruminate. Germination adjacent-ligular
or remote-tubular; eophyll entire. About 30 spp., S
America and Lesser Antilles.
167. Lytocaryum Toledo
Lytocaryum Toledo, Arquivos Bot. Est. Sao Paulo II,. 2(1): 6.
(1944).
Glaziova Mart. ex Drude (1881) (non Bureau 1868).
Solitary, short-stemmed. Petiole unarmed; leaflets
acuminate, discolorous. Inflorescence branched
to 1 order; peduncular bract deeply grooved.
Staminate flowers sessile or pedicellate, symmetri-
cal; sepals 3, distinct or basally connate; petals 3,
distinct; stamens 6, filaments inflexed, anthers
elongate, medifixed, latrorse; pollen sulcate or
trichotomosulcate, exine tectate. Staminodal ring
Palmae
irregularly 6-toothed. Fruit 1-seeded, beaked; epi-
carp smooth, splitting with mesocarp along 3 lines
exposing smooth, thin endocarp, pores lateral
near base. Endosperm homogeneous or ruminate,
hollow or solid. Germination adjacent-ligular;
eophyll pinnate. Two spp., SE Brazil.
168. Parajubaea Burret
Parajubaea Burret, Notizb. Bot. Gart. Mus. Berlin-Dahlem 11:
48 (1930); Moraes & Henderson, Brittonia 42: 92-99 (1990).
Solitary trees. Petiole unarmed; leaflets bifid or
oblique. Inflorescence branched to 1(2) orders; pe-
duncular bract deeply grooved. Staminate flowers
sessile, asymmetrical; sepals 3, distinct; petals 3,
distinct; stamens 15, filaments inflexed, anthers
linear, dorsifixed, introrse; pollen sulcate, exine
tectate, punctate. Staminodal ring 3-lobed. Fruit 1-
seeded, beaked; epicarp smooth, endocarp thick,
irregularly sculptured, 3-ridged or not, pores
basal, sunken. Endosperm homogeneous, hollow.
Three spp., Ecuador, Bolivia, and Colombia.
169. Voanioala J. Dransf.
Voanioala J. Dransf., Kew Bull. 44: 192 (1989); Dransf. &
Beentje, Palms Madag.: 444-450 (1995).
Solitary tree. Petiole unarmed; leaflets bilobed.
Inflorescence branched to 1 order; pedunculat
bract deeply grooved. Staminate flowers sessile,
asymmetrical; sepals 3, distinct; petals 3, distinct;
stamens 12-13, filaments not inflexed, anthers
linear, basifixed, latrorse; pollen sulcate, exine
intectate with granular processes and baculae.
Staminodal ring irregularly 9-toothed. Fruit 1-
seeded, beaked; epicarp with scaly indument,
endocarp thick, deeply grooved, internally with
irregular rounded protrusions; pores lateral below
equator. Endosperm homogeneous, with a very
narrow hollow. Germination remote-tubular;
eophyll entire. Only one sp., V. gerardii J. Dransf.,
Madagascar.
170. Allagoptera C.G.D. Nees
Allagoptera C.G.D. Nees in Wied-Neuwied, Reise nach
Brasilien 2:335 (1821); Moraes, Fl. Neotrop. 73: 1-34 (1996).
Caespitose, acaulescent, stems also branching ae-
rially by forking. Petiole unarmed; leaflet tips 2-
lobed. Inflorescence spicate; peduncular bract
grooved. Staminate flowers stalked or sessile,
asymmetrical; sepals 3, connate basally; petals 3,
 Palmae 381

distinct; stamens 6-18, filaments not inflexed, an-
thers linear, dorsifixed, latrorse; pollen sulcate,
exine tectate, foveolate or fossulate. Staminodal
ring forming a shallowly lobed collar. Fruit 1-
seeded, angled; epicarp smooth, sometimes with
scaly indument, endocarp thick, pores lateral near
base. Endosperm homogeneous, hollow. Germi-
nation remote-tubular; eophyll entire. Four spp.,
Argentina, Bolivia, Brazil, Paraguay.
171. Polyandrococos Barb. Rod.
Polyandrococos Barb. Rod., Contr. Jard. Bot. Rio de Janeiro 1:
7 {1901); Henderson eta!., Field Guide: 153 {1995).
Solitary, acaulescent to erect. Petiole armed or
unarmed; leaflets bilobed, discolorous. Inflores-
cence spicate; peduncular bract grooved. Stami-
nate flowers stalked or sessile, asymmetrical;
sepals 3, connate basally; petals 3, distinct; sta-
mens 60-ca. 100, filaments inflexed, anthers linear,
dorsifixed, introrse; pollen sulcate, exine tectate,
reticulate. Staminodal ring irregularly lobed. Fruit
1-seeded, angled, beaked; epicarp smooth, some-
times with scaly indument, endocarp thick, pores
lateral near base. Endosperm ruminate, hollow or
solid. Eophyll entire. Only one, very variable, sp.,
P. caudescens (Mart.) Barb. Rod., Brazil.
Sc. Subtribe Attaleinae Drude (1887).
carpels. In the past, several genera have been
separated on the basis of staminate flower form
(see Uhl and Dransfield 1987 for discussion).
These genera have now been included in the single
genus Attalea (Henderson et al. 1995). However,
the taxonomy of the group remains poorly
known.
172. Attalea Kunth Fig. 89A-N
Attalea Kunth in H.B.K., Nova Genera et Species Plantarum 1:
folio edition 248; quarto edition 309 {1816); Henderson
eta!., Field Guide: 153-164 (1996).
Maximiliana Mart., Palmarum Familia 20 (1824), nom. cons.
Orbignya Mart. ex Endlicher, Gen. Plant. 257 {1837).
Scheelea Karsten, Linnaea 28: 264 (1857) ("1856").
Small to massive, solitary, acaulescent or erect.
Leaflets single-fold, tips irregularly lobed. Stami-
nate flowers asymmetrical, sessile, or stalked;
sepals 3, distinct, imbricate, or briefly connate;
petals (2)3( -5), equalling sepals or longer, occa-
sionally basally connate, variously membranous
or terete, fleshy, varied in shape; stamens 3-75,
shorter than to much longer than petals, anthers
straight or spirally twisted, dorsifixed to medi-
fixed, sometimes thecae separate or joined,
coiled around each other, latrorse or introrse;
pistillode inconspicuous or absent. Twenty nine
or more spp., S and Central America and the
Caribbean.

Unarmed, pleonanthic, monoecious. Inflores- Sd. Subtribe Elaeidinae J.D. Hook. (1883).
cences solitary, interfoliar, entirely staminate, en-
tirely pistillate, or with flowers of both sexes, Pleonanthic, monoecious; inflorescences inter-
branched to 1 order; prophyll obscured by leaf foliar, bisexual or unisexual, branching to 1 order;
sheaths; peduncular bract tubular, enclosing inflo- prophyll short, becoming fibrous distally; stami-
rescence in bud; staminate rachillae bearing nate flowers spirally arranged, sunken in pits, pis-
paired or solitary flowers; bisexual rachillae either tillate flowers superficial; staminate flowers small;
similar to staminate but bearing a few basal pistil- sepals 3, distinct, widely separated; stamens 6,
late flowers or 1-several triads with a short slender filaments connate in a fleshy tube; gynoecium
apical portion bearing fertile or sterile staminate trilocular, triovulate; ovule atropous; mesocarp
flowers; pistillate rachillae lacking all trace of thick, fleshy, oily; endosperm homogeneous with
staminate flowers at maturity; pollen sulcate or a central cavity; embryo lateral; germination
trichotomosulcate, exine tectate; gynoecium of adjacent-ligular; eophyll entire, lanceolate.
3-several connate carpels; fruit 1-several-seeded, The subtribe is distinguished by the enclosure of
beaked; endocarp very thick, stony; seed basally staminate flowers in pits and by the staminal tube
attached; endosperm homogeneous; embryo formed by the connate filaments; the pistillate
basal; germination remote-tubular; eophyll entire, flower has massive stigmas.
lanceolate.
Attaleinae comprise the single genus Attalea.
The subtribe is distinguished by the presence of 173. Barcella (Trail) Trail ex Drude
more than one type of inflorescence - staminate, Barcella (Trail) Trail ex Drude in Mart., Fl. Brasil. 3{2): 459
androgynous, and sometime also pistillate - on {1881).
the same plant. Gynoecia may have more than 3 Elaeis subgenus Barcella Trail ( 1877).
382
'
. Se. Subtribe Bactridinae J.D. Hook. (1883) .
X
Fig. 89A-X. Palmae-Arecoideae. A Attalea exigua, acaules-
cent palm, showing "saxophone" growth of subterranean
stem. B-G Attalea maripa. B 2 interfoliar inflorescences, each
with a persistent, beaked peduncular bract. C Rachilla of an-
drogynous inflorescence with basal pistillate flowers and dis-
tal staminate flowers. D Staminate flower. E Fruit. F Endocarp,
lateral view. G Same, from below, showing 3 pores. H - N
Attalea speciosa. H Stamen, with coiled anthers. I Pistil with
staminodal cupule. J Ovary in cross-section. K Infructescence
with peduncular bract. L Fruit. M Fruit in cross-section. N
Seeds. 0-X Elaeis guineensis. 0 Stem base with persistent leaf
bases and staminate inflorescences. P Staminate rachilla. Q
Pistillate rachilla with spinose bracts. R Staminate flower. S
Same in longitudinal section. T Pistillate flower. U Pistillate
inflorescence. V Young fruit. W Endocarp. X Seed. (Takhtajan
1982)
Palmae
Solitary, acaulescent, unarmed. Petiole margins
sharp; leaflets single-fold, tips acuminate. Inflo-
rescences bisexual, protandrous; peduncular bract
much longer than prophyll, beaked, woody, stri-
ate, persistent; basal rachillae with few basal pistil-
late flowers, staminate flowers distally, other
rachillae bearing dense spirals of paired staminate
flowers in pits. Anthers rectangular, versatile,
latrorse; pollen sulcate, exine tectate, scabrate;
pistillode much shorter than connate filaments;
staminodal ring 6-toothed, membranous. Fruit 1-
seeded, beaked; endocarp 3-angled, pores 3, lat-
eral. Only one sp., B. odora (Trail) Trail ex Drude,
confined to a small area of the banks of the Rio
Negro and its tributaries in Brazil.
174. Elaeis Jacq. Fig. 890-X
Elaeis Jacq., Sel. Stirp. Americ. Hist.: 280 (1763).
Solitary, erect. Petioles armed with fiber spines
and spinelike eroded leaflet bases; leaflets single-
fold, tips acuminate. Inflorescences unisexual; pe-
duncular bract similar to prophyll, disintegrating
into fibers; staminate rachillae with spinelike tips;
pistillate rachillae with spinelike tips and spinelike
bracts. Anthers rectangular, versatile, introrse;
pollen elliptic or triangular, sulcate, trichotomo-
sulcate, or tetrachotomosulcate, exine tectate, re-
ticulate; pistillode slightly shorter than connate
filaments; staminodal ring 6-toothed, membra-
nous. Fruit 1-seeded, beaked; endocarp variously
angled, pores 3, apical. Two spp., E. guineensis
Jacq., humid tropical Africa, E. oleifera (Kunth)
Cortes, central and northern S America.
Spiny, pleonanthic, monoecious palms, the spines
formed from epidermal emergences, sometimes
unarmed by loss of spines; inflorescences axillary,
interfoliar; gynoecium irregularly ovoid, trilo-
cular, triovulate, variously bristly or tomentose;
stigmas 3, conspicuous, reflexed beyond the petals
at maturity; ovule laterally attached, atropous;
stigmatic remains apical; endocarp pores 3; seed
basally attached; endosperm homogeneous; ger-
mination adjacent-ligular.
This subtribe is immediately separable by the
almost constant presence of epidermal emergence
spines, on stems, leaf sheaths, leaf rachises, some-
times on laminae, and usually on peduncles and
peduncular bracts; sometimes even calyces and
fruit surfaces are spiny. There is considerable
 Palmae
diversity in habit from massive tree palms to
diminutive undergrowth palmlets, acaulescent
palms, and climbers.
175. Acrocomia Mart.
Acrocomia Mart., Hist. Nat. Palm. 2: 66 (1824); Henderson
eta!., Field Guide: 166-167 (1996).
Acaulescent to erect, solitary. Leaflets single-fold,
tips acute or bifid. Inflorescence branching to 1
order, protandrous; peduncular bract persistent,
cowl-like; rachillae with basal, bare portion, and 1-
several triads, distally bearing spirals of solitary
staminate flowers in pits. Stamens 6, anthers
rectangular, medifixed, latrorse; pollen tricho-
tomosulcate, exine tectate. Pistillate sepals free;
staminodal ring 6-toothed, free, or briefly adnate
to petals, with sterile anthers. Fruit 1-seeded, glo-
bose, epicarp smooth, or bristly, endocarp thick,
pores equatorial; endosperm sometimes hollow;
embryo lateral. Two spp., Caribbean and Mexico
southwards to Argentina.
176. Gastrococos Morales
Gastrococos Morales, Repertorio Fisico-Natural de Ia Isla de
Cuba 1: 57 (1865); Uhl & J. Dransf, Gen. Palm.: 519-521
(1987).
Solitary, erect, stem ventricose. Leaflets single-
fold, tips acute or bifid, discolorous. Inflorescence
branching to 1 order, protandrous; peduncular
bract, persistent, cowllike; rachillae with basal,
bare portion, and 1-several triads, distally bearing
spirals of solitary superficial staminate flowers.
Stamens 6, filaments not inflexed, anthers linear,
dorsifixed, latrorse; pollen trichotomosulcate,
exine tectate, reticulate. Pistillate calyx cupular;
staminodal ring 6-toothed, adnate to petals. Fruit
1-seeded, globose, epicarp smooth, endocarp
thick, pores equatorial; endosperm hollow;
embryo lateral. Eophyll entire. Only one sp., G.
crispa (Kunth) H.E. Moore, Cuba.
177. Aiphanes Willd.
Aiphanes Willd., Mem. Acad. Roy. Sc. Berlin 1804: 32 (1807);
Borchsenius & Bernal, Flora Neotrop. 70: 1-94 (1996).
Solitary or caespitose, acaulescent to erect.
Lamina undivided or with single-fold leaflets, tips
praemorse. Inflorescence spicate or branching to 1
or 2 orders, protandrous; peduncular bract persis-
tent or disintegrating; rachillae bearing triads
383
basally and staminate flowers distally. Stamens 6,
basally connate, anthers medifixed; pollen
sulcate or trichotomosulcate, exine tectate, with
supratectal processes. Pistillate sepals distinct;
staminodal ring 6-toothed, adnate to petals. Fruit
1-seeded, globose, epicarp smooth or spiny, en-
docarp thick, pores equatorial; endosperm hollow;
embryo lateral. Eophyll entire. About 22 spp.,
West Indies and northern S America, especially
diverse in Andean region.
178. Bactris Jacq. ex Scolopi Fig. 82G
Bactris Jacq. ex Scopoli, lntroductio ad Historiam Naturalem
70 (1777); Henderson, Palms Amazon: 167-224 (1994).
Guilielma Mart. (1824).
Solitary or caespitose, acaulescent to erect.
Lamina undivided or leaflets single- or multi-fold,
tips acute, rarely praemorse, concolorous or
discolorous. Inflorescence spicate or branching to
1 order, protogynous; peduncular bract persistent
or disintegrating; rachillae bearing triads through-
out or basally with staminate flowers distally, or
few triads scattered among staminate flowers. Sta-
mens 6( -12), distinct, anthers dorsifixed; pollen
sulcate or trichotomosulcate, exine tectate. Pistil-
late sepals and petals connate; staminodal ring 6-
toothed, free. Fruit 1-seeded, epicarp smooth or
spiny, pores equatorial; embryo lateral. Eophyll
bifid. Sixty four spp. recognized by Henderson et
al. (1996), but 239 described in the past; the genus
remains poorly known. Distributed from Central
to S America and the Antilles.
179. Desmoncus Mart.
Desmoncus Mart., Palmarum Familia: 20 (1824); Henderson,
Palms Amazon: 224-233 (1994).
Slender, caespitose, erect or climbers. Leaf usually
with terminal cirrus bearing acanthophylls; ocrea
present; leaflets single-fold, acuminate, con-
colorous. Inflorescence spicate or branching to 1
order, protandrous; peduncular bract persistent;
rachillae bearing triads basally and staminate
flowers distally. Stamens 6-9, distinct, adnate to
corolla, anthers rectangular, basifixed, latrorse;
pollen sulcate, exine tectate, fossulate. Pistill-
ate sepals and petals connate; staminodes 6,
epipetalous, not connate in a ring. Fruit 1-seeded,
epicarp smooth, pores slightly distal to equator;
embryo lateral. Eophyll bifid. Seven spp., Central
and S America, many more recognized in the
past.
384
180. Astrocaryum G.F.W. Meyer
Astrocaryum G.F.W. Meyer, Prim. Fl. Essequeb.: 265 (1818),
nom. cons.; Kahn & Millan, Bull. Inst. Fr. et. Andines 21:
459-531 (1992).
Acaulescent to erect, solitary or clustered. Leaflets
single-fold, discolorous, tips acute. Inflorescence
branching to 1 order, protandrous; peduncular
bract persistent; rachillae with or without basal,
bare portion, and !-several triads, distally bearing
spirals of solitary staminate flowers in pits. Sta-
mens 3-12, anthers dorsifixed, latrorse; pollen
sulcate or trichotomosulcate, exine tectate. Pistil-
late sepals and petals connate; staminodal ring
6-toothed, epipetalous. Fruit 1(-2)-seeded,
epicarp smooth or bristly, sometimes splitting
irregularly, displaying endocarp, pores subtermi-
nal. Endosperm hollow; embryo subterminal.
Eophyll bifid. Eighteen or more spp., Central to S
America.
6. Tribe Geonomeae Drude (1881).
Unarmed, pleonanthic, monoecious palms; leaves
reduplicately pinnate, sheaths fibrous, not form-
ing a crownshaft, leaflets acute; inflorescences
solitary, interfoliar, sometimes becoming infra-
foliar, protandrous; rachillae with bracts subtend-
ing a triad of flowers enclosed in a pit, floral
bracteoles 3, narrow, keeled, chaffy, sepal-like;
staminate flower with sepals 3, imbricate basally,
chaffy; petals 3, connate basally; stamens very di-
verse; pollen sulcate, exine tectate, vermiculate,
reticulate, fossulate; pistillodes 3-lobed, minute;
pistillate flower with sepals 3, imbricate basally,
chaffy; petals 3, connate basally; gynoecium
trilocular, triovulate, or pseudomonomerous,
with a central elongate style terminating in 3 ex-
serted, spreading, slender stigmas; ovules anatro-
pous, only one maturing. Fruit with basal
stigmatic remains; endosperm homogeneous;
embryo basal; germination adjacent-ligular;
eophyll bifid.
The Geonomeae comprise six genera restricted
to the New World. They are characterized by the
presence of deep pits on the rachillae, enclosing
triads. The genera show different specializations
of corolla, androecium, and staminodal ring, re-
lated to the presentation of pollen or nectar out
of the pits. Five of the genera have trilocular,
triovulate gynoecia, while Geonoma usually has
only 1 fertile ovule.
Palmae
KEY TO THE GENERA OF GEONOMEAE
1. Moderate to large tree palms; leaves regularly pinnate 2
- Acaulescent or small understory palms, stem short or if tall,
then slender and canelike; leaves undivided and bifid or
variously divided, rarely evenly pinnate 3
2. Leaflets broadly lanceolate, lacking a distinct midrib; inflo-
rescence stout, rachis very short bearing a few (ca. 8) long
pendulous rachillae; stamens numerous 36( -42); petals of
pistillate flower connate for 2/3 of their length, tips valvate,
glumaceous; staminodes ca. 15, awl-shaped where free
182. Welfia
- Leaflets narrowly lanceolate, midrib and 2 pairs of lateral
ribs evident abaxially; inflorescence moderate, bearing
many rachillae equal in length, clustered at the end of the
long peduncle, stamens 6; petals of pistillate flower connate
in a tube, opening by a circumscissile cap; staminodes con-
nate, forming a fleshy tube enclosing the style and stigmas
183. Calyptronoma
3. Bracts covering floral pits not "locked" into the rachilla
distally but upper margins rounded, truncate, or split, lat-
eral margins adnate to the rachilla beside the floral pits;
anthers terminal on the end of the connective, inflexed in
bud, thecae spread apart or parallel; ovary unilocular at
anthesis 186. Geonoma
- Bracts covering the floral pits overlapping laterally or im-
mersed in the rachilla; anthers sagittate or thecae terminal
on a bifid connective; ovary trilocular at an thesis 4
4. Stem solitary, short to moderate, leaves ± irregularly
divided into several-fold pinnae; petioles long, slender,
bracts covering the floral pits overlapping laterally, densely
tomentose, filaments of stamens and staminodes united in
a short tube but free and awl-shaped distally
181. Pholidostachys
- Stem solitary, usually very short or lacking, leaves bifid or
with usually unequal, several-fold pinnae; petioles short,
bracts covering the floral pits not overlapping laterally,
glabrous or lightly hairy, not densely tomentose 5
5. Leaves bifid, bracts covering the floral pits "locked" in bud
by a distinct rounded upper lip on the pit; thecae separated
on a bifid connective; staminodes free and fleshy distally
185. Asterogyne
- Leaves irregularly divided, floral pits without definite upper
lip, anthers sagittate; staminodes united in a tube, con-
stricted at the middle, rounded and very briefly 6-lobed
distally 184. Calyptrogyne
181. Pholidostachys H. Wendl. ex J.D. Hook.
Pholidostachys H. Wend!. ex J.D. Hook. in Benth. & Hook.,
Gen. Pl. 3: 915 (1883); Henderson eta!., Field Guide: 207-
208 (1996).
Calyptrogyne H. Wend!. subgen. Pholidostachys (H. Wend!.)
Wessels Boer, Verh. Kon. Ned. Akad. Wetensch. II, 58 (1);
73 (1968).
Solitary. Inflorescence spicate or branching to
1(-3) orders; peduncular bract inserted above
prophyll, caducous or marcescent; margins of pit
bracts sometimes overlapping. Stamens 6, anthers
sagittate, medifixed, exserted and spreading at
anthesis, introrse, connectives with pointed tips.
Pistillate flowers with only free parts of sta-
 Palmae
minodes and stigmas exserted at anthesis; sta-
minodes 6-8, fleshy, connate in basal 2/3, adnate
to corolla, free portions angled, terete, exserted
and spreading at anthesis. Mesocarp with thick,
curved, anastomosing fibers. Four spp., Costa Rica
to northern S America.
182. Welfia H. Wendl.
Welfia H. Wend!., Gartenflora 18: 242 (1869); Henderson et al.,
Field Guide: 209-210 (1996).
Solitary tree. Inflorescences branched to 1(-2)
orders; peduncular bract inserted near prophyll;
rachillae with 8 rows of pits. Stamens 36(27-42) in
antesepalous and antepetalous groups, anthers
linear-sagittate, basifixed, introrse, connectives
with pointed tips. Pistillate flower with 15-16
staminodes, adnate to corolla for 2/3 their length,
free above. Fruit compressed dorsiventrally, later-
ally ridged, pointed; mesocarp with slender paral-
lel fibers. Only one sp., W. Megia H. Wendl. ex
Andre, Honduras to west Colombia.
183. Calyptronoma Griseb.
Calyptronoma Griseb., Flora of the British West Indian Is-
lands 518 (1864); Zona, Principes 39: 140-151 (1995).
Calyptrogyne subgenus Calyptronoma (Griseb.) Wessels Boer,
Verh. Kon. Ned. Akad. Wetensch. II, 58 (1): 63 (1968).
Solitary, erect. Inflorescences branched to 1(-4)
orders; peduncular bract inserted halfway along
peduncle; pits closed with rounded bracts. Sta-
mens 6, filaments connate, forming a stalk, elon-
gating at anthesis, free filaments far exserted,
anthers sagittate, dorsifixed near base, introrse,
versatile. Pistillate flowers with petals connate
with short valvate apices, upper 1/4 shed as a cir-
cumscissile cap; staminodes completely connate,
urn-shaped, inflated above corolla tube at anthe-
sis, pushing off corolla cap. Mesocarp with stout
fibers anastomosing distally, one large fiber con-
spicuous. Three spp., Greater Antilles.
184. Calyptrogyne H. Wendl.
Calyptrogyne H.A. Wend!., Bot. Zeit. 17: 72 (1859); Henderson
et al., Field Guide: 211-212 (1996).
Solitary, acaulescent or erect. Inflorescence spi-
cate, rarely branched to 1 order; peduncular bract
inserted just below flowers, caducous; pits closed
by rounded bracts. Stamens 6, filaments connate
forming a stalk, free lobes recurved at anthesis,
anthers sagittate, dorsifixed near base, introrse.
385
Pistillate flowers with petals connate, very briefly
free, valvate distally, distal 1/3 shed as a cap;
staminodes connate, constricted near middle,
briefly 6-lobed distally, upper part shed, revealing
stigmas. Mesocarp with layer of large anastomos-
ing fibers, the largest median. Eight spp., Mexico
to northwestern Colombia.
185. Asterogyne H. Wendl. ex J.D. Hook.
Fig. 811
Asterogyne H. Wend!. ex J.D. Hook. in Benth. & J.D. Hook.,
Gen. Pl. 3: 914 (1883); Henderson et al., Field Guide: 213-
214 (1996).
Solitary, small. Inflorescences spicate or branched
to 1 order, peduncular bracts 1(-2), inserted near
prophyll; rachillae 1 or few (2-8), pits with distinct
upper "lips" locking bract over developing flow-
ers. Stamens 6-ca. 24, filaments connate in a solid
stalk, tubular above, distally free, connective bifid,
thecae separate, inflexed, erect at anthesis,
introrse. Pistillate corolla tubular in basal 2/3;
staminodes ca. 15, connate basally, adnate to
corolla, free distally. Mesocarp with a layer of
appressed, longitudinal fibers. Five spp., central
to northern S America.
186. Geonoma Willd. Fig. 82B
Geonoma Willd., Sp. Pl. 4(1): 174, 593 (1805); Wessels Boer,
Verh. Kon. Ned. Akad. Wetensch. II, 58 (1): 1-202 (1968).
Small, solitary or clustered. Inflorescences spicate
or branched to 1-3( -4) orders; peduncular bracts
(0- )1( -2), usually inserted near prophyll. Rachilla
pits decussate, spiral, or whorled, with or without
a distal lip. Stamens 3(6), (rarely more), filaments
connate in a stalk, tubular above, distally free,
inflexed, connective divided with divaricate
thecae or short and united, introrse. Pistillate
petals connate basally; staminodes connate,
tubular, truncate, or 6-lobed. Mesocarp thin,
with narrow longitudinal fibers. Seventy five,
possibly fewer spp., from Mexico to Brazil and
Bolivia.
VI. Subfamily Phytelephantoideae Drude
(1887).
Acaulescent or erect; unarmed; pleonanthic, dioe-
cious; leaves pinnate, reduplicate; inflorescences
dimorphic; peduncular bract 1; staminate inflo-
rescence spikelike or racemose, pistillate inflores-
cence headlike; flowers multiparted, the staminate
386 Pal mae

in monopodia! clusters, the pistillate solitary,

crowded; staminate flowers with much reduced
perianth and very numerous stamens; pistillate
flower with large, elongate, fleshy sepals and pet-
als, gynoecium of 5-10 connate carpels, styles
long; stylodia elongate; ovules hemianatropous to
anatropous; fruits 5-10-seeded, borne in a con-
gested headlike cluster; mesocarp cracking to
form irregular, fibrous, pyramidal warts; en-
dosperm very hard, homogeneous with smooth,
hard, thin pyrenes, enclosing individual seeds.
Eophylls pinnate.
The phytelephantoid genera add an extraordi-
nary dimension to inflorescence and flower struc-
ture in palms, and recently a growth habit, not
previously known in the family, has been discov-
ered in Phytelephas (Bernal1998). The flower clus-
ters are condensed monopodia! units rather than
sympodial ones (rhipidia) as throughout the rest
of the family. Flowers are multiparted, some of
them showing a tetramerous pattern in early
stages. Stamen numbers are very large with over
1000 reported in Ammandra, the highest number
known in the family. Development of stamens is
centrifugal. The subfamily has been resolved as
monophyletic (Barfod 1991; Barfod et al. 1998; Uhl
et al. 1995), and in some of the analyses was sister
to the Ceroxyleae. However, its relationships to
other palms remain unclear.

KEY TO THE GENERA OF THE

PHYTELEPHANTOIDEAE

l. Staminate inflorescence spicate 2

- Staminate inflorescence not spicate, flowers borne singly
but crowded on the ends of short branches; receptacles
angled, stamens ca. 500 or more, filaments short or lacking,
anthers rounded or slightly elongate; corky projections on
fruits few, large, fruits several-seeded. Colombia
189. Ammandra
2. Staminate inflorescence a spike with flowers sessile or in
groups of 4- more on short or long grooved stalks; recep-
tacles flat; stamens ca. 150-699; fruits 4- 10-seeded, seldom Fig. 90A-O. Palmae-Phytelephantoideae. Phytelephas seem-
with 6 seeds. Amazon Basin, Colombia, Venezuela, and annii. A Plant with pistillate inflorescences. B Stem base with
Panama 187. Phytelephas staminate inflorescences. C Staminate flower. D Stamen. E
- Staminate flowers in groups of 4( -6) flowers united by Pistillate flower. F Gynoecium and staminodes, the style bear-
pseudopedicels (elongate floral receptacles with adnate ing numerous stylodia. G Ovary with staminodia. H Ovary in
perianth members), stamens ca. 400- 659; fruits usually cross-section. I Sepals. J Petals. K Infructescence with several
6-seeded 188. Aphandra fruit crowded together. L Fruiting head, all fruits removed but
l. M Seed with partly removed endocarp. N Seed in longitudi-
nal section. 0 Seedling. (Takhtajan 1982)
187. Phytelephas Ruiz & Pavon Fig. 90A-O
Phytelephas Ruiz & Pavon, Syst. Veg.: 299 (1798); Barfod,
Opera Botanica 105: 5-73 (1991). ranged. Prophyll and peduncular bract glabrous;
Palandra O.F. Cook (1927). staminate inflorescences unbranched; flowers
sessile or short-stalked, in groups of 4; receptacles
Stem erect or procumbent. Petiole less than rounded, flat; stamens 150-600; pollen elliptic,
200 em long. Leaflets regularly or irregularly ar- often with one end dilated, sulcate or trichotomo-
 Palmae
sulcate, exine tectate, reticulate; pistillode lacking.
Germination remote-ligular. About five spp., with
a bicentric distribution, occurring in the Amazo-
nian Basin in Bolivia, Ecuador, and Peru, along the
northwest coast of Colombia, and into Venezuela
and Panama.
188. Aphandra Barfod
Aphandra Barfod, Opera Bot. 105: 5-73 (1991); Balslev &
Henderson, Syst. Bot. 12: 501-504 (1987).
Stem erect. Petiole less than 200 em long. Leaflets
regularly arranged, sheath margins with long
fibers. Prophyll and peduncular bract brown scaly;
staminate inflorescences with short branches
bearing clusters of 4( -6) flowers, floral receptacles
elongate, adnate to perianth; stamens 450-650;
pollen elliptic, often with one end dilated, sulcate
or trichotomosulcate, exine tectate, reticulate;
pistillode sometimes prominent. Pistillate heads
with 30-50 flowers. Germination remote-ligular.
One species (A. Natalia Balslev & Hend.) Barfod,
endemic to eastern foothills of the Ecuadorean
Andes, ranging east into lowlands of Amazonian
Ecuador.
189. Ammandra O.F. Cook. Fig. 82D
Ammandra O.F. Cook, J. Wash. Acad. Sci. 17: 220. 1927. A.S.
Barfod, Opera Bot. 105: 5-73 (1991).
Solitary or caespitose. Stem prostrate or decum-
bent. Petioles longer than 200 em, terete distally.
Leaflets regularly arranged, sheath margins with
flat, hairlike fibers. Prophyll and peduncular bract
brown-scaly; staminate inflorescences with short
branches bearing clusters of 6-9 flowers, recep-
tacles asymmetrical; stamens 300-1200; pollen
elliptic, monoporate, exine tectate, reticulate;
pistillode sometimes prominent. Pistillate heads
with 6-10 flowers. Germination remote-ligular.
Two spp., one from the western coastal regions of
Colombia and a second from eastern Colombia.
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Barrow, S. 1996. A monograph of Phoenix L. (Palmae:
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387
Bernal, R. 1998. The growth form of Phytelephas seemannii-
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 Palmae/Philydraceae
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389
Philydraceae
U. HAMANN 1
Philydraceae Link, Enum. Plant. Hort. Berol. Alt. 1: 5 (1821),
nom. cons.
Perennial herbs with rhizomes or corms and
fibrous roots; leaves basal or crowded near the
stem base, distichous, with equitant, sheathing
base and linear-ensiform or terete, unifacial blade.
Flowering stem with few upwards gradually re-
duced leaves; inflorescence a terminal simple or
compound spike; flowers sessile in the axils of dis-
persed, spathaceous bracts, bisexual, (nearly) hy-
pogynous, zygomorphic; perianth petaloid, pale
yellow or white to pink; segments 4, free or basally
connate, the outer 2 median larger than the 2 lat-
eral inner, the median upper frequently 3- or 2-
cuspidate, composed of the 2 lateral outer tepals
and the reduced median inner tepal; stamen soli-
tary, anterior, its filament broad, inserted at the
base of the lower tepal; anther bithecate and
tetrasporangiate, basifixed to dorsifixed, straight
and introrse, or curved (Philydrella), or helically
twisted (Philydrum), opening by 2 longitudinal
slits; staminodia usually absent; gynoecium of 3
united carpels; ovary superior, often slightly
united at the base with the perianth and filament,
::±:: zygomorphic (the median carpel often smaller),
::±:: trilocular with axile placentas, or for the great-
est part unilocular with deeply intruding parietal
placentas; each placenta bearing numerous ana-
tropous, heterotropous ovules; style simple, ter-
minal, with a capitate, slightly trilobed stigma;
septal nectaries or other nectariferous glands
absent. Fruit a loculicidal, 3-valved capsule, or
berrylike; seeds numerous, small; testa longitudi-
nally or spirally striate, operculate, with enlarged
chalazal cap or prolongation at micropylar and
chalazal end; embryo small, straight, cylindrical,
embedded in copious non-mealy endosperm con-
taining starch, protein and oil.
Four genera and five species in Australia and SE
Asia.
1 Deceased.
390 Philydraceae

VEGETATIVE MORPHOLOGY. The rootstock in

Philydraceae is either a rhizome or a corm. In
Orthothylax and Helmholtzia the rhizome is stout,
ascending to vertical and often branched. In
Philydrum the rhizome is short and erect.
Philydrella has a small corm similar to that found
in some Iridaceae. The corm consists mainly of 1
thickened internode and is renewed annually from
the axillary bud of the basal leaf of the flowering
shoot.
In all genera the leaves (in Philydrella often only
a single leaf) are mostly basal and distichous
(Fig. 91). They have an open sheath, and a
unifacial, equitant, ensiform or (in Philydrella)
terete, acuminate blade; a ligule is lacking. The
texture of the thick, erect, aerenchymatous leaf
blades of Philydrum is soft and spongy. The leaves
of Helmholtzia and Orthothylax have a flattened,
firmer and broader lamina which twists, becoming
± horizontal and secondarily differentiated into
an upper and lower surface of different colour and
anatomy.
The epigeal cotyledon and the small primary
leaves of young plants are bifacial (investigated in
Philydrum).
The elongated, terminal flowering stem has only
few leaves. These leaves have progressively
reduced blades and are transitional to the bifacial,
acuminate floral bracts. In the upper part the
phyllotaxis changes from distichous to alternate.
Long woolly hairs are pressed on the flowering
stem and bracts except in Phylidrella.

VEGETATIVE ANATOMY. The roots of Philydrum

possess an exodermis of 2 cell layers, a thiCk
cortex with many radial air cavities, a weakly
thickened endodermis, a closed, !-layered peri-
cycle and polyarch vascular bundle (Solereder and
Meyer 1929). The xylem of the roots in Philydrum
and Orthothylax has tracheids and vessels with
scalariform perforation plates (Fahn 1954;
Cheadle 1968). Persistent 1-celled root hairs with Fig. 91A-H. Philydraceae. Philydrum lanuginosum. A Habit.
B Flower. C Flower with perianth half cut away. D Stamen. E
thickened, brownish cell walls often form a dense Lateral tepa!. F Pistil. G Seed. H Same in longitudinal section.
felt on older roots of Helmholtzia and Ortho- a embryo; b micropylar endosperm; c chalaza! endosperm;
thylax. d operculum. (Takhtajan 1982)
The flowering stem of Philydrum has below the
cortex a lignified sclerenchymatous tissue, which
forms a peripheral sheath and includes small blade they form a complete ring arranged with the
collateral or semiconcentric vascular bundles. phloem outwards. The ensiform blades have
Bundles are also present in the cortex and in the numerous parallel veins which are connected by
parenchymatous pithlike central part of the axis. short transverse veinlets. In the secondarily dor-
Vessels are lacking in the stem and also in leaves siventral blades of Helmholtzia and Orthothylax a
of Philydraceae. complex of veins forms a "false costa", which is
In the leaf sheath the collateral bundles form a prominent on each surface. In these genera the
bow in transverse section, but in the unifacial assimilatory tissue of the upper side is more devel-
 Philydraceae 391

oped than on the lower. Stomata are here the outer whorl, which is usually only slighty
restricted to the lower surface (Skottsberg 1932). larger than the anterior, is derived by fusion of the
The leaf blades of Philydrum and Philydrella have 2 lateral outer tepals and the median-posterior
an isolateral anatomy. In Philydrum the leaf tepal of the inner whorl. In the past, the median
blades have a central aerenchyma oflarge lacunae, inner tepal was thought to be suppressed (Pilger
separated by narrow diaphragms of parenchyma- 1930; Skottsberg 1932); but it is often represented
tous tissue; this is surrounded by smaller lacunae by a weak median vein best developed in
and by assimilatory tissue which contains the Philydrum, between the 2 strong median (and fre-
peripheral collateral bundles with sclerenchyma- quently 4 lateral) veins of the 2 outer tepals. In
tous sheaths (Solereder and Meyer 1929). Orthothylax and Helmholtzia the 2-keeled poste-
The stomata of Helmholtzia and Orthothylax rior segment is frequently bidentate, while in
have 2 to 4 lateral and 2 polar subsidiary cells, Philydrum (sometimes also in the two other
some of which are only poorly distinct from other genera) it is regularly tridentate. Each of the 2
epidermal cells. In Philydrum and Philydrella gen- small lateral segments of the inner perianth whorl
erally only 2 lateral subsidiary cells are present, has 1 median vein, which may branch near the
which are much smaller than the elongated epi- base.
dermal cells. The subsidiary cells are derived from In Helmholtzia all perianth segments are con-
cells adjacent to the guard cell initials (Hamann nate into a short tube. In the other genera the
1966). anterior segments are ± free or only slightly con-
The long woolly hairs of the flowering stems of nate at their base; in Philydrella the 2 small lateral
Orthothylax, Helmholtzia and Philydrum are few- tepals are adnate to the base of the filament.
celled and unbranched. These hairs occur also on The single stamen has a median-anterior posi-
the inner surface of the leaf sheath in Helmholtzia tion and belongs to the outer whorl of the
novo-guineensis. androecium which is otherwise suppressed. In
Tanniniferous cells and idioblasts with styloids Philydrum a small lateral stamen or staminodium
of oxalate are frequent in all organs. The sieve- of the outer whorl may rarely be developed. The
tube plastids have cuneate protein bodies but lack inner whorl is always lacking. Helmholtzia and
starch. Orthothylax have straight, introrse anthers with a
broad, brown, tanniniferous connective. In
INFLORESCENCES. All Philydraceae have an inflo- Philydrella the young anther is introrse, later
rescence of the indeterminate, racemose type. It becoming reniform or semiorbicular by predomi-
consists of a terminal spike with many or (in nant growth of its adaxial part; in Philydrum the
Philydrella) few flowers each in the axil of a thecae become helically twisted by increased
spathelike bract. In all genera except Philydrella growth.
(and dwarfforms of Philydrum) additional lateral The gynoecium consists of 3 fused carpels with a
spikes develop below the terminal one. These are simple style and a capitate stigma. The ovary is
generally unbranched in Philydrum so that the basally connate to the perianth. Due to the weaker
terminal spike remains predominant. In Helm- development of the anterior carpel, it is ± zygo-
holtzia and Orthothylax at least some of these morphic, most strongly so in Orthothylax. The
numerous lateral spikes bear tertiary spikes, some ovary is 3-locular, with axile placentas, or in the
of which may branch again. Thus the whole inflo- upper part 1-locular. In Philydrum only the lower-
rescence becomes a large "panicle" composed of most part is completely 3-locular; in the main part
numerous spikes, in which the primary terminal of the ovary the 3 septa protrude deeply but do not
spike is a small component. fuse in the centre. In Orthothylax the fusion of the
septae is weak and the septa separate when the
FLORAL STRUCTURE. The characteristic zygo- capsule opens. The placentas bear many ovules,
morphic flower ofPhilydraceae, with its 4 perianth particularly in Philydrum.
segments and single stamen, is unique in the
monocotyledons (Fig. 91B,C). It has been derived EMBRYOLOGY. The embryology ofPhilydraceae is
from a trimerous-pentacyclic prototype by fusion fairly well known and apparently uniform (see
and suppression of organs. Hamann 1966), although completely studied only
Of the 2 members of the outer perianth cycle for Philydrum lanuginosum. The anther wall
only the median-anterior is a single organ. It has 1 develops according to the Monocotyledonous
median vein and 2 weaker lateral veins which type. The endotheciallayer has thickenings of the
branch <fnd anastomose. The posterior segment of Girdle type (Orthothylax) or transitions to the Spi-
392 Philydraceae
ral type (Philydrum). The tapetum is secretory; its
cells become 2- to 4-nucleate and always contain
raphides. The mature pollen grains are 2-celled.
The ovules are anatropous, mixed epi-, pleuro-,
and apotropous, bitegmic and weekly cras-
sinucellate. After fertilisation, the outer integu-
ment elongates at the micropylar region, while the
inner integument gives rise to an operculum. The
archesporia! cell cuts off a primary parietal cell
which usually divides once. The funiculus forms
an obturator, with an air cavity in Philydrum.
After meiosis an 8-nucleate, 7-celled embryo sac
develops from the chalazal megaspore according
to the Polygonum type. Endosperm development
follows a peculiar subtype of the Helobial type, in
which the small chalazal endosperm chamber
becomes cellular earlier than the large micropylar
chamber. Only the micropylar chamber serves as
storage tissue.
PoLLEN MoRPHOLOGY. The pollen grains are
monads, except in Philydrum lanuginosum, where
they remain united in tetrads. Rap hides of calcium
oxalate derived from the degenerated tapetal cells
are always intermixed with the pollen. The pollen
grains are sulcate (Erdtman 1952; Hamann 1966).
They have a broad distal aperture and reticulate
(in Orthothylax minutely and indistinctly reticu-
late) exine. The long axis ranges from ca. 30 f.tm
(Helmholtzia, Orthothylax) to 60 f.tm (Philydrella).
KARYOLOGY. For Helmholtzia acorifolia and
Orthothylax glaberrimus a chromosome number
of 2n = 34 has been determined (Briggs 1966), for
the latter species also 2n = 32 and for Philydrum
lanuginosum 2n = 16 (Hamann 1966).
PoLLINATION. No field observations on the polli-
nation of the nectarless and scentless pollen flow-
ers of the Philydraceae are known. Cultivated
plants of Philydrum lanuginosum, which regularly
set seeds, are autogamous. The pale yellow flowers
last only 1 day, and the few flowers of the whole
inflorescence bloom simultaneously. After anthe-
sis the tepals and bract close around the pistil and
stamen. The twisted anther comes into contact
with the stigma, and the pollen tetrads germinate
partly inside the dehiscent anther. It is unknown
whether entomogamy occurs in nature.
Philydrella and presumably the other genera are
at least facultatively autogamous. In the white
flowers of Helmholtzia acorifolia the style exceeds
the anther, which makes autogamy more difficult
and is perhaps the reason for the poor fruit set in
the greenhouse.
FRUIT AND SEED. In most genera the fruit is a
loculicidal capsule opening with 3 valves, to which
in Philydrum and Orthothylax the placenta is
attached. In the centre of the ovary of Orthothylax
the loosely connected septa become separated at
least in the upper part, while in Philydrella
the septa break and the valves separate from the
remaining central column. In Helmholtzia the
fruit is berrylike, finally with a leathery, appar-
ently indeshiscent (or tardily dehiscent?) peri carp.
The seeds (Fig. 91G,H) are small and generally
very numerous (in Helmholtzia often only few).
They have a starchy, but not mealy, endosperm with
large single (not compound) starch grains, peculiar
crystalloid aleuron bodies and also some oil. The
outer endosperm layer is clearly differentiated and,
like the remnants of the chalazal endosperm cham-
ber, does not contain large starch grains. The small,
linear, cylindric embryo is embedded in the
endosperm and lacks starch. It consists mainly of
the straight cotyledon; the radicle is very short and
the plumule only few-celled.
Both integuments participate in the seed coat,
which becomes longituginally or (especially in
Philydrum) spirally striate. The elongated
exotestal cells have thick cellulose walls. The
endotegmic layer contains tannin. Cuticules are
developed between the tegmen and the remnants
of the nucellus and (except in Philydrella) also
between testa and tegmen. An operculum devel-
ops in the micropylar region from both layers of
the inner integument. It is later pushed off by the
germinating embryo.
DISPERSAL. No observations on seed dispersal in
natural habitats are available, but the tiny seeds of
Philydrum and Philydrella may be dispersed by
water; they are known to float for long periods.
The same may be true for the elongated (scobi-
form) seeds of Orthothylax which also would be
suitable for wind dispersal. It is unknown whether
the berrylike fruits of Helmholtzia are endozo-
ochorous or become dry and ultimately dehisce.
PHYTOCHEMISTRY. The cell walls contain ferulic
and diferulic acids (Harris and Hartley 1980). In
Philydrum, syringetin and common flavonols
(Bohm and Collins 1975) and proanthocyanidins
(N. Amrhein, pers. comm.) are found. Pro-
cyanidin and prodelphinidin are also present in
the leaves of Orthothylax (Bate-Smith 1968). No
alkaloids or saponins are known so far.
RELATIONSHIPS WITHIN THE FAMILY. Two
groups of genera can be distinguished, one com-
 Philydraceae
pnsmg the montane rainforest genera Helm-
holtzia and Orthothylax, the other the lowland
genera Philydrum and Philydrella. The two former
are so closely related that sometimes they are
combined in the single genus, Helmholtzia, but
differences in perianth, ovary, fruit and seed
justify their generic separation. The geographi-
cally isolated Philydrella resembles a dwarf, or
flowering juvenile form, of Philydrum. Its tuber-
ous corm, terete leaf, reduced size and inflores-
cence, may be interpreted as adaptations to the
long dry period of the SW Australian summer.
AFFINITIES. The combined evidence of reproduc-
tive and vegetative characters indicates that
the closest relatives of the Philydraceae are the
Pontederiaceae. Common features include the
ovary position, embryology, embryo and endo-
sperm in the ripe seed, vegetative anatomy (para-
cytic stomata, styloid crystal idioblasts, etc.)
and similar proanthocyanins. The structure of
the exine and testa are, however, dissimilar,
and the Pontederiaceae usually have septal
nectaries. The basic chromosome numer of x = 9
in Philydraceae is consistent with a shared
ancestry with Pontederiaceae, which apparently
have x = 8.
The Haemodoraceae also agree with the
Philydraceae in many respects. Remarkable simi-
larities include the ensiform leaves, the pubescent
flowering stems, the paracytic stomata, features of
the Helobial endosperm formation and its starchy
contents, and the amoeboid tapetum.
The type of Helobial endosperm formation with
early cell-wall formation in a small chalazal cham-
ber that precedes wall formation in a large micro-
pylar chamber is found in the Philydraceae,
Typhaceae, Bromeliaceae, Velloziaceae and at
least some Burmanniaceae, Pontederiaceae and
Haemodoraceae. This seems to be an important
feature that may have appeared in their common
ancestor.
Thus the morphological data would suggest
a position of the Philydraceae adjacent to the
Pontederiaceae with further affinities to the
Haemodoraceae and perhaps Bromeliaceae and
Velloziaceae. With respect to the closeness of
Philydraceae, Pontederiaceae and Haemodora-
ceae, this conclusion is supported by recent
cladistic analyses based on morphological and
molecular (rbcL) data (Chase et al. 1995; Linder
and Kellog 1995).
DISTRIBUTION AND HABITATS. The family is
confined to Australia and tropical E Asia.
393
Philydrum is the only genus with a wider distribu-
tion, ranging from Victoria along the E and N
coastal parts of Australia to S New Guinea and
from Malaysia and Thailand to S China, Taiwan
and S Japan. Helmholtzia is restricted to tropical
NE Australia, New Guinea and the Moluccas and
Orthothylax to a limited area of (sub- )tropical E
Australia. Philydrella is isolated from the rest of
the family in SW Australia.
Most Philydraceae live in (sub)tropical climate,
occurring either in wet habitats of the montane
forest (Helmholtzia, Orthothylax) or of the low-
land (Philydrum). Philydrella grows in a season-
ally dry climate on temporarily moist, sandy
alluvial lands, surviving the dry period with its
corm.
KEY TO THE GENERA
1. Inflorescence much branched, at least with some tertiary
spikes; anther straight; seeds > 1.5 mm long 2
- Inflorescence a simple terminal spike or only with
unbranched secondary spikes; anther curved or helically
twisted; seeds < 1 mm long 3
2. Tepals united at the base, the outer dorsally pubescent;
ovary densely pubescent, not clearly zygomorphic; fruit
berrylike; seeds < 3 mm long 1. Helmholtzia
- Tepals free, glabrous; ovary (sub)glabrous, strongly zygo-
morphic; fruit capsular; seeds 3.5-4.5mm long
2. Orthothylax
3. Flowering stem 40-100 (or more) em long, with several30-
80-cm-long basal leaves; terminal spike many-flowered,
often with secondary spikes; inflorescence axis lanuginose;
anther helically twisted 3. Philydrum
- Flowering stem less than 20 em long, at an thesis mostly with
a single± 10-cm-long basal leaf; inflorescence a simple, few-
flowered spike; inflorescence axis glabrous; anther curved,
reniform 4. Philydrella
Genera of the Philydraceae
1. Helmholtzia F. Muell.
Helmholtzia F. Muell., Fragm. Phytogr. Austr. 5: 222 (1866).
Tall, stoutly rhizomatous herbs; leaves mostly
basal, large, ensiform, coriaceous; inflorescence a
much branched, densely hairy panicle of spikes;
tepals white, united at the base in a short cupular
tube; anther straight; ovary pubescent, ::±:: actino-
morphic; fruit berrylike, apparently indehiscent;
seeds 1.5-3 mm long. Two spp., NE Australia, New
Guinea, Moluccas, in rainforest.
2. Orthothylax Skottsb.
Orthothylax Skottsb., Bot. Jahrb. Syst. 65: 264 (1932), and in
Kew Bull. 3: 97-99 (1934).
394 Philydraceae

Tall, stoutly rhizomatous herb; leaves ::±:: basal,
ensiform, coriaceous; inflorescence a much
branched, glabrous or softly pubescent panicle
composed of spikes; tepals white to pink, free; fila-
ment short, anther straight; ovary (sub- )glabrous,
strongly zygomorphic; fruit capsular; seeds 3.5-
5 mm long, with elongated, often twisted chalaza!
process. 2n = 34. Only one sp., 0. glaberrimus
(Hook. f.) Skottsb., montane rainforests of E
Australia.
3. Philydrum Banks ex Gaertner Fig. 91
Philydrum Banks ex Gaertner, Fruct. 1: 62, t. 16 (1788).
Tall herb with short rhizome; leaves mostly basal,
rounded-ensiform, thick, soft, glabrous. Inflores-
cence a terminal composed spike, its axis and
bracts lanuginose; flowers numerous; tepals yel-
low, thin, the outer free, the inner nearly free from
flattened filament; anther with helically twisted
thecae; ovary imperfectly 3-locular; capsule with
the placentae on the middle of the 3 valves; seeds
numerous, < 1 mm long, testa spirally striate,
chalaza! cape enlarged. 2n = 16. Only one sp.,
P. lanuginosum Banks ex Gaertner, E and N
Australia, S, SE and E Asia from New Guinea to
Japan.
Dahlgren, R. et al. 1985. See general references.
Erdtman, G. 1952. See general references.
Fahn, A. 1954. Metaxylem elements in some families of the
Monocotyledoneae. New Phytol. 53: 530-540.
Hamann, U. 1966. Embryologische, morphologisch-
anatomische und systematische Untersuchungen an
Philydraceen. Willdenowia, Beih. 4: 1-178, 8 Taf.
Harris, P.J., Hartley, R.D. 1980. See general references.
Linder, H.P., Kellog, E.A. 1995. Phylogenetic patterns in the
commelinid clade. In: Rudall, P.J., Cribb, P.J., Cutler, D.F.,
Humphries C.J. (eds.) Monocotyledons: systematics and
evolution, Vol. 2. Royal Botanic Gardens, Kew, pp. 473-496.
Pilger, R. 1930. Philydraceae. In: Engler, A., Prantl, K.,
Natiirliche Pfianzenfamilien, 2nd edn. 15a: 190-191.
Leipzig: W. Engelmann.
Skottsberg, C. 1932. Bemerkungen tiber die Philydraceen. Bot.
Jahrb. Syst. 65: 253-274.
Solereder, H., Meyer, F.J. 1929. Systematische Anatomie der
Monokotyledonen, Pt. 4. Berlin: Borntraeger.
Takhtajan, A.L. 1982. See general references.

4. Philydrella Caruel
Philydrella Caruel, Nuovo G. Bot. Ita!. 10: 91 (1878).
Pritzelia F. Muell. (1843).

Slender, rarely more than 20cm high cormiferous

herb; leaf basal, terete, mostly solitary; inflores-
cence a single, glabrous, few-flowered spike; tepals
yellow, thin; outer tepals free, inner basally fused
with filament; anther curved, reniform; ovary gla-
brous, nearly perfectly 3-locular; capsule valves
separated from central placental column; seeds
< 0.5 mm long, shortly acuminate at chalazal end;
testa indistinctly striate. Only one sp., P. pygmaea
(R. Br.) Caruel, SW Australia.

Selected Bibliography

Bate-Smith, E.C. 1968. See general references.

Bohm, B.A., Collins, F.W. 1975. Flavonoids of Philydrum
lanuginosum. Phytochemistry 14: 315-316.
Briggs, B. 1966. Chromosome numbers of some Australian
monocotyledons. Contrib. N. S. W. Nat!. Herb. 4: 24-34.
Chase, M.W. eta!. 1995. See general references.
Cheadle, V.I. 1968. Vessels in Haemodorales. Phytomor-
phology 18: 412-420.
 Pontederiaceae
Pontederiaceae
C.D.K. CooK
Pontederiaceae Kunth in Humb., Bonpl. & Kunth, Nova Gen.
Sp. 1, ed. 40: 265 (1816), nom. cons.
Submerged, floating or emergent, freshwater
annuals or perennials. Stems of two kinds, vegeta-
tive and flowering; vegetative stems indetermi-
nate, with many leaves, with or without sympodial
rhizomes or stolons; flowering stems determinate,
leafless, bearing a spathe and a terminal inflores-
cence. Leaves distichous or spiro-distichous,
"pseudo-whorled" in Hydrothrix (see below),
often scalelike on rhizomes and stolons, cauline
or basal, usually sheathed at the base; stipules
axillary or median, either large and enveloping the
stem, forming an ochrea, or reduced to a small
sheath or ligule or absent; juvenile leaves simple
and linear; adult leaves either simple and linear or
clearly differentiated into petiole and blade; peti-
oles of variable length, sometimes inflated; blades
simple, submerged, floating or emergent, gla-
brous, linear to orbicular or sagittate, with entire
margin, venation acrodromous. Inflorescence ter-
minal, a panicle, umbel-like, a spike, or flowers
paired or solitary, subtended by 2 opposed spathes
(bracts), one often leaflike, the other sheathing or
both leaflike or sheathing; peduncles glabrous or
glandular pubescent. Flowers bisexual, zygomor-
phic or almost actinomorphic, some trihetero-
stylous. Tepals usually 6, sometimes reduced to 4
or 3, equal or unequal, all petaloid, usually wither-
ing early, free almost to the base or united basally
into a tube, white, blue, pink mottled with yellow
or yellow. Stamens all alike or dimorphic, 6, 4, 3 or
1; filaments inserted on the perianth, similar or
dissimilar, elongate or swollen, straight or twisted,
glabrous or hairy; the hairs linear and smooth,
glandular, or moniliform; anthers similar or dis-
similar, 2-thecate, basifixed or auriculate and thus
appearing dorsifixed, dehiscing introrsely by lon-
gitudinal slits or through apical pores, rem~ining
straight or coiling, yellow, blue, brown or black.
Gynoecium of 3 united carpels; ovary superior,
3-locular, sometimes only 1 locule fertile; placen-
tation axile or appearing parietal with intrusive
placentas, ovules anatropous, crassinucellar,
bitegmic, solitary or numerous; style single,
395
simple and elongate, glabrous or hairy; stigma
relatively small, entire or 3-lobed, Dry. Well-
developed septal nectaries present in Eichhornia
and Pontederia. Fruit a capsule or a 1-seeded
nutlet enclosed in a specialized perianth base
(anthocarp). Seeds numerous or solitary, ovoid,
usually with longitudinal ribs; endosperm starchy;
embryo central, cylindrical, linear; cotyledon ter-
minal; plumule lateral.
A family comprising 9 genera and ± 33 species;
distributed mostly in the tropics but extending
into temperate zones as well; almost cosmopolitan
through introduction. The centre of diversity is
tropical and subtropical New World.
VEGETATIVE MORPHOLOGY. The seeds germinate
under water. Seedling morphology has been
described and reviewed by Tillich ( 1995); the seed-
lings are very similar to those ofPhilydraceae. The
juvenile leaves are linear without differentiation
into petiole and blade, described in detail for
Eichhornia by Richards ( 1983 ). Some species show
paedogenesis and retain these juvenile leaves into
maturity. The mature plants may be emergent,
floating or submerged, with or without rhizomes
or stolons (Fig. 92). All species branch sympo-
dially, the development of "renewal shoots" is
described by Richards (1982). The roots are either
fixed in the substrate or free-floating. The leaves
are basically distichous, but the internodes may be
twisted so that the leaves appear to be spirally
arranged or the stem is truncate in parts so that
the leaves develop in rosettes. Leaf shape, even
within species, is highly plastic. In most species
the leaf sheath is laterally expanded, usually form-
ing an open or sometimes closed ochrealike struc-
ture sometimes with a variable number of free
lobes above, or forming an almost free ligule
(Schwartz 1926). The petioles are also euryplastic;
in Eichhornia crassipes (Fig. 93) they are often
swollen and function as floats. In Hydrothrix (Fig.
94) the leaves appear to be whorled; Rutishauser
(1983) refutes Goebel's theory that the leaf whorls
represent short shoots with annular insertion
along the long shoot and suggests, instead, that
the whorls are composed of leaves of the same
shoot order. Only the first leaf ( = the main leaf) of
the whorl starts as an outgrowth of the shoot apex
itself; the remaining whorl leaves are then prod-
ucts of an intercalary meristematic ring below the
real shoot apex, but with about the same morpho-
genetic potentiality.
VEGETATIVE ANATOMY. Cheadle (1970) studied
the vessel elements in 5 genera and 12 species.
396
4\ F
Fig. 92A-F. Pontederiaceae. A-D Pontederia cordata. A
Habit. B Inflorescence. C Flower in longitudinal section. D
Fruit. E, F Reussia rotundifolia. E Habit. F Fruit. (Orig. C.D.K.
Cook)
Vessels are mostly confined to roots but some
were found in the stems of Eichhornia and
Heteranthera. The leaves are without vessels. The
vessels have obliquely orientated, scalariform per-
foration plates with few to many perforations.
This elaborate structure has been used as evidence
Pontederiaceae
Fig. 93A-E. Pontederiaceae. A, B Eichhornia crassipes. A
Habit. B Flower in longitudinal section. C-E Monochoria
vagina/is. C Inflorescence. D Habit, with fruiting shoot. E
Flower. (Orig. C.D.K. Cook)
to support the theory that the aquatic habit in the
Pontederiaceae is not "primary'' and that they had
a "recent" terrestrial ancestor. The starch grains in
the vegetative parts are simple (centric and oval)
as well as compound (Hamman 1961). Trichomes,
when found, are associated with the inflorescence
and flowers. The stomata are of the paracyctic
 Pontederiaceae
kind. Tanniniferous idioblasts have been reported
from the petioles. Oxalate raphides are widely
present and cells with 1 or 2 prismatic crystals
have been found in some genera Hegnauer (1963).
INFLORESCENCE STRUCTURE. The inflorescences
are borne terminally and subtended by 2 spathes
(bracts) the lower of which is often leaflike or par-
tially leaflike, the upper sheathing and enveloping
the whole axis but they may both be leaflike or
scalelike. The lower spathe sometimes has a long
397
Fig. 94A-H. Pontederiaceae. A, B Heteranthera zosterifolia. A
Habit. B Inflorescence. C, D Zosterella dubia. C Habit. D Style
and stamens. E, F Hydrothrix gardneri. E Habit. F Flower. G, H
Eurystemon mexicanum. G Habit. H Flower. (Orig. C.D.K.
Cook)
sheathing stalk so the inflorescence often appears
to arise from a petiole. In the larger emergent spe-
cies the inflorescences are panicle-like, umbel-like
or spikelike; however, the flowers rarely open in
a simple sequence, so one may assume they are
398 Pontederiaceae
reduced from more complex types (Richards and
Barrett 1984; Cook 1989). In some more aquatic
species the inflorescences are reduced to paired or
solitary flowers. In Hydrothrix (Rutishauser 1983)
the paired flowers are mirror images of each other
and are pressed together within the upper spathe
(bract) making up a single functional unit
(pseudanthium). In some species the pedicels are
accrescent and grow downwards forcing the
developing fruits under water or in to mud, in other
species the peduncle merely bends downwards.
FLORAL STRUCTURE AND ANATOMY. The floral
structure is of the basic type for petaloid mono-
cotyledons. The tepals are usually petaloid and 3
+ 3 in number, but in some genera, such as
Monochoria, the outer whorl may be greenish and
somewhat sepal-like. In the large-flowered species
the flowers are distinctly zygomorphic and 2-
lipped, frequently with an enlarged adaxial tepal
with a yellow nectar guide. Septal nectaries occur,
at least, in Eichhornia and Pontederia (Daumann
1965). The flower usually wilts after 1 day. In
Pontederia and Reussia the perianth tube becomes
hardened and develops ridges with wings, teeth or
spines after anthesis and develops into a utricle
surrounding the fruit (anthocarp).
The floral anatomy, particularly of Eichhornia
crassipes has been studied in detail by Singh
(1962). Each tepal receives 3 vascular traces which
divide repeatedly; a number of smaller vascular
bundles in the midrib region deviate from the
others in being inverted. Calcium oxalate crystals
are found in perianth, stamen and carpel tissue.
The stigmatic surface is of the Dry type.
EMBRYOLOGY. The anthers are tetrasporangiate.
The endothecial cells have the girdle type of wall
thickenings and an amoeboid periplasmodial
tapetum, in which the cells of the early tapetum
are 1- or 2-nucleate. Microsporogenesis is succes-
sive, and the pollen tetrads isobilateral. The pollen
grains are monads and di- or trinucleate when
shed.
The ovules are anatropous, bitegmic and
crassinucellate. A parietal cell is cut off from the
archesporia! cell and gives rise to parietal tissue.
Embryosac formation is of the Polygonum type
and endosperm formation is helobial, with a small
chalaza! chamber where cell wall formation begins
much earlier than in the large micropylar chamber
(Davis 1966). Remarkable endosperm haustoria
have been described for Monochoria (Banerji and
Haldar 1942), where the micropylar chamber gives
rise to 2 lateral outgrowths 1 on each side of the
chalaza! chamber, growing downward and invad-
ing the chalaza! tissue. Subsequently the main
body of the micropylar chamber elongates and
unites with the haustoria to form a continuous
mass of endosperm cells with the chalaza! cham-
ber still recognisable at the base. The significance
of these haustoria is not known. The micropylar
haustorium is starchy. Embryogeny is of the
Asterad type.
POLLEN MORPHOLOGY. The pollen grains are hi-
or rarely trisulculate with 1 to 3 distal or subequa-
torial colpi (Rosatti 1987). The sexine is as thick
as the nexine or thicker, sometimes more or less
areoloidate. Exine sculpturing is verrucate, mi-
nutely, scabrate to finely reticulate. Plants with
dimorphic anthers have equally fertile, monomor-
phic pollen grains (Wang et al. 1995). The
triheterostylic species have trimorphic pollen;
the pollen grains are, of course, dimorphic within
a single flower. The pollen of Pontederiaceae is
very similar to that of Haemodoraceae according
to Simpson (1987).
KARYOLOGY. The base number of x = 8 is gener-
ally accepted for the family (Eckenwalder and
Barrett 1986) but both polyploidy and aneuploidy
have operated on the original base number. Three
species of Pontederia have been examined and all
found to be diploid with n = 8. In the closely
related Reussia the only species examined is tetra-
ploid, n = 16. Eichhornia paniculata and E.
paradoxa are diploid, with 2n = 16, while E.
azurea is 2n = 32 and E. crassipes is usually 2n =
32 but with isolated counts of n = 9, 2n = 30 and
2n = 58. The genus Monochoria has been exam-
ined several times (Cook 1989) and the following
counts have been reported: n = 14, n = 30, n = 40
(X4), n = 42, 2n = 26, 2n = 28 (X5), 2n = 52
(X6), 2n = 72, 2n = 74, 2n = 80 (X3) and 2n = 34
to 82. In Heteranthera n = 7 or 8. (2n = 14 in
H. limosa and H. rotundifolia, 2n = 32 in H.
multiflora, 2n = 48 in H. reniformis). The number
2n = 30 has been reported for Zosterella dubia.
POLLINATION AND REPRODUCTIVE SYSTEMS.
Cleistogamous flowers are frequently produced
on underwater stems in Hydrothrix, Scholleropsis,
Zosterella and some species of Eichhornia and
Heteranthera as well as emergent chasmogamous
flowers. The emergent flowers of all species are
relatively showy. Some species of Eichhornia and
Pontederia have well-developed septal nectaries
(Daumann 1965) which are certainly lacking in
Hydrothrix, Monochoria and perhaps also in
 Pontederiaceae
Heteranthera and Zosterella. It is assumed that all
chasmogamous flowers are pollinated by insects.
However, even in its native range, Eichhornia
crassipes is not visited very frequently and seed set
is often poor (Barrett 1980). Insect visitors include
nectar and pollen collecting bees. Eichhornia
azurea is visited by long-tongued solitary bees
(Anthophoridae and Megachilidae) and by small
pollen-collecting halictid bees (Barrett 1978).
Pontederia cordata is pollinated by bumble bees,
solitary bees and butterflies (Price and Barrett
1982).
Monochoria has somatic enantiostyly (left- and
right-handed flowers) accompanied by stamen di-
morphism (1 anther large and blue, the remaining
5 small and yellow). The flowers are visited mainly
by bees (Wang et al. 1995). The large blue anther
touches the same position of the bee's abdomen as
the style of the other morph, thereby apparently
facilitating intermorph pollination.
Triheterostyly occurs in all species of Reussia,
most species of Pontederia and three species of
Eichhornia. The biology of triheterostyly has re-
ceived considerable attention, the following are
some selected references: Eichhornia - Barrett
(1988), Barrett and Harder (1992); Pontederia
(including Reussia) - Barrett et al. (1983), Glover
and Barrett (1983, 1986), Price and Barrett (1984).
Homostyly or semi-homostyly has been observed
in several species of Eichhornia and in Pontederia
parviflora. Triheterostyly is accompanied by
incomplete physiological incompatibility. The
strongest incompatibility is found in short-styled
plants and the weakest in mid-styled plants.
Natural populations of Reussia and Eichhornia
usually contain unequal proportions of flower
morphs, often only a single morph.
FRUIT AND SEED. Most species have regularly or
irregularly dehiscent capsules. The capsules are
often shed before they open. In some species the
flowering stalks bend down after anthesis and
seeds will mature only in submerged carpels. In
most other members the seeds ripen in emergent
or submerged carpels. The functional disseminule
is often an unopened capsule. The seeds, at least
in Eichhornia and Heteranthera, remain viable
for many years if stored in wet anaerobic
conditions.
The genera Pontederia and Reussia have nutlets
surrounded by postfloral-modified perianth bases
which become hardened, rigid and in some spe-
cies toothed or spiny. In this utricle the fruits may
float for at least 15 days; the spines and teeth may
play a role in zoochory.
399
The seed coat anatomy has been investigated in
Heteranthera, Monochoria and Pontederia (Coker
1907). The outer integument is made up of 2
distinct layers. The outer layer has uniformly box-
shaped cells, which are elongated in the longitudi-
nal axis of the seed. The inner cells are highly
modified, being thin and elongated in the axis per-
pendicular to the wings of the seed. The walls at
either end of each cell are much taller, producing
the characteristic wings. The inner integument
becomes crushed and is deeply staining. The
embryo is straight, cylindrical and relatively large,
nearly reaching the length of the endosperm.
DISPERSAL AND REPRODUCTIVE BIOLOGY. Sev-
eral species are annuals which grow in tempo-
rary bodies of water; these are self-compatible and
often autogamous relying on seed for effective re-
production. Most species of Pontederia, Reussia
and some of Eichhornia are long-lived, partially
or wholly self-incompatible perennials. Seed set is
often poor in these species either because suitable
floral morphs are lacking or they are unsuccessful
in attracting pollinators. They spread by vegeta-
tive fragmentation. The vegetative reproduction
is efficient in spite of the lack of vegetative
propagules such as bulbils, corms, turions, etc.
The spread of Eichhornia crassipes, perhaps the
most troublesome weed in the world, is docu-
mented by Gopal (1987).
PHYTOCHEMISTRY. Hegnauer (1963, 1986) has
summarised the known, but somewhat erratic,
information on the family. Apart from common
phenolics, the presence of 0-methylated flavonols
and polyamines (the roots of Eichhornia crassipes
have blue anthocyaninlike cell wall-bound pig-
ments) are notable. Myrophyllin cells contain
inclusions of condensed tannins which have lost
their astringent properties.
SUBDIVISIONS OF AND RELATIONSHIPS WITHIN
THE FAMILY. The family is sometimes divided
into three tribes: Eichhornieae (trilocular cap-
sules), Pontederieae (1-seeded nutlets) and Heter-
anthereaa (unilocular capsules with numerous
seeds). Schwartz (1927, 1930) placed Monochoria
in the Heteranthereae but it certainly belongs
in the Eichhornieae on morphological grounds
and the genera Eichhornia and Monochoria share
a highly specific parasitic fungus Alternaria
eichhorniae Nag Ray et Ponnappa. The subdivi-
sion seems to reflect the phylogenetic relation-
ships within the family with the Heteranthereae
representing the most reduced or paedomorphic
400 Pontederiaceae

state (nearly actinomorphic flowers, tepals with- horniae Nag Ray et Ponnappa. Two weevils
out nectar guides and 3 stamens with basifixed (Coleoptera, Erirhiniae, Bagoini) Neochetina
anthers). Eckenwalder and Barrett (1986) made a bruchi Hustache and Neochetina eihhorniae
cladistic analysis of the family which was revalued Warner are very specific herbivores of Eichhor-
by Graham and Barrett (1995) (see also Barrett nia crassipes; these and a fungus Cercospora
and Graham 1997). They concluded that the rodmannii Conway are effectively used for bio-
Old World Pontederiaceae were derived from 4 logical control. Gopal (1987) has published a long
separate dispersals from S America to Africa at list of feeders, parasites and phytopathogens of
various times in the past. Triheterostyly is consid- Eichhornia.
ered to have arisen only once and is not the primi-
tive breeding system; dimorphic anthers and PALAEOBOTANY. The genus Pontederites Knowl-
triheterostyly represent different clades so one ton has been described from the Eocene in the
does not lead from or to another. Green River Formation of the USA. The upper
portion of a leaf fragment is preserved; it looks
AFFINITIES. Rosatti (1987) has evaluated the lit- like contenporaneous Pontederia. A fossil find in
erature on the affinities of the Pontederiaceae the Eocene Deccan Intertrappen beds of India has
based on morphological characteristics. The been described as a new species of Eichhornia by
combination of the presence of vessels with sca- Patil and Singh (1978); this is rather unlikely and
lariform perforations in the roots and sometimes the material could well be a Monochoria.
in the stems, UV-fluorescent ferulate in the cell
walls, oxalate raphides and styloids, helobial ECONOMIC IMPORTANCE. Eichhornia crassipes
endosperm with a diminutive chalazal chamber (water hyacinth) is one of the most troublesome
and amoeboid tapetum suggest relationships with weeds in the world (Gopal1987). In some regions
Philydraceae, Hamemodoraceae, Typhaceae and it is being replaced by even more noxious aquatics
Sparganiaceae, where all or most of these features such as Hydrilla verticillata and Salvinia molesta.
are present. However, affinities to the Com- Attempts are being made to utilise E. crassipes as
melinales, Bromeliales and Juncaceae can also animal feed, mulch, compost, paper fibre or as a
be supported. In the summary by Chase et al. source of biogas, alcohol and numerous chemi-
(1995a) using rbcL DNA sequences the affinities cals, and also for pollution abatement. In some
are with Philydraceae, Haemodoraceae and regions such as SE Asia it is used for fodder, fish
Commelinaceae. food and compost and is even regarded as desir-
able. Ash from burning large quantities of
DISTRIBUTION AND HABITATS. Monochoria (Asia Eichhornia (probably E. azurea) which contains
and Africa) and Scholleropsis (Africa) are native to much K2 0, is used as a substitute for common
the Old World; Pontederia, Reussia, Eurystemon, salt by the Indians of the Amazon lowland
Hydrothrix and Zosterella are indigenous to the (K. Kubitzki, pers. comm.).
New World. Eichhornia and Heteranthera are Some species of Heteranthera are pernicious
mostly New World with E. natans and H. callifolia weeds in rice fields. They are difficult to eradicate
in Africa. Eichhornia crassipes, Monochoria chemically because they are sensitive to the same
vaginalis, Pontederia cordata and three species of chemicals as rice. The seeds remain viable for at
Heteranthera are widely naturalised outside their least 4 years even when dried or exposed to frost
native range. (Schiele 1988). Many species, such as water hya-
All species of Pontederiaceae are aquatic, either cinth and Pontederia spp., are widely cultivated
obligate submerged plants with emergent flowers as ornamentals. Some species of Heteranthera,
of free-floating or bottom-rooted emergents. Zosterella, Hydrothrix and Eichhornia are grown
They are found in a wide range of oligotrophic as aquarium plants. Monochoria is used in medi-
to eutrophic freshwater habitats. Annual species cine in India and particularly against asthma and
are restricted to seasonally wet sites, the perenni- toothache; the leaves are eaten by humans in SE
als usually occur in permanent water. Some Asia.
species of Monochoria may occur in saline or
brackish habitats. Many species are gregarious
and may dominate large areas.

PARASITES. Eichhornia and Monochoria share a

highly specific parasitic fungus Alternaria eich-
 Pontederiaceae
KET TO THE GENERA
1. Leaves filiform, appearing whorled; functional stamen 1
7. Hydrothrix
- Leaves at least partly flattened, never filiform, alternate
along the stems or in rosettes, never in whorls; functional
stamens 3 or 6
2. Fruit a 1-seeded nutlet, enclosed by the persistent and
hardened base of the perianth tube (anthocarp)
- Fruit a 3-locular, many-seeded capsule, not enclosed by the
base of a persistent or modified perianth tube
3. Anthocarp wall toothed, ridged or winged but not spiny;
flowering stems erect 1. Pontederia
- Anthocarp wall with spiny ridges; flowering stems prostrate
2. Reussia
4. Stamens 6
- Stamens 3
5. Perianth distinctly tubular at base, distinctly zygomorphic;
anthers dorsifixed, subequal 3. Eichhornia
- Perianth divided almost at base, nearly actinomorphic;
anthers basifixed, 1 much larger (blue) than the other 5
(yellow) 4. Monochoria
6. Perianth of 4 (rarely 3) segments; at least the upper leaves
clearly differentiated into petiole and blade
9. Scholleropsis
- Perianth of 6 segments; leaves differentiated or not into
petiole and blade
7. Filaments of the 2 short stamens obliquely globular-
inflated; leaf with a small axillary ligule; perianth
zygomorphic 8. Eurystemon
- Filaments not markedly inflated; leaf sheath open, envelop-
ing the stem; perianth actinomorphic or nearly so
8. Anthers dissimilar, straight; spathe petiolate, not enclosing
the perianth tube; seeds numerous; perianth white or blue
5. Heteranthera
- Anthers all alike, curved after anthesis; spathe sessile, en-
closing perianth tube; seeds relatively few; perianth yellow
6. Zosterella
1. Pontederia L. Fig. 92A-D
Pontederia L., Sp. Pl. 1:288 (1753); Lowden, Rhodora 75:426-
483 (1973), rev.
Floating or rooted, creeping or ascending perenni-
als with long rhizomes or stolons. Leaves scalelike,
linear or petiolate, estipulate; blades elliptic to
ovate or sagittate. Inflorescence a complex spike
subtended by 2 dissimilar spathes. Tepals blue
to purple or whitish, basally tubular, 2-lipped;
abaxial 3 lobes partly united, the adaxial lobes
subequal and free. Stamens 6, the 3 upper shorter
than the 3 lower; filaments with glandular hairs.
Ovary with only 1 fertile locule. Fruit a 1-seeded
anthocarp with smooth or toothed ridges. Six spp.,
Lowden (1973) recognised only three spp. and two
varieties, America from Canada (50° N) to Argen-
tina (35° S).
401
2. Reussia Endl. Fig. 92E,F
Reussia End!., Gen. Pl. 1: 139 (1836), nom. cons. -Included as
subgen. in Pontederia by Lowden (1973); Lowden, Rhodora
75: 426-483 (1973), rev.
Submerged, floating or rooted, creeping or as-
3 cending annuals or perennials. Rhizomes short.
Flowering stems prostrate. Leaves scalelike or
4 elongate, linear or petiolate; blades lanceolate to
orbicular, reniform or sagittate. Inflorescence a
spike, subtended by 2 dissimilar spathes. Tepals
pale blue to purple or pinkish, basally tubular, 2-
5 lipped; adaxial lip large, simple; the abaxial with 5
6 subequallobes. Stamens 6; filaments with glandu-
lar hairs; anthers dorsifixed. Ovary with only 1
fertile locule. Fruit a 1-seeded anthocarp with
spiny ridges. Two spp., Central America to E
Argentina.
3. Eichhornia Kunth Fig. 93A,B
7 Eichhornia Kunth, Eichhornia, genus novum [Diss.]: 3 (1842),
Enum. Pl. 4: 129 (1843), nom. cons.
Submerged or floating, creeping or stoloniferous
8 annuals or perennials. Stems rhizomatous or
erect. Leaves basal or cauline, linear or differenti-
ated into petiole and blade; petiole occasionally
inflated; blade linear to cordate. Inflorescence a
panicle or complex spike; the lower spathe leaflike
or reduced to an inflated base, the upper scalelike.
Tepals blue to somewhat pinkish, basally tubular,
6-lobed; adaxial lobe largest or lobes subequal,
with glandular hairs. Stamens 6, the 3 upper ones
shorter than the 3lower ones; filaments with glan-
dular hairs. Fruit a capsule; seeds numerous, lon-
gitudinally winged. Seven or eight spp., tropical
America, one in Africa, E. crassipes (Mart.) Solms-
Laub., introduced to warmer regions throughout
the world.
4. Monochoria K.B. Presl Fig. 93C,E
Monochoria K.B. Pres!, Rei. Haenk. 1(2): 127 (1827); Cook in
Kit Tan (Ed.), The Davis and Hedge Festschrift. Edinburgh
University Press: 149-184 (1989), rev.
Annuals or perennials, in water or wet places.
Stems truncate and sometimes cormlike or
elongate and creeping. Leaves radical, spiro-
distichous; juvenile leaves submerged, linear or
scalelike; adult leaves emergent; petioles erect
or flaccid and floating, with large, ochrealike
stipules (enveloping the stem and developing
leaves); blades linear-lanceolate to ovate, cordate,
402 Pontederiaceae
sagittate or hastate. Inflorescence paniculate,
raceme- or umbel-like; both spathes leaflike
or lower leaflike and the upper scalelike (some-
times included within the lower one); the lower
spathe sometimes dominant the inflorescence
appearing to burst out of its petiole. Tepals blue
or outer 3 greenish, lobes free almost to base,
subequal. Stamens 6; filaments simple or 1 with 1
or rarely 2 lateral, oblique teeth; anthers unequal,
5 small and yellow, 1 large and blue, the blue
anther lateral at anthesis, the yellow ones in a
cluster above the ovary. Fruit a 3-valved capsule.
Seeds numerous, with 8-14 longitudinal nerves,
ribs or wings. Eight spp. (one new sp. recently
described from Hainan, China, is doubtfully dis-
tinct), tropical to warm temperate Africa (two
spp.), Asia (four spp.) and Australia (two spp.);
M. vaginalis K.B. Presl naturalised in Europe
(Ukraine and Italy), N America, Hawaii (Oahu)
and Australia.
5. Heteranthera Ruiz et Pavon Fig. 94A,B
Heteranthera Ruiz et Pavon, Fl. Peruv. Chi!. Prodr. 9 (1794),
nom. cons.; Horn, C.N. PhD. dissertation, Univ. Alabama
1-260 (1986), rev.
Submerged, floating or emergent, annuals or per-
ennials. Stems creeping or ascending. Leaves with
or without stipules, linear or differentiated into
petiole and blade; blades submerged, floating or
emergent, linear to ovate, reniform or cordate.
Inflorescence a spike or flowers paired, rarely soli-
tary. Flowers chasmogamous or the lower often
cleistogamous, chasmogamous flowers pedicel-
late, borne above the spathe. Tepals blue or white,
basally tubular or lobed almost to base; lobes 6,
subequal. Stamens 3 (or sometimes 1 in cleistoga-
mous flowers), one larger than the others, straight.
Ovary with 3 ± intrusive placentas, appearing 1-
locular. Fruit a loculicidal capsule. Seeds numer-
ous, longitudinally ribbed. Ten spp., tropical and
temperate America and tropical Africa; H. limosa
(Sw.) Willd., H. reniformis Ruiz et Pavon and
H. rotundifolia (Kunth) Griseb. naturalised in
Europe.
6. Zosterella J.K. Small
Zosterella J.K. Small in Small and Carter, Fl. Lancaster
Co. (Pennsylvania): 68 (1913). Included as subgen. in
Heteranthera by Rosatti (1987); Horn, C.N. PhD. disserta-
tion, Univ. Alabama 1-260 (1986), rev.
Submerged perennials or perhaps annuals. Stems
elongate, flaccid. Leaves cauline, linear. Flowers
solitary, emergent or submersed; partly enclosed
by the tubular upper spathe. Tepals yellow, basally
tubular, with 6 subequal lobes. Stamens 3;
filaments recurving after anthesis; anthers alike,
coiling. Ovary appearing 1-locular. Fruit an ir-
regularly dehiscent capsule. Seeds few. Two spp.,
temperate and subtropical N America.
7. Hydrothrix J.D. Hook. Fig. 94E,F
Hydrothrix J.D. Hook., Ann. Bot. 1: 89 (1887).
Submerged annual or perhaps sometimes peren-
nial. Stems elongate and flaccid. Leaves cauline,
filamentous in pseudowhorls of 7-30. Flowers
submerged and cleistogamous or emergent and
chasmogamous, paired (the 2 flowers are a mirror
images resembling and functioning as a single
flower), each pair subtended by 2, similar, green
spathes. Tepals 6, basally tubular; the outer 3lobes
large, yellow and petaloid; the inner 3 lobes fila-
mentous. Stamens 3, 1 functional, 2 staminodial
and filamentous. Ovary appearing 1-locular. Fruit
a lodicidal capsule. Seeds up to ±40. Only one sp.,
H. gardneri J.D. Hook., Ceara, NE Brazil.
8. Eurystemon Alexander Fig. 94G,H
Eurystemon Alexander, N. Am. Fl. 19 (19): 55 (1937).
Emergent annual. Stems contracted, corm-like.
Leaves radical, linear, grasslike, with ligular pro-
jection from the sheath. Inflorescence an erect
spike, subtended by 2 leaflike spathes. Tepals 6,
blue and white, basally tubular, zygomorphic;
adaxial lobes 5, erect to spreading; abaxial lobe
smaller, deflexed. Stamens 3, unequal, 1 large and
2 small; filaments globosely inflated at middle.
Fruit a capsule. Seeds numerous. Only one sp.,
E. mexicanum (S. Watson) Alexander, northern
Central America.
9. Scholleropsis H. Perrier
Scholleropsis H. Perrier, Not. Syst. (Paris) 5: 158 (1935).
Submerged, floating or sometimes terrestrial,
probably annual. Stems elongate, flaccid, branch-
ing. Leaves heterophyllous; basal leaves radical,
submerged, linear, to 7 em long; cauline leaves
linear or differentiated into an elongate petiole
and elliptic to ovate floating blade. Flowers soli-
tary, cleistogamous and sessile when submerged,
chasmogamous and pedicellate when emergent;
the lower spathe long-stalked; the upper spathe
 Pontederiaceae
sheath-like. Tepals (3- )4, basally tubular; the
lobes subequal, yellow. Stamens 3, I large, 2 small.
Fruit an irregularly dehiscent capsule. Seeds nu-
merous. Only one sp., S. lutea H. Perrier, Mada-
gascar and N Cameroon by Lake Chad.
Selected Bibliography
Banerji, I., Halder, S. 1942. A contribution to the morphology
and cytology of Monochoria hastaefolia Pres!. Proc. Indian
Acad. Sci. 16 B: 91-106.
Barrett, S.C.H. 1977. The breeding system of Pontederia
rotundifolia, a tristylous species. New. Phytol. 78: 209-
220.
Barrett, S.C.H. 1978. Floral biology of Eichhornia azurea
(Swartz) Kunth (Pontederiaceae). Aquat. Bot. 5: 217-228.
Barrett, S.C.H. 1980. Sexual reproduction in Eichhornia
crassipes (water hyacinth) II. Seed production in natural
populations. J. Appl. Ecol. 17: 113-124.
Barrett, S.C.H. 1988. Evolution of breeding systems in
Eichhornia: a review. Ann. Mo. Bot. Gard. 75: 741-760.
Barrett, S.C.H., Graham, S.W. 1997. Adaptive radiation in the
aquatic plant family Pontederiaceae: insights from phyloge-
netic analysis. In: Givnish, T.J., Sytsma, K.J. (eds.) Molec-
ular evolution and adaptive radiation. Cambridge:
Cambridge University Press, pp. 225-228.
Barrett, S.C.H., Harder, L.D. 1992. Floral variation in
Eichhornia paniculata (Spreng.) Solms II: effects of devel-
opment and environment on the formation of selfing
flowers. J. Evol. Bioi. 5: 83-107.
Barrett, C.C.H., Price, S.D., Shore, J.S. 1983. Male fertility and
anisoplethic population structure in tristylous Pontederia
cordata (Pontederiaceae). Evolution 37: 745-759.
Chase, M.W. et al. 1995a. See general references.
Cheadle, V.I. 1970. Vessels in Pontederiaceae, Ruscaceae,
Smilacaceae, and Trilliaceae. In: Robson, N.K.B., Cutler,
D.P., Gregory, M. (eds.) New research in plant anatomy.
Supplement 1 to J. Linn. Soc. Bot. 63: 45-50.
Coker, W.C. 1907. The development of the seed in the
Pontederiaceae. Bot. Gaz. 44: 293-301.
Cook, C.D.K. 1989. Taxonomic revision of Monochoria
(Pontederiaceae). Kit Tan (ed.) The Davis and Hedge
Festschrift. Edinburgh: Edinburgh University Press, pp.
149-184.
Dahlgren, R.M.T. et al. 1985. See general references.
Daumann, E. 1965. Das Bliitennektarium bei den Ponte-
deriaceen und die systematische Stellung dieser Familie.
Preslia 37: 407-412.
Davis, G.L. 1966. Systematic embryology of the angiosperms.
New York: Wiley.
Eckenwalder, J.E., Barrett, S.C.H. 1986. Phylogenetic systemat-
ics of Pontederiaceae. Syst. Bot. 11: 373-391.
Glover, D.E., Barrett, S.C.H. 1983. Trimorphic incompatibility
in Mexican populations of Pontederia sagittata. New
Phytol. 95: 439-455.
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404 Posidoniaceae

Posidoniaceae The leaves have open sheaths with distinct

J. Kuo and A.J. McCOMB
Posidoniaceae Lotsy, Vortr. Bot. Stammesgesch. 3: 658 (1911),
nom. cons.
Submersed marine or occasionally estuarine
robust perennial herbs with creeping, monopodi-
ally branched, cylindrical to flattened rhizomes,
the latter bearing a scale at each node and covered
with fibrous remains of old sheaths. Roots
branched, root hairs rare. Leaves alternate, dis-
tichously arranged on the rhizome, with distinct
blade and sheathing base. Blade linear, flat, or bi-
convex to terete, with 3-20 or more longitudinal
vascular bundles; squamules numerous; tannin
cells numerous; stomata absent. Inflorescence
racemose, spikelike on a long, flattened peduncle.
Flowers actinomorphic, hermaphroditic, perianth-
less; stamens 3; anthers sessile, tetrasporangiate,
pollen filiform; pollination hydrophilous; carpel
1 with a sessile ornate stigma; ovule 1. Fruit with
spongy pericarp, dehiscent. Testa membranous;
hypocotyl enlarged storing mainly starch.
One genus of nine species, one in the Mediterra-
nean, the other eight from the temperate to sub-
tropical coasts of Australia.
strap-shaped to filiform blades. The sheath is
amplexicaul, ligulate, auriculate and persistent
after the fall of the blade. The leaf blade is flattened
or biconvex to terete, with a rounded to truncate
or spathulate apex, and contains three to
more than twenty longitudinal, parallel vascular
bundles connected periodically by transverse
veins (Kuo 1978; Cambridge and Kuo 1979; Kuo
and Cambridge 1984). The blade epidermal cells
are uniformly chlorenchymatous, rectangular to
tall and spindle-shaped in transverse section. The
cell walls do not have ingrowths, and the epider-
mis is covered by a thin, porous cuticle (Kuo
1978). The mesophyll tissue is of homogeneous,
highly vacuolated parenchyma cells containing a
few chloroplasts. Lacunae are present, but less
prominent than in other seagrass families; they
have regular, transverse diaphragms.
Each longitudinal vascular bundle of the leaf is
enclosed by a distinct layer of sheath cells with

VEGETATIVE MORPHOLOGY AND ANATOMY. The

rhizomes are cylindrical to laterally compressed
and are covered with persistent, fibrous re-
mains of old leaf sheaths (Fig. 95). The rhizome
internode has a conspicuous stele with 4 to 10
cortical vascular bundles, numerous lignified
fibrous strands, tannin cells and some small starch
grains in cortical parenchyma cells. It has both
exodermis and epidermis (Kuo and Cambridge
1978, 1984; Albergoni et al. 1978). Branching is
monopodia} and irregular (Tomlinson 1974,
1982).
Each node bears 1 or 2 roots, which are long and
branched; root hairs are rarely seen. Each root
has a well-defined stele surrounded by an endo-
dermis with a distinct Casparian strip (Kuo and
Cambridge 1978). Large lacunae are embedded in
the cortex. The exodermis consists of 1 or 2 layers
of cells with thickened, suberised but not lignified
walls. There are numerous bacteria and fungi in Fig. 95A-E. Posidoniaceae. Posidonia australis. A Habit. B
the rhizosphere (Kuo et al. 1981). Part of rhizome. C Flower. D, E Fruits. (den Hartog 1970)
 Posidoniaceae 405

thin, lignified walls, and contains several sieve callose deposited only at the wall junctions with
tubes with thin walls and typical monocotyledon- the pollen tube.
ous proplastids with protein crystalloids. The
xylem is represented by several tracheids which EMBRYOLOGY. Endosperm formation is helobial.
have little secondary wall thickening and are The embryo has an elongated hypocotyl rich in
poorly lignified. Vascular parenchyma cells lack starch, a well-developed plumule and a short
cell-wall ingrowths (Kuo 1978, 1983). Fibrous radicle, situated at the distal end of the hypocotyl.
strands of thick-walled, unlignified cells are abun-
dant in the leaf margins and in the hypodermal POLLEN MORPHOLOGY AND POLLINATION. The
and subhypodermallayers in those species having pollen is filiform and comparatively short and
flattened blades, but in addition, occur deep in the has a swollen central region, lacking an exine
mesophyll tissue in species with biconvex to terete (McConchie and Knox 1989a). Posidonia has no
blades (Ostenfeld 1916; Cambridge and Kuo 1979; pollen-stigma recognition system to prevent self-
Kuo and Cambridge 1984). Squamules are numer- pollination (McConchie and Knox 1989b).
ous and palisadelike.
The leaf sheath differs from the blade in having KARYOLOGY. All species have 2n = 20, with 5
a non-porous cuticle and lacking chloroplasts in larger and 5 smaller pairs of chromosomes (den
the epidermis and mesophyll. Fibrous strands are Hartog et al. 1987; Kuo et al. 1990; Semroud et al.
more pronounced in the sheath than the blade. In 1992).
the sheath these strands, apart from those associ-
ated with the adaxial epidermis, are lignified and FRUIT AND SEED. Fruits are soft when shed, with
persistent on the rhizomes (Kuo 1978). a fleshy pericarp, and are buoyant. Being fleshy
Water depth apparently affects some morpho- and dehiscent, the fruit is not readily classified as
logical and anatomical features of Posidonia drupe, berry or achene. Each fruit splits longitudi-
oceanica such as leaf blade thickness, tannin cell nally and releases a single developing seed, which
and fibre frequency (Colombo et al. 1983). The sinks to the bottom. The seed is covered by a
reproductive cycle seems to be related to a delay in thin membrane but lacks a distinct seed coat.
leaf production in deeper plants. There is little or no dormancy phase. The main
Starch is present in the rhizome and leaves, and storage organ of the seed develops as an out-
the plants are rich in tannin cells. growth of the hypocotyl. A well-developed pri-
mary root is found on the seedlings of some
INFLORESCENCE STRUCTURE. Flowers are carried species (P. oceanica), but not others (P. australis,
on long peduncles. The axillary inflorescences are P. coriacea). The hypocotyl may remain attached
racemose and bear leafy bracts (Fig. 95A); the ulti- to the seedling for up to 18 months, but nutrient
mate units are spikelike, with several apparently storage may last only 8 months (Kuo and Kirkman
naked flowers (Cambridge and Kuo 1979; Kuo and 1996). Nutrient accumulation in fruits and its
Cambridge 1984; Kuo and McComb 1989). utilisation during seed germination and seedling
establishment have been described for two
FLORAL STRUCTURE. Flowers are actinomorphic, Australian Posidonia (Hocking et al. 1980, 1981).
bisexual, and lack a perianth. Each has 3 stamens
and 1 single carpel. Each stamen has 2 bilocular REPRODUCTION. In Australia, flowers are initi-
thecae attached to a broad connective. The carpel ated in autumn to winter, flowering occurs in
is ellipsoid with a short, irregularly lobed style and spring, and fruit is shed in late spring-early
a solitary atropous, pendulous ovule. The stigma summer. Flowering may be sparse to prolific.
is sessile and ornate (Fig. 95C). It has a surface Numerous pericarp remains accumulate on
pellicle, cuticle, and reacts positively for esterase beaches, but seeds germinate in ripple marks as
(McConchie and Knox 1989b). A diffuse secretion soon as they are detached, and seedlings develop.
covering the surface is thought to trigger preco- Few seedlings appear to survive, as most available
cious pollen germination, before pollen-stigma habitats are already occupied by established
contact (McConchie and Knox 1989a). When the meadows (Kuo and Kirkman 1996). Flowering and
pollen of Posidonia comes in contact with a recep- fruiting are common, but the establishment of
tive stigma, a meniscus is formed between the pol- seedlings is episodic in the Mediterranean species
len and stigma coatings. In Posidonia the pollen (Procaccini and Mazzella 1996). In contrast, veg-
tube is occluded by a plug of callose where it en- etative propagation is very important in the inva-
ters the pistil and the adjacent cells have lenses of sion of areas at the edges of seagrass meadows, in
406 Posidoniaceae
the colonisation of small areas disrupted by storm
action, and in ongoing meadow regeneration.
Leaves are being continually replaced at rates
which show seasonal trends at higher latitudes,
but are seasonal at lower latitudes.
PHYTOCHEMISTRY. Posidoniaceae have 2 or 3
sulphated phenolic acids, but sulphated flavones
are absent from the family, though present in
other seagrass families (McMillan et al. 1980a).
AFFINITIES. Molecular data indicate a close rela-
tionship between Cymodoceaceae, Posidoniaceae,
and Ruppiaceae (Waycott and Les 1996; Les et al.
1997).
DISTRIBUTION AND HABITATS. The distribution
of the family is strikingly disjunct. One Posidonia
species occurs in the Mediterranean, and the
others are from temperate Australia. In that conti-
nent, P. australis occurs along the southern,
southwestern and southeastern coastline, but
other species are restricted to southern and south-
western Australia.
In Australia Posidonia may occur in small
clumps, or as vast underwater meadows which
may be monospecific or consist of several
Posidonia species, interspersed with other
seagrasses, especially Amphibolis spp. They occur
in estuaries, sheltered embayments, and near-
shore, relatively protected oceanic regions; they
are almost invariably subtidal, and if the water is
clear, may occur down to 40m. They are ecologi-
cally important because of their role in supporting
food chains and stabilising sediments. There is
some direct grazing by echinoderms and fish, but
much of their importance lies in the production of
nutrient-rich organic detritus which is used by
filter feeding organisms. They support growth of
epiphytic plants and animals and infauna which
graze on epiphytes; they are important as a
nursery area for fish, and provide shelter
(Shepherd and Womersley 1981). Detached old
leaves, with materials from other seagrasses and
macroalgae, accumulate on beaches in wracks up
to 1m high for up to 4 months each year, decom-
posing in the wracks or surf zone, where they are
again important in food chains and nutrient
cycling. Persistent fibres are rolled into balls by
wave action, and such "marine balls" have been
known since ancient times (Cannon 1979).
PARASITES. The rhizomes of P. oceanica are
frequently infected with the parasitic Pyrenomy-
cete Halotthia posidoniae (Durieu & Montagne)
Kohlmeyer, and less often with Pontoporeia
bilurbinata (Durien & Montagne) Kohlmeyer.
Brearley and Walker (1996) observed that small
lynseiid isopods burrow through the leaf meso-
phyll, leaving the epidermis intact. Two species
of lynseiid isopods are specific, Lynseia annae
Cookson & Poore and L. dianae Cookson & Poore
being found in the P. australis group and P.
ostenfeldii group, respectively, but lynseeid iso-
pods are yet to be found in the Mediterranean
species.
PALAEOBOTANY. According to den Hartog (1970)
two Posidonia fossils are known, Posidonia
cretacea Hos. a.v.d. Mark from the Cretaceous,
and Posidonia parisiensis (Brongt.) Freitel ( =
Caulinites parisiensis Brongt. Posidonia
perforata Saporta & Marion) from the Eocene
of the Paris Basin. They have thick rhizomes
with short internodes and flattened, acuminate-
tipped leaves. However, the actual affinities of
remains of this type are far from certain (Daghlian
1981).
ECONOMIC IMPORTANCE. The main economic
importance is indirect, in the support of various
food chains which lead to commercially important
animals such as fish and prawns; the loss of
seagrass meadows after industrial or other devel-
opments is therefore viewed with concern.
Another indirect effect lies in the ability of
seagrass meadows to stabilise sediments and
damp wave action. Large submerine deposits of
persistent Posidonia fibres (e.g. in St. Vincent's
Gulf, S Australia) have been viewed as potential
sources of fibre for textile and paper manufacture,
and for insulation (Winterbottom 1917; Kuo and
Cambridge 1978).
 Posidoniaceae
Only one genus:
Posidonia Konig Fig. 95
Posidonia Konig, Ann. Bot. (Konig & Sims, London): 95
(1805).
Description as for family.
Nine spp., temperate and subtropical waters, all
endemic to Australia except P. oceanica (L.)
Delile, which is restricted to the Mediterranean.
Among the Australian species, two groups can be
distinguished: the P. australis group with mem-
branous leaves and growing in shallow, sheltered
waters; and the P. ostenfeldii group with leathery
leaves and occurring in deeper, more open waters.
Selected Bibliography
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sur l'anatomie de Posidonia oceanica (Zosteraceae). Plant
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Brearley, A., Walker, D.I. 1996. Burrow structure and effects
of burrowing isopods (Limnoriidae) in southerwestern
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Cambridge, M.L., Kuo, J. 1979. Posidonia sinuosa and P.
angustifolia (Posidoniaceae); two new taxa of seagrasses
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Cambridge, M.L., Kuo, J. 1982. Morphology, anatomy and his-
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Cannon, J.F.M. 1979. An experimental investigation of
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Colombo, P.M., Rascio, N., Cinelli, F. 1983. Posidonia oceanica
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Dahlian, C.P. 1981. A review of the fossil record of monocoty-
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Hartog, C. den, Hennen, J., Norten, Th. M.P.A., Van Wijk, R.J.
1987. Chromosome number of the European seagrasses.
Plant Syst. Evol. 156: 55-59.
Hocking, P.J., Cambridge, M.L., McComb, A.J. 1980. Nutrient
accumulation in the fruits of two species of seagrass,
Posidonia australis and Posidonia sinuosa. Ann. Bot. 45:
149-161.
Hocking, P.J., Cambridge, M.L., McComb, A.J. 1981. Nitrogen
and phosphorus nutrition of developing plants of two
seagrasses, Posidonia australis and Posidonia sinuosa.
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Kirkman, H., Kuo, J. 1996. Seagrasses of the southern coast of
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Kuo, J. 1978. Morphology, anatomy and histochemistry of the
Australian seagrasses genus Posidonia Konig (Posidonia-
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Kuo, J. 1983. Notes on the biology of Australian seagrasses.
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Kuo, J., Cambridge, M.L. 1978. Morphology, anatomy and his-
tochemistry of the Australian seagrasses genus Posidonia
Konig (Posidoniaceae ). II. Rhizomes and roots of Posidonia
australis Hook f. Aquat. Bot. 5: 191-206.
Kuo, J., Cambridge, M.L. 1984. A taxonomic study of
Posidonia ostenfeldii complex (Posidoniaceae) with de-
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Kuo, J., Kirkman, H. 1996. Seedling development of selected
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the biology of seagrasses with special reference to the
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Kuo, J., McComb, A.J., Cambridge, M.L. 1981. Ultrastructure
of the seagrass rhizosphere. New Phytol. 89: 139-143.
Kuo, J., James, S.H., Kirkman, H., Hartog, C. den 1990.
Chromosome numbers and their systematic implications in
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Larkum, A.W.D., McComb, A.J., Shephard, S.A. (eds.) 1989.
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Les, D.H. eta!. 1997. See general references.
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in the seagrass Posidionia australis. Ann. Bot. 63: 235-248.
McConchie, C.A., Knox, R.B. 1989b. Pollination and reproduc-
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A.J., Shephard, S.A. (eds.) Biology of seagrasses. A treatise
on the biology of seagrasses with special reference to the
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McMillan, C., Zapata, 0., Escobar, L. 1980a. Sulphated phe-
nolic compounds in seagrasses. Aquat. Bot. 8: 267-278.
McMillan, C., Parker, P.L., Fry, B. 1980b. C13 /C 12 ratios in
seagrasses. Aquat. Bot. 9: 237-249.
McMillan, C., Williams, S.C., Escobar, L., Zapata, 0. 1981.
Isozymes, secondary compounds and experimental cul-
tures of Australian seagrasses in Halophila, Halodule,
Zostera, Amphibolis and Posidonia. Aust. J. Bot. 29: 247-
260.
Ostenfeld, C.H. 1916. Contribution to Western Australian
botany, part I. Dan. Bot. Ark. 2: 1-44.
Phillips, R.C., McRoy, C.P. (eds) 1981. Handbook of seagrass
biology. New York: Berland, STPM Press.
Procaccini, G., Mazzella, L. 1996. Genetic variability and
reproduction in two Mediterranean seagrasses. In: Kuo, J.,
Phillips, R.C., Walker, D.I., Kirkman, H. (eds.) Seagrass
biology. Proceedings of an International Workshop,
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Perth: Science, UW A, pp. 85-92.
Robertson, E.L. 1984. Seagrasses, pp. 57-122. In: Womersley,
H.B.S. (ed.) The marine benthic flora of southern Australia,
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408 Posidoniaceae/Potamogetonaceae
1992. On a broad-leaved form of the seagrass Posidonia
oceanica (Posidoniaceae) from Algiers (Algeria). Aquat.
Bot. 43: 181-198.
Shepherd, S.A. Womersley, H.B.S. 1981. The algal and seagrass
ecology of Waterloo Bay, South Australia. Aquat. Bot. 11:
305-391.
Tomlinson, P.B. 1974. Vegetative morphology and meristem
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Aquaculture 4: 107-130.
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seagrasses. In: Metcalfe, C.R. (ed.) Anatomy of the mono-
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Waycott, M., Les, D.H. 1996. An integrated approach to the
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Walker, D.I., Kirkman, H. (eds.) Seagrass biology.
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Western Australia. 25-29 January 1996. Perth: Science,
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Potamogetonaceae
R.R HAYNES, D.H. LES, and L.B. HoLM-NIELSEN
Potamogetonaceae Dumort., Anal. Pam. Pl. 59, 61 (1829),
nom. cons.
Perennial or rarely annual floating-leaved or
submersed glabrous herbs growing in fresh
or brackish waters, sometimes stranded by reced-
ing water levels. Roots fibrous, few, nonseptate,
from lower nodes of stems. Stems slender,
branched or unbranched, often dimorphic, the
lower stems rhizomatous or stoloniferous, occa-
sionally with tubers, the upper erect, leafy, the tips
often modified into turions. Leaves alternate
and the upper 2-12 subopposite, or opposite
throughout, entire to serrulate, sessile, or peti-
olate, 1-many-veined, mostly ligulate; ligules
forming a tubular sheath around the stem, free
from or adnate to the base of the blade; veins
when several or numerous parallel or arching,
often connected by cross-veins; axillary infra-
vaginal scales present, membranous; submersed
leaves thin, linear to orbicular; floating blades
often leathery, lanceolate to elliptic or ovate.
Inflorescences pedunculate, axillary, or terminal,
with 2 opposite flowers or more than 2 flowers
in a capitate to interrupted spike, with a sub-
tending spathe. Flowers perfect, hypognous,
actinomorphic; tepals 4( -2), in one series, dis-
tinct, rounded, flat to convex, short -clawed,
greenish, reddish, or brownish. Stamens 4, rarely
2-3; anthers extrorse, 2-locular, dehiscing by
longitudinal slits; filaments adnate to the perianth
claw; pollen grains shed singly, spherical.
Gynoecium of (1-) 4 (-7) distinct carpels, the
carpels unilocular, sometimes with an indistinct
stipe; ovules solitary, attached to the ventral
margin of the carpel; styles short; stigmas
capitate. Fruits usually drupelets with a membra-
nous exocarp, fleshy mesocarp, and stony en-
docarp, or rarely berries, opening by decay of
the pericarp. Seeds solitary, exalbuminous, with
coiled embryo.
A subcosmopolitan family with 2 (or 3)
genera and 85-90 species. Additionally, 40-50
putative hybrids have been described, which
are concentrated predominantly in Great Britain,
Scandinavia, and Japan (Preston 1995).
 Potamogetonaceae
VEGETATIVE MoRPHOLOGY. The Potamogetona-
ceae are aquatic or rarely semiaquatic herbs grow-
ing either entirely submersed or with emergent
inflorescences and submersed leaves or with both
submersed and floating leaves and emergent inflo-
rescences (Figs. 96, 97). Those that are semi-
aquatic are stranded by receding water levels,
often appearing terrestrial. The roots are adventi-
Fig. 96A-I. Potamogetonaceae. A, B Potamogeton lucens. A
Fruiting shoot. B Drupelet. C- E P. perfo /iatus. C Fruiting
shoot. D Flower. E Drupelet. F- 1 Groenlandia densa. F Flower-
ing shoot. G Two-flowered spike. H Flower, from above. I
Drupelet. (Takhtajan 1982)
409
tious and unbranched, ansmg singly or, more
commonly, in whorls of up to 10 from the nodes of
rhizomes, stolons, or lower nodes of erect stems.
The stems often are divided into a short to elon-
gate, non-chlorophyllous rhizome and an erect
chlorophyllous stem. The rhizome varies in length
up to 2m or more. It may be clearly different from
the erect stem.
Fig. 97A-J. Potamogetonaceae. A-F Potam ogeton natans. A
Fruiting shoot. B Flowering spike. C Fruiting spike. D Flower.
E Tepa! with stamen. F Drupelet. G-H P. gram ineus. G Shoot
with submersed and floating leaves. H Wholly submersed
shoot. I, J P. a/pinus. I Flowering shoot. J Drupelet. (Takhtajan
1982)
410 Potamogetonaceae
Rhizomes lack chlorophyll and have scalelike,
non-chlorophyllous leaves. These consist essen-
tially of a sheathing base that nearly or completely
encircles the stem. The 1st scale is solitary and
lacks an axillary bud, the next 2 scales are paired
and subopposite, each with an axillary bud. The
bud of the 1st scale of the subopposite pair in
some species gives rise to a regenerative rhizome.
This supersedes the rhizome apex and makes
branching sympodial. The bud of the 2nd scale of
the subopposite pair gives rise to the erect stem.
Some Potamogeton species produce elongate
overwintering rhizomes. These develop in the axil
of the 1st of the paired scales. In Groenlandia and
other species of Potamogeton which do not pro-
duce overwintering rhizomes, the bud of the 1st
scale of the subopposite pair remains dormant.
The erect stems vary in length to 6 m, depending
partly on water depth. Each node produces one or
rarely more axillary buds. These generally remain
dormant, but sometimes grow into branches. The
apex of Potamogeton shoots produces an inflores-
cence subtended by 2 subopposite leaves. Each
leaf has an axillary bud, one or both of which may
break dormancy, in which case a sympodium is
formed.
The leaves are predominantly alternate, oppo-
site or in whorls of 3 in Groenlandia, and consist
of a sessile or petiolate blade and, usually, a basal
stipular sheath. The leaves in Groenlandia are
sessile and usually exstipulate, or stipulate on
young shoots (Fig. 96F). The leaf axils have 2-
several intravaginal squamules. The squamules
lack veins and are mostly linear to subulate, rarely
ovate, entire, and acute.
The blades vary in shape from linear to lan-
ceolate or oval. They are glabrous and predo-
minantly entire, although serrations are rarely
present. Serrations are evenly spaced along the
margins (P. crispus). Some species of Potamogeton
such as P. puleher have minute denticles along the
margin that become dislodged shortly after leaf
maturation. Venation is basically parallel, with
an obvious midvein and from 0-30, rarely more,
lateral veins. Cross-veins are normally present
between the longitudinal veins. The leaves of
many species are entirely submersed. Floating
leaves are present in several species of Potamoge-
ton. They are usually coriaceous, whereas the sub-
mersed leaves are delicate. The two types are often
widely different in shape. . lacunae. The nodes are probably the only break in
The ligules (often called stipular sheaths) are
normally free from the leaf base but in some
species adnate to it from the leaf base up to 3/4 of
its length. The ligules, when separate from the
blade, encircle the stem. The margins of the ligules
may be fused, forming a tube, but are more com-
monly free and overlap basally. The sheath is
mostly delicate and hyaline.
Vegetative reproductive structures and peren-
nating organs are produced at the tips of erect
stems and branches by several species of
Potamogeton. These are variously called winter
buds, turions or hibernacula, and primarily pro-
duced in species without elongate rhizomes.
Turions are short, hard stem apices with short
internodes. The effective propagation of turions
was demonstrated by Yeo (1966) who planted 1
turion of P. crispus on 1 April1963 and, at the end
of the growing season, found 23 520 turions
produced.
The inflorescences are pedunculate, solitary,
and terminal or axillary. The penduncles vary in
length from few to 50 mm or more. They are cylin-
dric and filiform to clavate and inflated and may
be erect to recurved.
VEGETATIVE ANATOMY (from Tomlinson
1982). The sieve tube plastids have cuneate pro-
tein bodies but no starch or filamental protein.
Silica bodies and calcium oxalate crystals are
lacking. Vessels with scalariform perforations are
often present in the roots of Potamogeton but
absent in Groenlandia; they are absent in stems
and leaves. The root epidermis is differentiated
into long and short cells, from the latter of which
root hairs are produced in Groenlandia, but this
exodermis is absent in Potamogeton.
The erect stem lacks stomata and has a thin to
slightly thickened cuticle. The epidermal cells
contain chlorophyll and surround a fairly wide
cortex that contains chlorenchyma and is copi-
ously lacunose. The outer 1 to 3 cell layers of the
cortex often comprise a hypodermis (pseudohy-
podermis in some literature), which is absent in
Groenlandia and some species of Potamogeton.
Immediately inside the cortex is an endodermis.
The endodermal cells in Groenlandia are
thin-walled but are variously thickened in
Potamogeton. Vascular and supporting tissues are
restricted to the stele in most species but in some
species of Potamogeton are present both in stele
and interlacunar and/or subepidermal bundles.
The cortex in the nodal area is compact and lacks
the longitudinal gas canals.
The leaf anatomy varies considerably between
genera and within Potamogeton.
 Potamogetonaceae
Groenlandia leaves have 3 veins at the base and
apex but 5 to 7 in the widest parts. The epidermal
layer is continuous except at the apex where a
well-developed apical pore is present, exposing
the tracheids. The cuticle is thin or absent.
The mesophyll consists of only 1 layer, except
along the veins. Lacunae are restricted to the
midvein area. All mesophyll and epidermal cells
contain chlorophyll. A fibrous strand extends
along the margin from the base nearly to the
apex.
Floating and submersed leaves of Potamogeton
differ in anatomy, although a pore is always
present. In floating leaves, the cuticle is thick on
the upper surface and thin on the lower, stomata
are common on the upper surface and rarely
present on the lower. Epidermis on both sides is
uniseriate, and the mesophyll consists of normal
palisade and spongy parenchyma. Large lacunae
traverse the spongy mesophyll on each side of the
midvein. In submersed leaves, stomata are rare
and nonfunctional, the cuticle is thin or nearly
absent, and the mesophyll varies from 1 to 4
or 5 cell layers, but a palisade is not developed.
Lacunae are common along the midvein and
sometimes occur in the mesophyll. All cells of
Potamogeton leaves, except those associated with
the veins, contain chlorophyll.
INFLORESCENCE STRUCTURE. The plants are
monoclinous. The inflorescences are usually
pedunculate with capitate to elongate spikes, or
reduced to 2 flowers.
FLORAL STRUCTURE. The flowers (Figs. 96D,H,
97D,E) are sessile and without subtending bracts.
There is a perianth of2-4 tepals that are adnate to
the filaments or connectives of the opposing
stamens. The perianth segments initially com-
pletely enclose the other floral parts. The struc-
tures here regarded as tepals in Potamogetonaceae
are frequently interpreted as staminal processes,
an interpretation supported by the stamen struc-
ture in the distantly related Posidoniaceae. Sattler
(1965) and Posluszny and Sattler (1973) demon-
strated that for the Potamogetonaceae, however,
the structures are best considered to be tepals.
There are mostly 4 stamens, rarely 2-3. Each theca
dehisces by a longitudinal slit.
The carpels are separate, mostly 4, rarely 1-3,
but there may be up to 7. The carpels are shortly
stipitate in P. robbinsii, these never exceeding
1-2mm. After fertilization, the peduncle often
bends laterally or becomes recurved. The pen-
411
duncle curvature serves to bring the developing
fruits under water.
EMBRYOLOGY. Anther-wall formation type is ap-
parently unknown. The epidermis persists during
anther wall development, and the endothecium
has Girdle-type thickenings. The tapetum cells
are predominantly 1-nucleate, rarely, e.g., in
Potamogeton natans, 2-nucleate. The walls even-
tually disintegrate and a periplasmodium is
formed. Microsporogenesis is of the successive
type. The tetrads are usually isobilateral. The
mature grains are then 3-celled.
The ovules are bitegmic and atropous in Pota-
mogeton (Davis 1969). A parietal cell is formed
and gives rise up to 7 parietal layers below the
nucellar epidermis. The micropyle is formed by
the inner integument.
Embryo sac formation is of the Polygonum type.
Endosperm formation is Helobial. Embryo forma-
tion follows the Caryophyllad type in which the
basal cell does not divide but enlarges. It com-
prises the terminal cell of the suspensor.
The ripe seeds lack endosperm and perisperm.
The embryo is coiled, macropodous, and linear. It
fills the entire seed cavity and lacks chlorophyll.
PoLLEN MORPHOLOGY. Pollen grains are in
monads, inapertuarate, ellipsoid to spheroidal,
and semitectate. The exine is less than 1.5 ~-tm
thick, homobrochate, consisting of muri with a
tectal solid part and an infratectal columellar part.
Lumina number varies from 10-31 per 50~-tm2
(Sorsa 1988).
KARYOLOGY. Chromosome reports are available
for about half of the species of the family. The
basic chromosome number for Potamogeton was
proposed as x = 7 by Les (1983). Potamogeton
exhibits extensive polyploidy, ranging from 2n =
14 to 2n = 52, and even 2n = 104, and several
aneuploid series. Groenlandia is reported to be
2n = 30.
POLLINATION. Most species are considered to be
wind-pollinated. Dragon flies have been observed
to land on P. nodosus spikes and to carry the
pollen, but there is no evidence of actual pol-
len transfer. Daumann (1963) detected minor
amounts of pollenkitt on the exines of P. fluitans
and P. natans, which led to slight stickiness of the
pollen. Potamogeton lucens, however, was found
never to have any pollenkitt. Daumann suggested
that the presence of pollenkitt may indicate that
412 Potamogetonaceae
anemophily in this situation has originated from
zoophily. Self-incompatibility is likely in at least
some species thought to be wind-pollinated (Voss
1972; Philbrick 1983). Some species regularly
flower under water. Philbrick (1983) observed
three species, one with aerial flowers only (P.
epihydrus), one with both emersed and submersed
flowers but similar inflorescences (P. pusillus var.
tenuissimus), and one with both emersed and sub-
mersed flowers but dimorphic inflorescences (P.
spirillus). The aerial flowers of P. epihydrus and P.
spirillus were facultatively autogamous, although
the stigmas expand before an thesis. Consequently,
outcrossing is possible and does occur; however,
any flowers not previously pollinated at anthesis
usually are selfed, resulting in near 100% fruit
set. The stamens of the submersed flowers of P.
spirillus dehisce prior to flower opening, resulting
in self-fertilization, again near 100% fruit set.
Potamogeton pusillus var. tenuissimus, however,
never had 100% fruit set. All of its flowers, both
submersed and emersed, appeared to be self-
incompatible. Aerial flowers were apparently
wind-pollinated, but the submersed flowers were
pollinated by air bubbles. The bubbles are released
from the opening anther, and some pollen adheres
to the surface of the bubble. The bubble moves to
the stigma of that flower or another flower, the
former of which is not fertilized, the latter,
however, resulting in outcrossing. Guo and Cook
(1989, 1990) determined that Groenlandia, in ad-
dition to being wind-pollinated, and P. pectinatus
are pollinated on the water surface, and below the
water surface, as described by Philbrick ( 1983)
above for P. pusillus var. tenuissimus.
FRUIT AND SEED. The fruit is a drupe or a berry.
The fruits of Potamogeton have a membranous
exocarp, fleshy mesocarp, and stony endocarp,
and are best considered a drupe, although they do
not fit the classical definition of any fruit type. The
fruits of Groenlandia lack a stony endocarp and
are thus 1-seeded berries.
DISPERSAL AND REPRODUCTIVE SYSTEMS. In
many species, the fruits after being released float
on the water for a period of time varying from less
than a week to several months. Spence et al. (1971)
reported fruits floating for 18 months under labo-
ratory conditions. The floating fruits are green
and photosynthetic, and the oxygen released by
photosynthesis keeps them buoyant. Eventually
the fruits sink and the pericarp decays. Muenscher
(1936) demonstrated that germination rate of
Potamogeton seeds is greatly reduced by drying,
and that many species, especially from colder re-
gions reguire freezing temperatures for months.
Lohammar (1954) demonstrated that seeds di-
gested by waterfowl retain the endocarp and
germinate at a high percentage.
Dispersal by feeding waterfowl is important
and probably accounts for, at least some, long-
distance dispersal. The vegetative reproductive
structures and buoyant seeds are obviously
capable of floating downstream in running waters.
Phibrick and Anderson (1987) studied breeding
systems of several species of Potamogeton and
suggested that those with submersed inflores-
cences are predominately autogamous and those
with aerial inflorescences are at least facultatively
autogamous, although some, as mentioned
earlier, are self-incompatible.
PHYTOCHEMISTRY. Saponins and alkaloids are
absent in the family. Tannins are known from the
family. Harborne (1982) reports that flavone agly-
cones, flavone glycosides, and C-glycosylated fla-
vones are common in Potamogetonaceae. Les and
Haynes (1995) summarized the flavonoid compo-
nents for the family as of that time. Anthocyanins
are absent, the red pigment characteristic of many
taxa consists of rhodoxanthin. Groenlandia ap-
parently has not yet been surveyed.
AFFINITIES. Taxonomists have historically en-
compassed the genus Potamogeton in a variety
of family concepts that broadly included diverse
genera such as Cymodocea, Halodule, Najas,
Phyllospadix, Potamogeton, Ruppia, Zannichellia,
and Zostera. Later authors (Tomlinson 1982;
Dahlgren et al. 1985; Cook 1990) adopted a
more conservative circumscription of Potamoge-
tonaceae with only three genera: Groenlandia,
Potamogeton, and Ruppia. The genera Althenia,
Lepilaena, Pseudalthenia, and Zannichellia are
usually segregated as a separate family (Zan-
nichelliaceae), although there is merit to consi-
dering their merger with the Potamogetonaceae
(see Zannichelliaceae and below).
Molecular data have been particularly helpful at
resolving questions of relationships among these
aquatic genera. A phylogenetic analysis of rbcL
data resolves a strongly supported, monophyletic
clade consisting of four genera: Groenlandia,
Lepilaena, Zannichellia, and Potamogeton (Les
and Haynes 1995; D.H. Les, unpubl.) Potamogeton
subgen. Potamogeton is quite distinct in these
analyses, but relationships among subgenus
Coleogeton and the remaining genera are unre-
solved due to an inadequate level of molecular
 Potamogetonaceae
divergence. As we mention in the treatment of
Zannichelliaceae, it is arguable whether the
Potamogetonaceae and Zannichelliaceae should
be retained taxonomically as separate families.
The molecular data also clearly separate both
Groenlandia and Potamogeton subgen. Coleo-
geton from P. subgen. Potamogeton. Together
with a number of morphological differences, this
result would help to rationalize the recognition of
three genera in the pondweed family. Phylogenetic
analysis of rbcL data also indicates that linear-
leaved species of Potamogeton subgen. Coleogeton
did not evolve from heterophyllous ancestors as
hypothesized by Les and Sheridan (1990), but
represent a distinct lineage that independently
acquired linear submersed leaves.
Although their relationship to Potamogetona-
ceae is occasionally hypothesized, none of the
genera of Cymodoceaceae, Najadaceae, Posidonia-
ceae, and Ruppiaceae associate as sister groups to
the Potamogetonaceae/Zannichelliaceae clade
(Les and Haynes 1995; D.H. Les, unpubl.). The
marine family Zosteraceae comprises the closest
sister group to this clade, but it is sufficiently di-
vergent at both the morphological and molecular
level to warrant the maintenance as a separate
family, and resolves as a separate, monophyletic
group (Les and Haynes 1995). The Juncaginaceae
are the closest sister group to the clade that in-
cludes the Potamogetonaceae, Zannichelliaceae,
and Zosteraceae (Les and Haynes 1995). All of
these groups share similar adaptions to aquatic
life, such as a lack of stem vessels and pollen aper-
tures. They also have in common root hair short-
cells, intravaginal squamules, caryophyllad type of
embryo, exendospermous seeds, and uniovulate
ovaries.
DISTRIBUTION AND HABITATS. The Potamo-
getonaceae are subcosmopolitan, with most spe-
cies occurring in temperate regions, especially in
the northern hemisphere.
Nearly all Potamogetonaceae grow in aquatic
freshwater ecosystems; several species can survive
short periods of drought in a terrestrial state. A
few species occur in brackish waters, but none are
marine. The family ranges in elevation from sea
level to 4500m. Hellquist (1980) studied the distri-
bution of Potamogeton (s.l.) in relation to alkalin-
ity and divided the species into six groups, based
on the mean of the alkalinity of the waters in
which they occurred. Some were obligate to low
alkalinity, some to intermediate alkalinity, and
some to high alkalinity. Finally, a few could occur
in nearly all of the six groups.
413
PALEOBOTANY. The earliest records of the family
are from the Paleocene. Records become more
common in Eocene and younger deposits
(Daghlian 1981 ). Some earlier reports of the family
are based on vegetative remains, but these cannot
be satisfactory assigned to the family (Daghlian
1981). The first fruits of Potamogetonaceae are
assigned to a separate form genus, and are from
the Paleocene, while Potamogeton is first recorded
in the Eocene.
EcoNOMIC IMPORTANCE. As source of food for
waterfowl, Martin and Uhler (1939) rank
Potamogeton first in importance in the United
States and Canada, and Gaevskaya (1966) lists at
least 135 species of animals that are known to feed
on the family. Some are regarded as water weeds
warranting control. Some species of Potamogeton
are also used in the aquarium trade.
KEY TO THE GENERA
1. Leaves all opposite; fruit without stony endocarp
2. Groenlandia
- Leaves alternate except those that subtend an inflorescence;
fruit with stony endocarp 1. Potamogeton
1. Potamogeton L. Fig. 96A-E, 97
Potamogeton L., Sp. Pl. 1: 126 (1753); Hagstrom, J.O., Critical
researches on the potamogetons. K. Sven. Vetenskapsakad.
Hand!. 55(5): 1-281 (1916); Dandy, J.E., The genus Pota-
mogeton L. in tropical Afric. J. Linn. Soc. Bot. 50: 507-540
(1937); Fernald, M.L., The linear-leaved North American
species of Potamogeton, section Axillares. Mem. Gray Her.
3: 1-183 (1932); Ogden, E.C., The broad-leaved species of
Potamogeton of North America north of Mexico. Rhodora
45: 57-105, 119-163, 171-214 (1946); Haynes, R.R., A revi-
sion of the clasping-leaved Potamogeton (Potamogetona-
ceae) Sida 11: 173-185 (1985).
Stuckenia Borner (1912).
Coleogeton Les & R.R. Haynes (1996).
Propagated by seeds, tubers, turions, or rhizomes;
stems terete or compressed, nodes occasionally
with oil glands; leaves alternate to subopposite;
submersed leaves sessile or petiolate, linear to or-
bicular, subulate to obtuse at apex, acute to perfo-
liate at base, margins entire or serrate, rarely
crimped, veins 1-35; floating leaves coriaceous,
mostly petiolate, rarely subsessile, elliptic to
ovate, acute to obtuse at apex, cuneate to rounded
or cordate at base, margins entire, veins 1-51;
stipules tubular, sheathing stem and young inflo-
rescences, connate or convolute, either free or
adnate to base of submersed leaves, free from base
of floating leaves. Inflorescence a capitate or cylin-
drical spike or panicle of spikes with 1-20 whorls
414 Potamogetonaceae

of flowers, compact or interrupted, with 2-4 Cronquist, A. 1981. See general references.
flowers in each whorl. Perianth segments 4, free, Daghlian, C.P. 1981. A review of the fossil record of monocoty-
ledons. Bot. Rev. (Lancaster) 47: 517-555.
rounded, short-clawed; stamens 4; carpels 4 or 1. Dahlgren, R.T.M., Clifford M.T. 1982. See general references.
Fruit abaxially rounded or keeled, beaked. Two Dahlgren, R., Rasmussen, F.N. 1983. Monocotyledon evolu-
subgenera: subgen. Potamogeton, stipular sheaths tion, characters and phylogenetic estimation. Evol. Bioi. 16:
of submersed leaves free from leaf base, or if ad- 255-395.
nate, then the adnate portion less than 1/2 length Dahlgren, R.M.T. eta!. 1985. See general references.
Daumann, E. 1963. Zur Frage nach dem Ursprung der
of stipule; leaves both submersed and floating Hydrogamie. Zugleich ein Beitrag zur Bliitenokologie von
or all submersed, the submersed pellucid, non- Potamogeton. Preslia 35: 23-30.
canaliculate; peduncles stiff and, if long enough, Davis, G.L. 1969. See general references.
projecting above surface of water; ca. 80 species, Gaevskaya, N.S. 1966. The role of higher aquatic plants in the
nutrition of the animals of fresh-water basins. (Trans!. from
near cosmopolitan; subgen. Coleogeton (Reichb.) Russian by D.G. Maitland Muller 1969). Yorkshire.
Raunk. (1896), stipular sheaths of submersed Guo, Y.-H., Cook, C.D.K. 1989. Pollination efficiency of
leaves adnate to leaf blade for 2/3 or more of Potamogeton pectinatus L. Aquat. Bot. 34: 381-384.
stipule length; leaves all submersed, opaque, Guo, Y.-H., Cook, C.D.K. 1990. The floral biology of Groen-
canaliculate, turgid; peduncle fexible, not project- landia densa (L.) Fourreau (Potamogetonaceae). Aquat.
Bot. 38: 283-288.
ing inflorescence above surface of water; turions Hagstr0m, J.O. 1916. Critical researches on the potamogetons.
lacking. About ten spp., near cosmopolitan. Kong!. Sven. Vetenskapsakad. Hand!. 55(5): 1-281.
We are including Coleogeton within Potamo- Harborne, J.B. 1982. Flavonoid compounds. In: Dahlgren,
geton at the insistence of the editor. In our opin- R.T.M., Clifford, H.T. (eds.) The monocotyledons. A com-
ion, data from a variety of sources, including parative study. London: Academic Press.
Haynes, R.R. 1974. A revision of North American Potamo-
molecular, cytological, hybridization, anatomical, geton subsection Pusilli (Potamogetonaceae). Rhodora 76:
morphological, and pollination studies, indicate 564-649.
strongly that subgen. Coleogeton should occupy Haynes, R.R. 1978. The Potamogetonaceae in the Southeastern
generic rank. United States. J. Arnold Arbor. 59: 170-191.
Haynes, R.R., Holm-Nielsen L.B. "1985" (1987). A generic
treatment of Alismatidae in the neotropics. Acta Amazonia
2. Groenlandia J. Gay Fig. 96F-I 15: 153-193.
Hellquist, C. B. 1980. Correlation of alkalinity and the distribu-
Groenlandia J. Gay, C. R. Acad. Sci. Paris 38: 703 (1854). tion of Potamogeton in New England. Rhodora 82: 331-
344.
Irmisch, T. 1858. Ueber einige Arten aus der natiirlichen
Propagated by stolons, buds or seeds. Leaves Pflanzenfamilie der Potameen. Berlin: G. Bosselmann.
submersed, opposite, pellucid, sessile, linear- Les, D.H. 1983. Taxonomic implications of aneuploidy and
lanceolate to broadly lanceolate, acute at apex, polyploidy in Potamogeton (Potamogetonaceae). Rhodora
longitudinally folded, clasping at base, the 85: 301-323.
margins entire, often crimped, the veins 5-7. Les, D.H., Haynes, R.R. 1995. Systematics of subclass
Alismatidae: a synthesis of approaches. In: Rudall, P.J.,
Stipules only present on young shoot. Turions Cribb, P.J., Cutler, D.F., Humphries, C.J. (eds.) Monocotyle-
absent. Inflorescence axillary or terminal, of 2-6 dons: systematics and evolution. Royal Botanic Gardens,
opposite flowers, submersed or held slightly Kew, pp. 353-377.
above surface of water. Perianth of (2-) 4, free, Les, D.H., Philbrick, C.T. 1993. Studies of hybridization and
rounded, short -clawed segments; stamens (2-) 4; chromosome number variation in aquatic angiosperms:
evolutionary implications. Aquat. Bot. 44: 181-228.
carpels (2-3)-4. Fruit dorsally keeled, beaked. Les, D.H., Sheridan, D.J. 1990. Biochemical heterophylly and
Only one sp., G. densus (L.) Fourr., Spain to Nor- flavonoid evolution in North American Potamogeton
way, east to Siberia and Afghanistan, and south to (Potamogetonaceae). Am. J. Bot. 77: 453-465.
Algeria. Les, D.H., Garvin, D.K., Wimpee, C.F. 1993. Phylogenetic
studies in the monocot subclass Alismatidae: evidence for a
reappraisal of the aquatic order Najadales. Mol. Phylogenet.
Evol. 2: 304-314.
Selected Bibliography Lohammar, G. 1954. Matsmaltningens inverkan pa
Potamogeton-fronas groning. Fauna Flora 49: 17-32.
Aalto, M. 1970. Potamogetonaceae fruits. I. Recent and sub- Martin, A.C., Uhler, F.M. 1939. Food of game ducks in the
fossil endocarps of the Fennoscandian species. Acta Bot. United States and Canada. U.S.D.A. Tech. Bull. No. 634.
Fenn. 88: 1-85. Muenscher, W.C. 1936. The germination of seeds of
Ascherson, P., Graebner, P. 1907. Potamogetonaceae. In: Potamogeton. Ann. Bot. (Lond.) 50: 805-821.
Engler, A. (ed.) Das Pflanzenreich IV. 11: 1-184. Leipzig: Philbrick, C.T. 1983. Aspects of floral biology in three species
W. Engelmann. of Potamogeton (pondweeds). Mich. Bot. 23: 35-38.
Cook, C.D.K. 1990. Aquatic plant book. The Hague: SPB Philbrick, C.T., Anderson, G.J. 1987. Implications of pollen/
Academic Publishing. ovule rations and pollen size for the reproductive biology of
 Potamogetonaceae/Rapateaceae
Potamogeton and autogamy in aquatic angiosperms. Syst.
Bot. 12: 98-105.
Posluszny, U., Sattler, R. 1973. Floral development of Potamo-
geton densus. Can. J. Bot. 51: 647-656.
Posluszny, U., Sattler, R. 1974. Floral development of Potamo-
geton richardsonii. Am. J. Bot. 61: 209-216.
Preston, C.D. 1995. Pondweeds of Great Britain and Ireland.
BSBI Handbook No. 8. London: Botanical Society of the
British Isles.
Sattler, R. 1965. Perianth development of Potamogeton rich-
ardsonii. Am. J. Bot. 52: 35-41.
Singh, v. 1964. Morphological and anatomical studies in
Helobiae. I. Vegetative anatomy of some members of
Potamogetonaceae. Indian Acad. Sci. 60: 214-231.
Singh, V. 1965. Morphological and anatomical studies in
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Voss, E.G. 1972. Michigan flora. Pt. 1. Gymnosperms and
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I. Weeds 14: 110-113.
415
Rapateaceae
D.W. STEVENSON, M. CoLELLA, and B. BooM
Rapateaceae Dumort., Anal. Pam. Pl.: 60 (1829).
Terrestrial, epiphytic, or amphibic perennial
herbs with an erect or prostrate rhizome. Leaves
unifacial to bifacial, condensed on a short stem,
with an open asymmetrical conduplicate sheath;
distichously or spirodistichously arranged; rarely
petiolate, the petiole and the leaf blade occasion-
ally bearing short spines; ligules rarely present.
Inflorescences solitary or crowded, axillary or ter-
minal, on leafless scapes, often subtended by 2
or more spathaceous bracts, globose or flat; the
spikelets composing the inflorescences bearing
numerous coriaceous bracts and a terminal
flower; inflorescences that have an indeterminate
main axis and lateral cymose units are more com-
mon than those with a determinate main axis and
cymose lateral units (Monotremeae); flowers tri-
merous, hypogynous, perfect, and actinomorphic
or slightly zygomorphic; sepals 3; petals 3, occa-
sionally spotted; stamens 3 + 3, tetrasporangiate,
connate at the base or adnate to the corolla tube,
basifixed; anthers sometimes bearing terminal
appendages, dehiscing by 4, 2, or 1 apical or
subapically lateral pores, extrose or latrose; ovary
syncarpous, superior, 3-locular or 1-locular by
abortion; style 1, erect; stigma capitate, often mi-
nutely ciliate; ovules 1-8 per locule, anatropous,
bitegmic, and crassinucellate; placentation axile
or basal; fruit a septicidal capsule; seeds globose,
prismatic, or oblong, occasionally bearing ap-
pendages; endosperm containing abundant
starch; embryo lenticular, at micropylar end with
flat surface appressed to endosperm; mechanical
tissue of seed coat derived from both integuments.
A basically neotropical family of 16 genera and
about 80 species, with the greatest diversity in the
Guayana region and with one disjunct genus in W
Africa.
VEGETATIVE MORPHOLOGY. Plant size varies
from 30 em to 2( -4) m. Roots are produced at the
nodes. The axis consists of 2 portions, 1 that is a
subterranean rhizome bearing scale leaves and
occasionally branching sympodially, and another
that is an aerial and mostly monopodia! leaf-
416 Rapateaceae
bearing stem. Supernumerary axillary buds are
present in the nodal region in the Rapateeae and
Monotremeae. The aerial stem can bear 1 of 3
types ofleaves: a unifacialleaf, a bifacialleaf, or an
intermediate leaf with a rudimentary unifacial
blade; the leaf base is conduplicate, symmetrical
in distichous plants, and asymmetrical in
spirodistichous plants, sheath open.
VEGETATIVE ANATOMY. In general, a rhizoder-
mis is absent at maturity, or if persistent, then it
becomes suberized and accompanied by hypoder-
mal sclereids (velamen). In roots, the outer cortex
contains a well-developed exodermis, the middle
cortex contains parenchyma cells with arms, the
inner cortex is composed of spheroidal cells
arranged in radial files, and the endodermis is
uniseriate and the cells have secondary walls at
maturity. The root central cylinder (stele) has a
uniseriate to multiseriate pericycle, is polyarch,
has sclerified ground tissue, and occasionally met-
axylem vessels.
Stems have a broad outer cortex composed of
starch-containing thin-walled parenchyma cells
and an inner cortex composed of 2-5 layers of
sclerids. The central cylinder of the stem consists
of some sclerified ground tissue and both collat-
eral and amphivasal vascular bundles that have
some metaxylem vessels. Slime is produced from
slime canals and cavities in some genera.
Leaf epidermal cells are similar on both sur-
faces. Epicuticular waxes are either lacking or in
some cases present in the form of amorph-
ous globules (Frolich and Barthlott 1988). The
paracytic stomata are located only on the abaxial
surface and the guard cells are thick-walled with
dumbbell-shaped lumina and the accessory cells
contain oil (Carlquist 1969). Bifacial leaf blades
develop most of the blade from 2 marginal mer-
istems and have a rudimentary radial leaf tip,
whereas unifacialleafblades develop from a single
adaxial meristem representing the leaf tip. Both
unifacial and bifacial lamina may occur in the
same genus (Schoenocephalium, Cephalostemon,
Stegolepis) as in some other monocot genera (e.g.,
Sansevieria). When a unifacial leaf blade is
present, the mesophyll is more or less undifferen-
tiated into palisade and spongy tissues (Arber
1922). When a dorsiventrallamina is present, the
mesophyll is differentiated into palisade and
spongy tissues. The palisade tissue is composed
of chlorenchyma cells with anticlinally infolded
walls and the spongy tissue is composed of arm-
parenchyma and mostly collapsed nonphoto-
synthetic spheroidal cells. Collateral vascular
bundles have a sheath of thin-walled parenchyma
cells and fibers either surrounding the vascular
tissue or only next to the phloem in smaller
bundles. In the largest vascular bundles, the fibers
form bundle extensions to the hypodermis.
Vessels are not present in leaves.
Uniseriate 2-6-celled trichomes are located
only in the axils of leaves and bracts. Al-
though nonglandular in appearance, the tri-
chomes secrete copious amounts of slime that
completely covers the emerging leaves and young
inflorescences.
Cellular inclusions in form of silica and tannins
are present. Epidermal cells of the leaves contain
either solitary or aggregated spherical silica bod-
ies. In general, the epidermal silica cells are con-
fined to costal regions. Tannin-containing cells
occur in all vegetative parts of the plants of some
genera and are particularly abundant in the inner
cortex of roots and inflorescence bracts. The
tanniniferous cells may occur singly or may be
aggregated into longitudinal files.
Carlquist (1969) distinguished anatomical
variations according to the subfamilial and tribal
scheme proposed by Maguire (1958). The follow-
ing is a summary of the unique features in these
groups. Saxofridericioideae have, in the roots,
asymmetrical, lignified, arm-parenchyma cells;
long, narrow tanniferous cells arranged in vertical
series in the central cortex; very wide metaphloem
sieve tubes, and a sclerified pith. Stems have
tannin-filled parenchyma cells aggregated in both
the cortex and the stele. In Schoenocephalieae,
cells with tanniferous contents are abundant in
the inner cortex of roots, and in stems slime canals
and cavities are present in the cortex and the stelar
parenchyma is unlignified. In Saxofridericieae, the
slime canals and cavities are absent and the stelar
parenchyma is slightly lignified. The roots of
Rapateoideae have cells of the central cortex ar-
ranged in radial plates, narrow metaphloem sieve-
tube elements, and simple and scalariform vessels.
In stems, tannin-filled parenchyma cells are dis-
tributed idioblastically in both cortex and stele or
sometimes absent (Spathanthus) and simple ves-
sels are present. The tribe Monotremeae lacks
silica cells in the epidermis, has silica sand in cor-
tical parenchyma cells, lacks hypodermal fibrous
strands, and has nonlignified stelar parenchyma.
In contrast, the tribe Rapateeae has silica cells in
the epidermis, and lacks silica in the cortical pa-
renchyma, has hypodermal fibrous strands, and
strongly lignified stelar parenchyma. Spathanthus
is anomalous in that it has some stem anatomical
features of the Monotremeae such as the presence
 Rapateaceae 417

of silica sand in cortical parenchyma cells, lacks

hypodermal fibrous strands, and has nonlignified
stelar parenchyma cells but otherwise conforms to
the Rapateeae in other stem features and in root
anatomy and floral morphology.

INFLORESCENCES. The inflorescence terminates a

long nonappendaged peduncle and is generally
enclosed by 2 or more bracts. Two different lines
of inflorescence construction are present: the in-
determinate thyrse (thyrse s. str., Weberling 1989)
with an indeterminate main axis and cymose par-
tial inflorescences (Figs. 98, 99), and the determi-
nate thyrse (thyrsoid, Weberling 1989) with the
main axis and partial inflorescences determinate.
The partial inflorescences are referred to as spike-
lets by convention, but they are most likely based
upon a bostryx. This is difficult to ascertain in the
Rapateaceae because each spikelet contains only a
seemingly terminal flower. The thyrse has spike-
lets whose length ranges from 0.5 to 5 em and
which consist of a variable number of coriaceous,
spirally arranged bracts. The number of spikelets
is variable, and some genera have thyrses with
70 spikelets (Saxofridericia, Spathanthus) or
T
¢
only 1-3 spikelets per inflorescence (Stegolepis,
Monotrema). The number of spathaceous bracts
subtending the inflorescence and their degree I
of fusion is variable. Shape, size, and develop-
mental patterns of the thyrse vary among the sub- ,./ ·: ..
E

~1 1 (~
families. The thyrse is common in the tribes
Saxofridericieae, Schoenocephalieae, and
Rapateeae; the determinate thyrse is present in
only 2 members of the Monotremeae: Windsorina K
and Maschalocephalus. The position of inflores- 1111 A

cences varies among the tribes. Thus, in Saxo-

fridericieae, the inflorescences are axillary; in
Schoenocephalieae, they are terminal with a dor-
mant axillary bud at the base; and in Rapateeae,
they are axillary with a proximal axillary vegeta-
tive bud.
Fig. 98A-K. Rapateaceae. A-D Saxofridericia grandis. A
Habit. B Inflorescence. C Capsule. D Seed, dorsal and ventral
view. E S. petiolata, spikelet. F-G S. spongiosa. F Spikelet. G
Anther. H-L Rapatea membranacea. H Inflorescence. I Same,
sectioned lengthwise. J Spikelet. K Ventral and lateral view of
anther. (A-G Maguire 1958, H-K Maguire 1965)

FLORAL STRUCTURE. The flowers have 3 coria-

ceous, lanceolate, imbricate sepals that are mem-
branous at the base and with hyaline margins. The
sepals are free from each other or basally : :': : con-
nate. The petals vary in size, shape, and color
within the family. They are usually imbricate, con-
nate at the base, lanceolate, ovate, obcordate, or
obovate. Petal color includes white, pink, yellow,
orange, red, and greenish. The stamens are 3 + 3
and are usually connate basally. The stamens may
be adnate to the flower tube.
Anther opening is poricidal (Figs. 98G,K, 99E).
Rarely, each of the 4 pollen sacs opens with a dis-
tinct pore (Schoenocephalium); mostly the pollen
sacs merge more or less toward the anther apex,
and opening is by 1 pore for each theca or by a
single pore for the whole anther. The pores may
occupy a subterminal ventral position or may be
strictly terminal, the former position possibly
being indicative of an originally longitudinal
anther dehiscence.
EMBRYOLOGY. Anther-wall ontogeny follows the
monocotyledonous type (Tiemann 1985). The mi-
crospore mother cells undergo simultaneous divi-
418
w F
Fig. 99A-F. Rapateaceae. Rapatea celiae. A Habit. B Lateral
view of inflorescence. C Spikelet. D Same, with open flower. E
Anther, ventral view. F Cross-section of anther. (Maguire
1965)
sion and the tetrads are decussate or tetrahedral.
Endothecial thickenings are of the spiral type, the
tapetum is secretory and binucleate. Pollen grains
are shed in the 2-celled stage. A funicular obtura-
tor forming a rim around the funicular region is
sometimes present. The micropyle is formed by
the 2 integuments. Embryo-sac development fol-
lows the Polygonum type, endosperm formation
is nuclear, and embryogenesis is of the Asterad
type. Supernumerary antipodals are common in
Rapateoideae. Fertilization is porogamous. The
reserve substances in the endosperm are simple
starch grains, oil, and proteins. The little-
differentiated mature embryo is lenticular, at the
micropylar end, and with the broad surface
appressed to the endosperm.
PoLLEN MoRPHOLOGY. Pollen shape ranges
from oblong to elliptical with some grains often
flattened laterally. Pollen-wall structure is tectate-
Rapateaceae
columellate with a thick foot-layer and a granular
endexine (Zavada 1983). All taxa studied have ex-
ine flakes scattered over the aperture. Aperture
types and sculpturing are different between the
two subfamilies. In Saxofridericioideae, the pollen
is sulcate with a fusiform sulcus; however, both
trichotomosulcate and sulcate grains have been
reported from the same anther in Schoeno-
cephalium (Carlquist 1961). In the Saxo-
fridericieae, the exine pattern is reticulate,
whereas it is usually foveolate (scrobiculate in
Guacamaya) in the Schoenocephalieae. Within
Rapateoideae, the pollen is sulcate with the sulcus
narrowed in the center and reticulate in the
Monotremeae and zonisulcate (disulcate in
Spathanthus) and scrobiculate in the Rapateeae.
Except for Rapatea, unusually large pollen grains,
up to 110 X 85~-tm in Cephalostemon (Malme
1933b), have been reported for the Rapateeae
(Carlquist 1961; Tiemann 1985).
KARYOLOGY. Data on karyotypes and chro-
mosome numbers in Rapateaceae are sparse
and concentrated in the subfamily Rapateoi-
deae. Spathanthus bicolor and Cephalostemon
angustatus have 2n = 52, whereas the African
Maschalocephalus dinklagei is 2n = 22.
POLLINATION. Most Rapateaceae (except Mono-
tremeae) seem to lack nectaries and possess
melittophilous pollen flowers (Vogel 1981);
Kunhardtia is ornithophilous (Steyermark 1974);
its nectary is unknown. Saxofridericia is buzz-
pollinated by bees and the Schoenocephalieae
are hummingbird-pollinated (Givnish et al.
1994).
SEED STRUCTURE. The seed coat is formed by
both integuments in all known cases, but other-
wise seed shape, size, sculpturing, and ontogeny
differ among the 4 tribes. In general, the mature
endosperm shows a protein-rich outer layer and
exotegmic cells have a zigzag pattern, while the
exotesta is siliciferous. In the Saxofridericioideae,
the seeds are pyramidal, prismatic, or lens-shaped
and striate or alveolate. Silica is present as silica
bodies, there are only 1-2 exotegmic middle lay-
ers, and the exotegmen functions as a mechanical
layer. In some genera, the exotegmen is sculptured
by protruding endotestal cells. In Rapateoideae,
the seeds are oval or oblong and have longitudinal
striae or are punctuate or granulose. In Rapateeae,
the seeds are longitudinally striate and occasion-
ally the seed appendage is papillate-mitriform.
There are silica incrustations on endotestal walls,
 Rapateaceae
the hilar scar is large, there are more than 2 middle
layers in the exotesta, and tracheoidal plates have
been reported. The seeds of Cephalostemon bear a
conical spongy caruncle or elaiosome at the cha-
laza! end and in Cephalostemon angustatus, the
seed becomes detached from the fruit by an ab-
scission layer of elongated cells with amyloplasts.
The endotesta of Spathanthus unilateralis has U-
shaped pectinaceous, siliceous walls, and contains
globular lipids. The seeds of the Monotremeae are
white and granulate, and have an apical flat
appendage.
PHYTOCHEMISTRY. Stored tannins and copious
amounts of slime are produced, but their identity
has not been elucidated. A remarkable feature of
Rapateaceae is their ability to accumulate up to 13
600 ppm of aluminum.
SUBDIVISION AND RELATIONSHIPS WITHIN THE
FAMILY. Based on the number of ovules per
carpel, placentation type, and seed, pollen, and
inflorescence morphology, Maguire (1958) di-
vided the family into two subfamilies each with
2 tribes, Saxofridericioideae and Rapateoideae,
with Saxofridericieae and Schoenocephalieae, and
Rapateeae and Monotremeae, respectively. Recent
anatomical, palynological, and embryological data
tend to confirm the subfamilial classification;
however, discrepancies occur at the tribal level.
The greatest number of plesiomorphic states such
as numerous ovules, usually massive inflores-
cences, pollen and ovular morphology occur in
the Saxofridericioideae. Within this subfamily it is
clear that Stegolepis, Epidryos, and Phelpsiella are
very close, and the same is true of Marahuacaea
and Amphiphyllum. Stegolepis retains some
primitive characters but shows apomorphies in
the reduction of the size of spathaceous bracts
that culminate in their absence and in the decreas-
ing size of the inflorescence. The spathaceous
bracts enclose the inflorescence after anthesis
in Saxofridericia and in some members of the
Schoenocephalieae, and ovary nectaries together
with ornithophily are apomorphic. Pollen mor-
phology does not differ greatly among the
Saxofridericioideae. Cymose inflorescence con-
struction and nectaries are present only in the
Monotr.emeae which also have white seeds with
flat appendages and a powdery exudate. Spa-
thanthus is the most enigmatic of genera in terms
of placement. Anatomically, it appears closer to
Monotrema, but in terms of seed structure and
floral morphology it is more similar to members
of the Rapateeae.
419
Recent analyses of sequence date from ndhF, a
chloroplast gene, indicate that the Rapateoideae
may be paraphyletic, but that the Saxofrideri-
cioideae are monophyletic with a paraphyletic
tribe Saxofridericieae and a monophyletic tribe
Schoenocephalieae (Givnish et al. 1995). It is
claimed that many of the morphological charac-
ters used to circumscribe the infrafamilial groups
recognized by Maguire (1958) are homoplasious.
However, because morphological characters were
simply mapped upon a cladogram produced by
molecular data only, these conclusions have not,
in fact, been tested by simultaneous analysis.
AFFINITIES. The Rapateaceae do not show a
unique set of embryological features other than
the simultaneous type of microsporogenesis and
supernumerary antipodals. However, they do
share many anatomical as well as embryological
and palynological features with the Commelina-
ceae, Xyridaceae, and, to a lesser extent, with the
Eriocaulaceae and the Mayacaceae, but the combi-
nation of embryological characters distinguishes
the family from related families. The stomata in
Rapateaceae, Xyridaceae, and Eriocaulaceae show
accessory cells with oil; Mayacaceae and Com-
melinaceae differ in the stomata type and their
accessory cells lack oil. Silica crystals are lacking
in Mayacaceae, Xyridaceae, and Eriocaulaceae.
The ovules are bitegmic, anatropous (in Com-
melinaceae they may be orthotropous to hemi-
anatropous) and crassinucellate in Rapateaceae
and Commelinaceae, the micropyle is formed by
the inner and outer integuments in Rapateaceae,
Xyridaceae, and Eriocaulaceae. The embryo-sac
formation conforms to the Polygonum type in all
families except in some Xyris and in some
commelinaceous genera, where the Allium type
also occurs. There are equal numbers of layers in
the outer integument in Rapateaceae and Com-
melinaceae. Endosperm is Nuclear in all families
except the Xyridaceous genus Abolboda, which
has Helobial endosperm. Poricidal anthers are
present in Commelinaceae, Mayacaceae, and
Rapateaceae. The endothecium has spiral thicken-
ings in Rapateaceae, Eriocaulaceae, and Com-
melinaceae. Secretory and binucleate tapetum is
present in Xyris, Eriocaulaceae, and Rapateaceae.
Pollen grains with exine flakes in the sulcate aper-
ture are present in Xyris but not in other
Xyridaceae and Rapateaceae. The exceptionally
large pollen grains of the Rapateeae are paralleled
in the Xyridaceae. The fruit is a capsule, but
in Commelinaceae berries are common. Bees
have been reported as the common pollinators
420 Rapateaceae

in all families but Eriocaulaceae. Some authors bilities remaining as the potential nearest rela-
(Hutchinson 1973; Cronquist 1981; Dahlgren et al. tives: Bromeliaceae, Juncaceae-Cyperaceae, and
1985; Thorne 1992; Takhtajan 1997) have accepted Eriocaulaceae-Xyridaceae-Mayacaceae.
a relationship between Rapateaceae and Xyrida-
ceae. However, there is little evidence to sup- DISTRITUTION AND HABITATS. The Rapateaceae
port this other than general leaf morphology. are nearly exclusively neotropical. They have their
Stevenson and Loconte (1995) proposed that center of distribution in the lowlands and high-
Rapateaceae are related to a group containing lands of the Guayana Shield, but extend to
Eriocaulaceae, Xyridaceae, and Mayacaceae. Thus, Panama and Bolivia; one genus, Maschalo-
Rapateaceae fall between Commelinaceae on the cephalus, is restricted to West Africa. Rapateaceae
one hand and the aforementioned group on the are most often found on oligotrophic white sand
other. Tiemann {1985) presents comparisons be- substrates or bogs, but occur also in a variety
tween the embryological features of the Rapatea- of other habitats such as forest understory,
ceae, Xyridaceae, Commelinaceae, Eriocaulaceae, shrublands, seasonally flooded savannas, and
Mayacaceae, Juncaceae, and Cyperaceae, and con- open rock and bog formations (Fig. 100). Exclu-
cludes that the Rapataceae is more closely related sively highland taxa occur in bogs and on sand-
to the Juncaceae-Cyperaceae than to the Com- stone and granitic substrates, where they have a
melinalian families. This is, however, based upon narrow distribution or are endemics to particular
shared general features rather than shared derived table mountian (tepui) summits; e.g., in the
characters. In fact, the Rapateaceae share no Saxofridericieae, Phelpsiella, Amphiphyllum,
derived embryological features, except simultane- and Marahuacaea are each endemic to a single
ous microsporogenesis, with the Juncaceae or tepui. The majority of the Rapateoideae occupy
Cyperaceae (see Table VII in Tiemann 1985). Ana-
tomical features that might show a relationship
with Juncaceae and Cyperaceae are, in fact, also
found in the Xyridaceae and Eriocaulaceae and to
a lesser extent in the Commelinaceae. Root ana-
tomical features such as middle cortex differenti-
ated into long and short parenchyma cells are a
synapomorphy of Rapateaceae, Eriocaulaceae,
Xyridaceae, and Mayacaceae. The presence of
amphivasal stem bundles, axillary glandular and/
or secretory trichomes, and differentiated peri-
anth are other synapomorphies of these familes.
The latter two also include the Commelinaceae.
These features are not found in other Com-
melinidae and thus are not homoplasious.
Recent data analyses derived from sequences of
two chloroplast genes, rbcL (Duvall et al. 1993;
Chase et al. 1995; Givnish et al. 1995; Linder and
Kellogg 1995) and ndhF (Givnish et al. 1995), have
placed the Rapateaceae in a strikingly different
position than have analyses of morphological
data. Molecular data support the sister-group re-
lationship of the Rapateaceae with the Bromelia-
ceae, as previously suggested by Pilger {1930).
This result is also obtained whenever rbcL data is
combined with morphological data (Chase et al.
1995; Linder and Kellogg 1995). When Linder and
Kellogg analyzed only morphological data, the
results supported the Rapateaceae as the sister
group to the Juncaceae-Cyperaceae, as suggested
by Tiemann (1985). Thus conflicting results are Fig. 100. Rapateaceae. Stegolepis angustata. Venezuela,
obtained both between and among morpholo- swampy place in Gran Sabana; note fire-resistant, blackened
gical and molecular data sets with three possi- leaf sheaths. (Photo K. Kubitzki)
 Rapateaceae
lowlands where they have probably diversified
more recently. Characteristically, the African
Maschalocephalus grows in white sand habitats as
well.
KEY TO THE GENERA
1. Carpels pluriovulate, placentation axial; seeds prismatic or
pyramidal: Subfam. Saxofridericioideae
- Carpels uniovulate; placentation basal or subbasal, seeds
oblong or oval: Subfam. Rapateoideae
2. Petals obcordate or obovate, exserted: Tribe Sxofri-
dericieae
- Petals lanceolate, included within the sepals (except in
Kunhardtia): Tribe Schoenocephalieae
3. Inflorescences bracteate
- Inflorescences ebracteate, or the bracts represented by
vestigial membranes; scapes several, rarely 1
4. Inflorescences tri- or pluribracteate
- Inflorescences bibracteate, bivalvate
5. The 2 outer bracts valvate, connate, the inner enclosed
3. Amphiphyllum
- The outer 2 conduplicate, the inner, smaller and condupli-
cate; leaves strongly auriculate 6. Marahuacaea
6. Anthers 4-celled, with apical appendages, dehiscing by a
single subapical pore; spikelets piercing spathaceous
bracts 1. Saxofridericia
- Anthers 2-celled, exappendiculate, dehiscing by 2 circular
apical pores; heads small, compressed; spikelets not pierc-
ing the inflorescence bracts 2. Phelpsiella
7. Anthers transversally rugose; 4-celled; seed pyramidal;
endocarp not separating from exocarp 4. Stegolepis
- Anthers smooth, 1-celled; seed flat and disciform; en-
docarp separating from exocarp 5. Epidryos
8. Corollas red, zygomorphic, exserted; anthers 2-celled
7. Kunhardtia
- Corollas red or green, actinomorphic, included within the
sepals; anthers 4-celled at least toward the base
9. Anther at apex bilobed, dehiscing by 2 apical pores;
spathaceous bracts ruptured by individual spikelets
8. Guacamaya
- Anther at apex cruciform, dehiscing by 4 apical pores;
cephalar bracts reflexed 9. Schoenocephalium
10. Involucra! bracts surpassing the inflorescence; seed ob-
long, with longitudinal striae; apical appendage lacking or
the appendage papillate: Tribe Rapateeae
- Involucra! bracts equal to or shorter than the inflorescence
(maybe longer in Monotrema bracteatum); seed oval to
subglobose, white granulate, with a flat apical appendage,
powdery: Tribe Monotremeae
11. Spathaceous bract fused to the axis at the base, laterally
adnate to the elongated inflorescence; ovary 1-locular by
abortion 12. Spathanthus
- Involucra! bracts 2; ovary 3-locular
12. Anthers with an appendage; seeds exappendiculate
10. Rapatea
- Anthers exappendiculate; seeds appendiculate or not
11. Cephalostemon
13. Inflorescence short -pedunculate, barely exserted from the
subtending leaf sheath; filaments pubescent, exceeding the
corolla; anthers 1-celled; W Africa 16. Maschalocephalus
- Inflorescence usually exceeding the leaf sheaths; anthers 4-
celled
14. Spikelets stalked; anthers 4-celled, dehiscing by a subapi-
cal lateral pore 15. Windsorina
421
- Spikelets sessile; inflorescence capitate; dehiscing by an
apical pore 15
15. Inflorescence bibracteate; anther pore obliquely terminal
13. Monotrema
- Inflorescence polybractetae; anther pore transversely
terminal, capsule deeply emarginate 14. Potarophytum
I. Subfam. Saxofridericioideae Maguire (1958).
2
1. Tribe Saxofridericieae Maguire (1958).
10
3 1. Saxofridericia R.H. Schomb. Fig. 98A-G
8 Saxofridericia R.H. Schomb., Rapatea Frid. Aug. Saxo-Frid.
4 Reg.: 13 (1845).
7 Usually robust; peduncles subterminally axillary,
5 solitary or several, subtended by 1-2 bracts; in-
6
volucra! bracts spathaceous, bivalved, the margins
connate, therefore the involucre saccate, per-
forated by the expanding spikelets and full of
mucilage; inflorescence globose or hemispherical;
spikelets numerous; sepals free; petals obovate or
suborbicular, basally connate into a tube; fila-
ments adnate to the corolla tube; anthers lan-
ceolate, rugose, 4-lobed, 4-locular, with a single
subapical pore, ending in a somewhat cucul-
late appendage; seeds semiannulate, rugose
and striate. Nine spp., Guayana highlands and
lowlands.
2. Phelpsiella Maguire
9
Phelpsiella Maguire, Mem. N.Y. Bot. Gard. 10: 28 (1958).
Small; leaves linear; peduncles subterminal, soli-
tary. Inflorescence small, compressed, bispathate;
involucral bracts orbicular-obovate, the margins
basally connate; heads few-flowered, spikelets
11 sessile, bracteoles gradate; anthers bilocular,
dehiscing with 2 apical pores. Only one sp.,
Ph. ptericaulis Maguire, summit of Cerro Paru,
13 Venezuela.
3. Amphiphyllum Gleason
12
Amphiphyllum Gleason, Bull. Torrey Bot. Club 58: 332 (1931).
Leaves equitant; thyrse bracts several, the outer 2
rigid, more or less valvate and connate; spikelets
few, sessile; anthers 4-locular, dehiscing by a lat-
eral subapical pore, appendaged. Two spp., en-
demic to the summits of Cerro Duida and Cerro
14 Marahuaca, Venezuela.
422 Rapateaceae
4. Stegolepis Klotzsch ex Kornicke Fig. 100
Stegolepis Klotzsch ex Kiirnicke, Linnaea 37: 480 (1872);
Maguire, Mem. N.Y. Bot. Gard. 12: 69-89 (1965), rev.
Caudex unbranched; leaves ensiform or gramine-
ous, with rounded to elongated auricles located at
the top of the sheaths; peduncles several, axillary,
1-3m long; spathaceous bracts mostly lacking or
vestigial to small (subsect. Gleasoniana); heads
with 1-many spikelets, bracteoles often gradate in
size, imbricate; sepals membranous at base, often
connate into a short tube; petals differentiated
into a claw and limb, the claws connate into a tube;
filaments free or usually connate at the base, the
tube discrete or adnate to the corolla tube; anthers
4-locular, dehiscing by 1 or 2 subterminal pores,
the posterior anther lobes forming a somewhat
cucullate appendage; capsules strongly indurate,
membranaceous at the base, seeds prismatic or
pyramidal. Over 30 species, centered in Guayana.
Three sections and four subsections: sect.
Guianensis with many-flowered spheroidal heads
with little evidence of lateral compression, the
flowers in an thesis not reflexed; sect. Pungens with
laterally compressed many-flowered heads; and
sect. Pauciflora, number of spikelets reduced and
sepals reflexed.
5. Epidryos Maguire
Epidryos Maguire, Taxon 11: 57 (1962).
Epiphytic or occasionally terrestrial; axillary pe-
duncles numerous; blades chartaceous; thyrse
ebracteate, with few spikelets; petals lanceolate,
connate at the base; anthers bilocular, dehiscent
by 2 confluent apical pores; endocarp indurate
and separating from the exocarp; seed disciform,
testa black, distally apiculate, alveolate. Two dis-
junct spp., Colombia (Choc6), Panama, and the
Guianas.
6. Marahuacaea Maguire
Marahuacaea Maguire, Act. Bot. Venez. 14: 16-17 (1984).
Leaves auriculate; inflorescences many, axillary;
thyrse strongly compressed; thyrse bracts several,
2 spathaceous, rigid, valvate, connate; spikelets
few; anthers 4-locular, dehiscing by a lateral sub-
apical pore, appendaged. Two spp., Gran Sabana,
Venezuela.
2. Tribe Schoenocephalieae Maguire (1958).
7. Kunhardtia Maguire
Kunhardtia Maguire, Mem. N.Y. Bot. Gard. 10: 32 (1958).
Peduncles solitary, inflorescence spheroidal
or hemispheric; subtending involucra! bracts
bivalvate; spikelets red; bracts gradate in size;
flowers numerous, with zygomorphic corollas; se-
pals free; petals red, lanceolate, strongly exserted,
connate at the base; anthers linear, bilocular,
muriculate, dehiscing by 2 annular terminal pores;
seeds subpyramidal, striate. Two spp., sandy, wet
lowlands and tepuis of Amazonas, Venezuela.
8. Guacamaya Maguire
Guacamaya Maguire, Mem. N.Y. Bot. Gard. 10: 36 (1958).
Leaves glaucous; peduncles axillary; inflorescence
hemispherical, solitary, subtended by saccate,
connate involucra! bracts; spikelets red, numer-
ous, spirally arranged; sepals free; petals included
within the sepals and connate towards the base;
filaments adnate to the corolla tube; anthers ba-
sally 4-locular, upwardly bilocular, dehiscing by 2
transverse apical pores; seeds subprismatic and
striate. Only one sp., G. superba Maguire, Guayana
lowlands, Venezuela to Colombia.
9. Schoenocephalium Seubert
Schoenocephalium Seubert in Mart., Fl. Bras. 3 (1): 130 (1847);
Maguire, Mem. N.Y. Bot. Gard. 10: 37 (1958).
Sheaths partially open; ventral ligules lanceolate;
heads globose, bibracteate; spikelets radiate;
bracts gradate in size; sepals free; petals lan-
ceolate, included within the sepals, basally con-
nate; filaments adnate to the corolla tube; anthers
4-locular, apex cruciform, 4-porous; seeds sub-
prismatic to subpyramidal. Five spp., Upper
Orinoco and Rio Negro drainages.
II. Subfam. Rapateoideae Maguire (1958).
1. Tribe Rapateeae Endl. (1839).
10. Rapatea Aubl. Fig. 98H-K, 99
Rapatea Aubl., Hist. Pl. Gui. 1:305 (1775), Maguire, Mem. N.Y.
Bot. Gard. 12: 92-102 (1965), rev.
Rhizome short; leaves dorsiventral to ensiform;
inflorescences many, each with 2 involucra!
 Rapateaceae
bracts, closely subtending the spheroidal, com-
pressed or elongated thyrse; flowers sesssile or
pedicellate; receptacle paleaceous; petals rather
ephemeral; stamens mostly adnate to the short
corolla tube; anthers 4-locular, prolonged into a
cochlear appendage with a subterminal ventral
pore; ovary trilocular, each bearing a single
basally attached ovule, normally 2 of the 3 ovules
aborting, capsule indurate above the middle; seeds
oblong, longitudinally striate. Three sections: sect.
Paludosa with a nonelongate fiat to convex floral
receptacle, and spikelet bracts gradate, Ten spp.,
low wet habitats from the Guianas southward to
Bolivia and Mato Grosso, Brazil; sect. Elongata
with an elongate floral receptacle and spikelet
bracts nongradate, three spp., Guianas, Colombia,
and Brazil; and sect. Longipes with a floral recep-
tacle convex or hemispheric, bracts of the spike-
lets nongradate, five spp., Colombia, Venezuela,
and Brazil (Amazon and Orinoco basins).
11. Cephalostemon R.H. Schomb.
Cephalostemon R.H. Schomb., Rapatea Frid. Aug. Saxo-Frid.
Reg.: 9 (1845).
Duckea Maguire (1958).
Leaves gramineous; leaf sheath partially closed;
inflorescence subglobose to cylindrical, bibract-
eate, multispicate; bracts linear-lanceolate,
conspicuous; petals yellow, obcordate-obovate,
retusely apiculate, ephemeral; anthers 4-locular,
dehiscing by a longitudinally subterminal introse
slit or pore; capsule globose; seeds elliptic-oblong,
appendaged or not. Nine spp., Guayana region to
the Orinoco and Amazon basins, south to Mato
Grosso, Brazil, and Bolivia.
Maguire (1958) segregated four spp. of Cephalo-
stemon with nonappendaged seeds into a separate
genus, Duckea. Subsequently, Steyermark (1989)
placed these species back into Cephalostemon, be-
cause the two genera are not separable by gross
morphological features. Most likely, the species
with nonappendaged seeds are a monophyletic
group and will remain as a subgenus within
Cephalostemon.
12. Spathanthus Desvaux
Spathanthus Desvaux, Ann. Sci. Nat. 13: 45 (1828).
Leaves distichous; inflorescences 1-several,
solitary in leaf axils; involucra! bract solitary,
spathaceous, lanceolate; inflorescence spicate, ad-
nate to the spathaceous bract; spikelets numerous,
423
bracts numerous, distichous; petals lanceolate;
anthers 4-locular, dehiscing by a single apical
inaequilaterally toothed pore; ovary trilocular, 2
locules abortive, the fertile 1-ovulate; capsule
bivalvate, monospermous; seed oblong-elliptic,
striate. Two spp., wet lowland habitats from the
Guianas and S Venezuela and Colombia to the
Amazon basin of Brazil.
2. Tribe Monotremeae Maguire (1958).
13. Monotrema Kornicke
Monotrema Kornicke, Linnaea 37: 475 (1873).
Leaves linear, gramineous; inflorescences axillary,
1-2 per leaf axil, monocephalous, small, sub-
globose, 1-2 bracteate; spikelets subtended by
secondary bracts, the lower empty, the middle
fertile; petals yellow, lanceolate; anthers basally
4-locular, dehiscing by a single circular, oblique
apical pore; ovules sublaterally attached; seed
oval, white, finely muriculate, basally apiculate,
with an applanate appendage at the apex. Five
spp., wet savannas of the drainages of the upper
Rio Orinoco and upper Rio Negro of Venezuela
and Colombia.
14. Potarophytum Sandwith
Potarophytum Sandwith, Kew Bull. 1939: 20 {1939).
Rhizomes unbranched; inflorescences 1-2,
axillary, shorter than the leaves, subtended by
1-several prophylloid bracts, terminal, globose,
thyrse subtended by 4 free deltoid-ovate involu-
cra! bracts; spikelet bracts 6, imbricate; sepals free;
petals free, spathulate, yellow, small; filaments
free, anthers 4-locular, dehiscing by a single, ob-
lique, subterminal pore; ovules attached to the
basal portion of the central axis; capsule indurate,
ovoid, 3-lobed, each valve deeply bilobed; seed
oblong-subglobose, white, muriculate, with an
applanate appendage. Only one sp., P. riparium
Sandwith, restricted to the Kaieteur Plateau,
Guyana.
15. Windsorina Gleason
Windsorina Gleason, Bull. Torrey Bot. Club 50: 148 (1923).
Peduncles 3-9 per leaf axil, each peduncle sub-
tended at the base by 1 or 2 prophylloid bracts;
inflorescence cymose, cymose portion subtended
424 Rapateaceae

by a lanceolate bract; flowers borne on pedicels of Linder, P., Kellogg, E. 1995. Phylogenetic patterns in the com-
differing length and each subtended by succes- melinid clade. In: Rudall, P.J., Cribb, P.J., Cutler, D.F.,
Humphries, C.J. (eds.) Monocotyledons: systematics and
sively smaller imbricate bracteoles; sepals free; evolution. Royal Botanic Gardens, Kew, pp. 473-496.
petals free; anthers basally 4-locular, dehiscing by Maguire, B. 1958. Rapateaceae, in: The botany of the Guayana
a single subterminal pore; capsule ovoid, indurate; Highland. Part III. Mem. N.Y. Bot. Gard. 10: 19-49.
seed oval or subglobose, muriculate, with an Maguire, B. 1965. Rapateaceae. In: The botany of the Guayana
applanate appendage. Only one sp., W. guianensis Highland. Part VI. Mem. N.Y. Bot. Gard. 12: 69-102.
Malme, G. 1933a. Beitrage zur Kenntnis der siidameri-
Gleason, Potaro river in Guyana. kanischen Xyridazeen. Ark. Bot. 25: 1-18.
Malme, G. 1933b. Einige wiihrend der zweiten Regnellschen
Reise gesammelte Phanerogamen. Ark. Bot. 26A: 1-32.
16. Maschalocephalus Gilg & K. Schum. Mangenot, S., Mangenot, G. 1957. Nombres chromosomiques
nouveaux chez diverses dicotyledones et monocotyledones
Maschalocephalus Gilg & K. Schum., Bot. Jahrb. Syst. 28: 148
d' Afrique occidentale. Bull. Jard. Bot. Etat Brux. 27: 639-
(1900).
654.
Pilger, R. 1930. Rapateaceae. In: Engler, A. (ed.) Die
Gramineous; inflorescences sessile to very short natiirlichen Pflanzenfamilien, 2nd edn. lSa. W. Engelmann,
pedunculate, axillary, each subtended by a single Leipzig, pp. 59-65.
bract, cymose; bracteoles membranous; sepals Stevenson, D., Loconte, H. 1995. Cladistic analysis of monocot
families. In: Rudall, P.J., Cribb, P.J., Cutler, D.F.,
membranaceous, the tube subequal to the lobes; Humphries, C.J. (eds.) Monocotyledons: systematics and
petals basally fused; filaments attached to the co- evolution. Royal Botanic Gardens, Kew, pp. 543-578.
rolla tube; anthers introse, 4-locular; styles ex- Steyermark, J.A. 1974. The summit vegetation of Cerro
ceeding the perianth; seed ellipsoid, with a flat Autana. Biotropica 6: 7-13.
appendage. Only one sp., M. dinklagei Gilg & K. Steyermark, J.A. 1989. Flora of the Venezuelean Guayana-VI.
Ann. Mo. Bot. Gard. 75: 1565-1570.
Schum., seasonally wet sandy soils in Sierra Leone Takhtajan, A. 1997. Diversity and classification of flowering
and Liberia. plants. New York: Columbia Univ. Press.
Thorne, R. 1992. Classification and geography of the flowering
plants. Bot. Rev. 58: 225-348.
Selected Bibliography Tiemann, A. 1985. Untersuchungen zur Embryologie, Bliiten-
morphologie und Systematik der Rapateaceen und der
Xyridaceen-Gattung Abolboda (Monocotyledonae). Disser-
Arber, A. 1922. Leaves of the Farinosae. Bot. Gaz. 74: 80-94.
tationes Botanicae, Band 52. Vaduz: J. Cramer.
Augier, J. 1956. Les glucides de quelques monocoty!edones de
Venturelli, M., Bouman, F. 1988. Development of ovule and
l'ordre des Farinosae. Bull. Soc. Bot. Fr. 103: 251-253.
seed in Rapateaceae. Bot. J. Linn. Soc. 97: 267-294.
Carlquist, S. 1961. Pollen morphology of Rapateaceae. Aliso 5:
Vogel, S. 1981. Bestaubungskonzepte der Monokotylen und
39-66.
ihr Ausdruck im System. Ber. Dtsch. Bot. Ges. 94: 663-675.
Carlquist, S. 1966. Anatomy of Rapateaceae: roots and stems.
Weberling, R. 1989. Morphology of flowers and inflorescences.
Phytomorphology 16: 17-38.
Cambridge: Cambridge Univ. Press.
Carlquist, S. 1969. Rapateaceae. In: Tomlinson, P.B., Anatomy
of the monocotyledons (Metcalfe, C.R., ed.), Vol. 3. Oxford Zavada, M. 1983. Comparative morphology of monocot pollen
and evolutionary trends of apertures and wall structures.
Univ. Press, Oxford, pp. 128-145.
Bot. Rev. 49: 331-379.
Chase, M.W. eta!. 1995. See general references.
Cheadle, V., Kosakai, H. 1982. Occurrence and specialization
of vessels in Xyridales. Nord. J. Bot. 2: 97-109.
Chenegry, E.M. 1949. Aluminum in the plant world. Part II.
Monocotyledons and gymnosperms. Kew Bull. 1949: 463-
466.
Cronquist, A. 1981. See general references.
Dahlgren, R. et al. 1985. See general references.
Duvall, M.R., Clegg, M.T., Chase, M.W., eta!. 1993. Phyloge-
netic hypoteheses for the monocotyledons constructed
from rbcL sequence data. Ann. Mo. Bot. Gard. 80: 607-619.
Friilich, D., Barthlott, W. 1988. See general references.
Givnish, T., Evans, T., Sytsma, K. 1994. Molecular evolution
and adaptive radiation in South American elements of the
plant family Rapateaceae. Am. J. Bot. 81 (6, Suppl.): 158.
Givnish, T., Evans, T., Sytsma, K. 1995. The order Com-
melinales is polyphyletic and represents a striking case
of convergent evolution in monocots. Am. J. Bot. 82
(6, Suppl.): 132.
Huber, 0. 1988. Guayana highland versus Guayana lowlands, a
reappraisal. Taxon 37: 595-614.
Hutchinson, J. 1973. The families of flowering plants. Oxford:
Oxford Univ. Press.
 Restionaceae
Restionaceae 1
H.P. LINDER, B.G. BRIGGS, and L.A.S. }OHNSON2
Restionaceae R. Br., Prodr. 1: 243 (1810) (Restiaceae).
Evergreen, rushlike plants, generally dioecious
or rarely monoecious; caespitose, rhizomatous
or stoloniferous. Culms photosynthetic, simple or
branching verticillately or dichotomously; terete,
compressed or quadrangular; surface smooth,
tuberculate, pitted, striate or sulcate. Leaves gener-
ally reduced to sheaths in the adult plants; sheaths
caducous or persistent, tightly or loosely convo-
luted or flat, split to the base; sheath apex often
with large membranous lobes or a smaller mem-
branous margin; mucro absent or developed into a
small awn or rarely a leaflike blade. Leaf morphol-
ogy of juvenile culms often different, with rela-
tively larger mucro or leatblade; in some taxa this
morphology coexisting with adult morphology.
Inflorescence spicate or paniculate, sexually differ-
entiated or the two sexes alike. Spathes persistent
or caducous, obscuring the flowers or relatively
minute. Flowers aggregated into spikelets or soli-
tary, subtended by 1 or rarely 2 membranous to
bony bracts, which may be longer or shorter than
the flower; spikelets with 1-many flowers, often
with the lower bracts sterile. Flowers unisexual,
rarely hermaphrodite. Perianth of 6 tepals in
2 whorls, occasionally less or completely absent;
segments all similar or variously differentiated,
membranous to bony, frequently the outer lateral
tepals villous-carinate, occasionally deeply keeled
and then flowers appearing winged. Male flowers
with 3 stamens opposite the inner tepals, filaments
free or rarely fused, slender; anthers dorsifixed, 1(-
2)-thecate, dehiscing by longitudinal slits; usually
with a minute pistillode. Female flowers with sta-
mens wanting or reduced to staminodes; ovary
superior, free, 1-3-celled, with 1-3 styluli which
may or may not be fused below into a style, inner
surface stigmatic and finely branched, ovules soli-
tary in each cell and pendulous. Fruit a 1-3-locular
capsule or a 1-locular nut, which may be sessile or
1 H.P.L. has been responsible for the generic classification
of
the African and Madagascan taxa while the treatment of the
extra-African groups is by B. G.B. and L.A.S.J.
2 Deceased.
425
stipitate on a fleshy pedicel, and which may be
variously ornamented. Seeds rotundate to oblong-
triangular, smooth, striate to variously tubercled.
Endosperm starchy; embryos disc-shaped, small,
adpressed to the micropyle.
Fifty five genera and ca. 490 spp. in Africa,
Madagascar, Australia, Malesia, New Zealand and
South America, with the great majority of the
species restricted to southwestern Africa and
southwestern Australia.
VEGETATIVE MoRPHOLOGY. The Restionaceae
are evergreen, rush-like plants, growing to a maxi-
mum height of3.5 m (Cannomois virgata). Most of
the species form low mat-forming or tussocked
plants, not more than 1m tall. The plants are sym-
podial in growth, and can either form tight tus-
socks or develop short to long rhizomes, which
are covered in scales and/or hairs (Figs. 101-103).
Some of the taxa are stoloniferous (Anthochortus,
Restio harveyi, etc.).
The culms are photosynthetic and green, and
are the main photosynthetic organs of the plants.
Generally, the culms are terete, but quadrangular
culms occur in Restio tetragonus and Lepidobolus
quadratus, and more or less compressed culms
have evidently evolved several times, and occur
in several genera (Platycaulos, Restio, Elegia,
Hypodiscus, Lepyrodia, Desmocladus, Tremulina,
Platychorda and Eurychorda). Tuberculate culms,
often with stomata situated on the tubercles, occur
in Restio, Calopsis, Desmocladus and Onycho-
sepalum while striations due to various anatomi-
cal features occur in both African and Australian
taxa, as well as in Apodasmia chilensis in South
America. Branching culms are characteristic of
several genera. Branches develop from the axils of
sheaths, and the first branch sheath is a typical
prophyll, which in some species of Ischyrolepis
may be hairy. In several genera verticillate
branches occur (Elegia, Ischyrolepis, Rhodocoma,
Desmocladus, Baloskion).
The leaves are reduced to sheaths, which in
some genera (Elegia, Chondropetalum, Dovea,
Askidiosperma, Lepidobolus) are caducous. The
sheaths are split to the base, and frequently have
membranous apical lobes, which may be ligule
homologues. The leaf apices usually have mucros,
which in some species, especially on immature
plants, may be developed as terete or flattened
"blades" (Lepyrodia, Anthochortus graminifolius).
Plants may be glabrous or, especially in the
Australian genera, have sparse or dense unicellular
(e.g. Thamnochortus) or (more commonly) multi-
cellular hairs. These are uniseriate and simple (e.g.
426 Restionaceae
Fig. lOlA-F. Restionaceae. A-E Lyginia barbata. A Habit. B
Apex of staminate inflorescence. C Staminate flower with long
fused stamen tube. D Anther, note bilocular opening by two
slits, an unusual condition in Restionaceae. E Pistillate inflo-
rescence. FL. imberbis, dehisced capsule. (D. Mackay)
Desmocladus, Harperia) or multiseriate and plate-
like (fan hairs) or branched (Lepidobolus,
Kulinia). Fan hairs are a distinctive type in Lepta-
carpus, Meeboldina, Hypolaena, Chaetanthus
and allies; they are multiseriate, stalked, some-
times pel tate, and are often appressed and overlap-
Fig. 102A-F. Restionaceae. Hypodiscus procurrens. A Stami-
nate plant with long creeping rhizome. B Staminate spikelet. C
Staminate flower, with exserted anthers and lateral sepals
shorther than the petals. D Pistillate spikelet with solitary
flower. E Pistillate flower. F Nutlet. (Linder 1985)
ping or interlocking, forming a smooth covering
over the epidermis.
VEGETATIVE ANATOMY. The culms of the
Restionaceae show a high degree of specialisa-
tion for the combined functions of mechanical
strength, photosynthesis, and resistance to da-
mage due to desiccation (Gilg 1891; Malmanche
1919; Noel 1959; Cutler 1969; Botha et al. 1972;
Johnson and Cutler 1973; Van Greuning and Van
der Schijff 1973, 1974; Cheadle and Kosakai 1975;
Botha and Van der Schijff 1976; Linder 1984;
 Restionaceae
If
c the base of the support cells (e.g. in Platycaulos).
B However, characteristic of the African taxa are
Fig. 103A-D. Restionaceae. Leptocarpus elegans. A Plant with
pistillate inflorescence. B Staminate inflorescence. C Pistillate
spikelet. D Pistillate flower with enlarged bracts which are
shed with the fruit. (D. Mackay)
427
Briggs et al. 1990). The family has a characteristic
culm anatomy. One to several layers of palisade-
like chlorenchymatous peg cells are found directly
inside the epidermis. The chlorenchyma is sepa-
rated from a sclerenchymatous cylinder by a ring
of parenchyma 1-several cells thick. The pith has
scattered vascular bundles, and the centre of the
pith may be hollow.
The culm anatomy of different genera show
specialisations of the various tissues, which are
very useful for taxonomic and phylogenetic
purposes (Briggs and Johnson l998a; Meney and
Pate 1998).
The epidermis is usually of a single layer, but
several genera of African Restionaceae have 2 lay-
ers of epidermal cells. The epidermal cells are vari-
able in shape, often with sinuous and thickened
anticlinal walls. In a group of Australian genera
the epidermal cells flanking the stomata are elon-
gated and line the substomatal cavity. These may
be analogous to the "protective cells" found in
some African genera. The position of the guard
cells in the stomata varies from superficial to near
The chlorenchyma cells are in l - 3 rows, and are
interconnected by numerous "tubes", giving the
cells a very distinct appearance. This type of cell
is termed a peg cell. The form and number of rows
of chlorenchyma cells is taxonomically significant.
protective cells, which are modified chlorenchyma
cells lining the substomatal cavity. These have
been lignified forming a strengthened basket.
Various developments of these protective cells are
characteristic of different generic groups in Africa
(see Linder 1984). Similar but separate develop-
ments of protective substomatal cells occur in
Desmocladus and allied genera.
The sclerenchyma ring in the African genera
may have ridges, which alternate with the vascu-
lar traces. In some taxa these ridges reach the epi-
dermis, giving the culms a striate appearance
(Hypodiscus, Willdenowia, Anthochortus). Occa-
sionally, the ridges may be capped by modified
chlorenchyma cells (false pillar cells), which con-
nect the ridge to the epidermis. Analogous ridges
also occur in many Australian taxa. These, how-
ever, are developments from the sclerenchyma-
tous cap of the vascular traces, and so are opposite
the vascular bundles. These ridges are often
capped by modified parenchyma cells (pillar cells)
which reach the epidermis, and also result in a
striated culm appearance.
The occurrence of silica is widespread in the
family, either as crystals or as sand. The location
428 Restionaceae

of silica is often constant in each genus, being The inflorescence axis itself may be simple or
either in the epidermis (e.g. Lepyrodia) or in the branching. If it is branching, then each branch is
parenchymatous cells (e.g. Ischyrolepis), or as subtended by a foliar structure (spathe if on the
sand in the pith, or in the sclerenchymatous cells, main axis, spathellae if on primary or secondary
or in various combinations. The shape of the crys- branches). Frequently there is also striking sexual
tals does not appear to be constant. Silica bodies dimorphism (Fig. 103). Several lines of sexual
almost filling the cell in the parenchyma sheath dimorphism can be traced ("Oberfeld 1925). A
and outer layers of the sclerenchyma cylinder are common development in the female inflorescence
common (Cutler 1969). is the reduction of the number of flowers per
The vascular bundles are collateral. The periph- spikelet, finally resulting in a single fertile flower
eral bundles, which are confined to the scleren- subtended by numerous sterile bracts (e.g.
chyma sheath, have small xylem and phloem Hypodiscus, Fig. 102D; Willdenowia, some species
poles. The medullary bundles are largest near the of Desmocladus). A similar result may be achieved
centre of the culm. These possess protoxylem. by the reduction in the number of spikelets, and
Vessels occur in all the organs, and usually have the elongation of the spikelet nodes, resulting in
scalariform lateral wall pitting, although alternate solitary flowers, but without the sterile subtending
and reticulate pitting occurs occasionally. The pits bracts (e.g. Elegia).
are simple or bordered. Perforation plates are In the male inflorescences the tendency is
oblique or transverse, simple, scalariform, to maximise pollen production. Consequently all
scalariform-fenestrate or reticulate. bracts are often fertile, but some genera usually
The rhizomes have variable, parenchymatous have sterile lower bracts (e.g. Harperia,
cortices, usually with distinct endodermoid Baloskion). Frequently, male spikelets are pendu-
sheaths and amphivasal vascular bundles. The lous (Staberoha, Thamnochortus, Leptocarpus,
central ground tissue contains sclerenchymatous Meeboldina, Hypolaena, Chordifex microcodon).
cells next to the lignified bundle sheaths, and par- In Willdenowia the tepals and bracts are much
enchymatous or aerenchymatous cells in strands reduced, exposing the anthers, and so presumably
amongst the thickened cells. facilitating pollen dispersal.
The roots in transverse section show that root
hairs develop from normal epidermal cells. The FLOWER STRUCTURE. The Restionaceae are basi-
cortex is variable and may consist of parenchyma- cally dioecious, except for Coleocarya (which is
tous cells in various arrangements. The endoder- monoecious), while in some species of Lepyrodia
mis is 1(2)-layered, with passage cells present in and in Restio mahonii ssp. humbertii herma-
some species, and the pericycle is 1-2(10)-layered. phrodite flowers may occur. Two species of
Some species have well-developed aerenchyma Desmocladus and Ischyrolepis and also some
tissue. populations of Cytogonidium leptocarpoides and
Restio duthieae are parthenocarpic and female
INFLORESCENCE STRUCTURE. Although variation only. The basic condition of the perianth is of 6
in inflorescence structure is extensive in the tepals in 2 whorls.
family, it has received little attention (Linder 1984; The perianth of the male flowers shows little
Kircher 1986). variation, and for the most are similar to the
Flowers are usually aggregated into "spikelets" female flowers, only somewhat smaller, and more
(not homologous to grass spikelets), subtended chartaceous. In Willdenowia, however, the male
by a single bract, and shortly pedicellate. In flowers are very different, with the tepals and
several genera, however, the flowers are not in bracts reduced to membranous hairs.
spikelets, and in most of these, they are also sub- The perianth of the female flowers varies
tended by bracteoles. The spikelets sometimes dramatically (Linder 1992b). The most common
have at their bases prophylls. In genera allied to structure is a somewhat flattened flower, with the
Leptocarpus the flowers may be crowded obscur- outer lateral segments conduplicate, and often
ing spikelet structure. Generally the inflorescences villous-carinate. From this, several developments
contain numerous spikelets, but in a number of can be postulated. In Elegia, Chondropetalum,
genera or species the inflorescence is reduced to a Dove a and Askidiosperma the tepals are very hard,
single spikelet. This is taken to its extreme in usually rounded dorsally, and the flower is
Alexgeorgea, where there is a single subterranean rounded rather than flattened. The inner whorl is
spikelet in each female inflorescence (Carlquist often longer than the outer whorl. In Thamno-
1976; Briggs et al. 1990; Meney et al. 1990). chortus and Leptocarpus the keel of the outer
 Restionaceae 429

lateral segments is often much widened, so that micromorphology of the seed (Linder 1984). The
the flower is winged (Fig. 102D). In Cannomois, micropyle is formed by both integuments, and
Ceratocaryum, Hypodiscus, Willdenowia and there is no enlargement of the integuments in
the Desmocladus group of genera (including the micropylar region. The embryology is of the
Harperia, Onychosepalum etc.) the perianth lobes Polygon urn type. Proliferation of the antipodals is
are membranous, very reduced and sometimes occasional in the African taxa (Borwein et al. 1949;
vestigial or altogether lost. Krupko 1966; Kircher 1986; Rudall and Linder
The androecium shows little variation but is 1988), resulting in the "Poaceae variant of the
reduced to 2 stamens in 2-merous flowers such as Polygonum type" (Anton and Cocucci 1984). The
those of Eurychorda complanata. In most genera embryo sac at maturity contains large starch
the anthers are exerted from the perianth at an- grains (Hamann 1962) and the nucellar epidermis
thesis, but in Elegia, Chondropetalum, Dovea, at the micropylar end is uniseriate and enlarged
Askidiosperma, Leptocarpus, Hypolaena and (Rudall and Linder 1988). Endosperm formation
Meeboldina the anthers stay inside the perianth is Nuclear (Mlodzianowski 1964).
lobes, which gape at anthesis. The anthers are
mostly unilocular and open by a longitudinal slit; POLLEN MORPHOLOGY. The single pollen grains
bilobed anthers are found in Hopkinsia and are spheroidal to oblate and ulcerate with a polar
Lyginia (Fig. lOlD), while in Lyginia the filaments diameter of 15-50 [!ill. (Chanda 1966; Chanda and
are fused. Rowley 1967; Ladd 1977; Chanda and Ferguson
The ovary is fundamentally 3-locular (Kircher 1978; Linder 1984, 1987). The pollen walls show a
1986; Linder 1992a), with 3 free styluli from the well-developed foot-layer, colummellate inter-
apex. The inner surface of the styluli is receptive, stitium, and a well-developed tectum, with nu-
and the stigmas are much branched, with the merous pores penetrating the tectum and the
branches papillose. In the Desmocladus group the foot-layer (scrobiculi), as well as pores that only
single style is very long and slender. The stigma(s) penetrate the tectum. The pollen surface varies
are Dry. The ovary walls are heavily impregnated from smooth to rugose, usually with micro-
with tannins, which also occur in the anther walls. verrucate ornamentation, rarely with dense verru-
Reduction to a 1-locular ovary has occurred cae or spinules.
numerous times within the family, often leaving The apertures are variable within the family. In
traces of the aborted locules. Unilocular ovaries Ischyrolepis, Staberoha, Hopkinsia and Lyginia the
are either dehiscent or indehiscent. The number of annulus consists of a heavily thickened foot-layer,
styluli has also been reduced in several lineages, and the aperture diameter is less than 6 [!ill. A thin
but reduction in stylulus number is not always layer of endexine covers the ulcus and extends
matched in the reduction of the locule number. some distance under the margins. This aperture
The details of stylar morphology vary from genus type has been termed graminoid.
to genus, but still require investigation. The basal In the African genera, excluding Ischyrolepis
portions of the styluli may be fused into a style, and Staberoha, the foot layer is not thickened in
but the stylar traces still remain distinct through- the annulus region, but the tectum is raised to
out the fused portion. form a ring. This often results in a vestibulate an-
nulus. The margins are often somewhat irregular,
EMBRYOLOGY. The endothecium of the anthers with exine fragments present in the ulcus. These
contains spiral thickenings (Manning and Linder fragments are attached to the thin layer of endex-
1990). The epidermis of the anthers is heavily ine which covers the ulcus. The ulcus has a diam-
impregnated with tannins. The pollen mother cell eter of 4-10[tm. Linder and Ferguson (1985) call
divisions are successive. In two taxa (Hypodiscus this type African restioid (Fig. 104).
aristatus and Elegia racemosa) pollen is shed in The extra-African genera, excluding Lyginia and
the trinucleate stage, while in Chondropetalum Hopkinsia, also lack a thickened foot layer, and the
hookerianum it is shed in the binucleate stage tectum is not raised, so that there is no distinct
(Krupko 1962, 1963). annulus. The ulcus is still covered by a thin endex-
The ovules are atropous, bitegmic and tenuinu- ine, carrying exine fragments; the ulcus margin
cellate, each integument with an inner and outer is usually irregular and the diameter range
epidermis. The inner epidermis of the inner in- approximately 8-25 [!ill. Chanda (1966) refers to
tegument is heavily impregnated with tannins this aperture type as centrolepidioid, while Linder
(Rudall and Linder 1988), while the outer epider- and Ferguson (1985) call it Australian restioid
mis of the outer integument forms the surface (Fig. 105).
430
Fig. 104. Restionaceae. Restio bolusii, pollen showing raised
aperture margins and associated coarse granules ("African
restioid"), SEM X2000. (Photo H.P. Linder)
KARYOLOGY. The karyology of the African genera
is still largely unknown, except for a few isolated
counts. Some Australian genera have been studied
in some detail (Briggs 1963, 1966; Johnson and
Cutler 1973; Bell and Pate 1993) and display a
range of basic numbers: n = 6, 7, 9, 11, 12. The
chromosomes are small. Within the Australian
Restionaceae there is little variation in numbers
within genera: the only exception is in Baloskion,
but most genera are inadequately sampled. The
three counts published for the African taxa (n =
16, 20, Krupko 1962, 1966) differ from the avail-
able Australian counts.
FRUIT AND SEED. The fruit of the Restionaceae
develops from a 3-locular ovary, with 1 seed per
locule. In many taxa this basic structure is main-
tained, and the ovary develops into a 3-locular
capsule. The seed is 1- 3 mm long, except in
Alexgeorgea which has seeds 7-10mm long. Seeds
are ovoid, globular or oblong, in detail variously
shaped, with complex surface ornamentation
(Linder 1984; Briggs and Johnson 1998a,b). Fre-
quently, the number of fertile locules is reduced;
in many genera the ovary has a single fertile locule
and is indehiscent. In the Willdenowia group in
Africa the ovary walls become heavily lignified,
and the floral pedicels become fleshy and function
as elaiosomes. Hypolaena in Australia has a func-
tionally similar hard nut on a fleshy axis, but the
latter includes the thickened axis of a reduced
spikelet. The perianth in these groups is much
reduced, usually membranous, and shorter than
the nuts. Wind-dispersed small nuts occur in
Thamnochortus, Staberoha, Leptocarpus and in
Restionaceae
Fig. 105. Restionaceae. Leptocarpus tenax, pollen showing
large aperture with irregular margin and fragments of the
exine on the ulcus, the surface of the grain marked by
scrobiculi and small pores ("Australian restioid"), SEM
X 1900. (Photo C. Porter)
some species of Calopsis and Meeboldina. The pe-
rianth is persistent, cartilaginous or membranous,
and longer than the nut, and the outer lateral
tepals have well-developed keels that function
as wings. In these nuts the ovary wall is not or is
only slightly lignified. In other species of
Meeboldina the tepals bear slender awns and
long hairs that may aid dispersal. In Elegia and
Calopsis, and in several species in various other
genera, and some non-African taxa, nuts are
formed that may be shed with or without
the perianth, but the mode of dispersal is not
known. In the bizarre Alexgeorgea the flowers are
borne underground (Carlquist 1976), and the
nut develops underground, without obvious
dispersal mechanisms. The very extensive rhi-
zome system presumably results in some dis-
persal, and burrowing mammals may also play
a part. The endosperm is starchy, the starch
grains are described as the Pteridophyte type
(Czaja 1978). The embryos are disc-shaped,
adpressed to the micropylar wall of the seed and
largely undifferentiated.
REPRODUCTIVE BIOLOGY. The whole family
is wind-pollinated, and with the exception of
Coleocarya and Lepy rodia, all are dioecious. A few
species are known in which males may be very
scarce or are unknown, while the females have
 Restionaceae
very good seed set. This suggests the occurrence of
apomixis.
PHYTOCHEMISTRY. The flavonoid chemistry of
the Anarthriaceae (Williams et al. 1997) and
Restionaceae has been studied (Harborne 1979;
Harborne et al. 1985, Williams et al. in press). The
flavonoid patterns of the family are different from
those of either the Juncaceae or the Poaceae.
Within the family, several broad patterns are
evident.
The extra-African taxa lack proanthocyanidins,
which occur almost universally in the African
taxa. In addition, the extra-African taxa are
characterised by the presence of gossypetin. This
compound, which is very rare in the monocots,
also occurs in a few African species. The African
taxa contain common flavonols, flavones and
glycoflavones. Within the African group, the
genera Elegia and Chondropetalum are distin-
guished by the presence of glycosides of myricetin,
larycitrin and syringetin, flavonols not reported in
the Poales outside this group.
Two allied genera, Lepyrodia and Sporadanthus,
differ in that the former contains the amino acid
analogue S-methylcystine (which is unpalatable to
both vertebrate and invertebrate herbivores) and
lacks starch in the rhizome. Lyginia possesses an
unusual sugar, allose.
SUBDIVISION OF AND RELATIONSHIPS WITHIN
THE FAMILY. The first subdivision of the family
was based on the number of locules in the ovary,
with the Restionoideae (Restioideae) with 1-3
locular capsules and the Willdenowieae with 1-
locular nutlets (Masters 1878). Gilg-Benedict
{1930) produced an improved classification. This
system is primarily based on the number of
anther-thecae, with the Diplantherae (Lyginia and
Hopkinsia) with 2-thecate anthers, and the
Haplantherae with unithecate anthers. Within the
Haplantherae the African taxa can be separated by
their possession of protective cells.
AFFINITIES. The Restionaceae are allied to the
Poaceae, Ecdeiocoleaceae, Joinvilleaceae, Flagel-
lariaceae and Anarthriaceae by ulcerate, spheroi-
dal pollen grains, the presence of silica, apical
placentation, atropous ovules and Nuclear
endosperm formation. These families, excluding
Flagellariaceae, are also tenuinucellate (Linder
and Rudall 1993; Kellogg and Linder 1995). This
grouping is supported by variation in rbcL se-
quences (Duvall et al. 1993) and by chloroplast
inversions (Doyle et al. 1992). The relationship
431
between the Restionaceae and Centrolepidaceae is
still unclear. Hamann {1962, 1975) suggested that
Centrolepidaceae may be embedded in Restiona-
ceae, an opinion that has received much sup-
port (Dahlgren and Clifford 1982; Dahlgren and
Rasmussen 1983; Linder and Rudall1993; Kellogg
and Linder 1995), but this is inconsistent with pol-
len data (Linder and Ferguson 1985). The exact
relationship between Ecdeiocoleaceae, Anarthria-
ceae, Restionaceae and Centrolepidaceae is still
unclear, but is currently under investigation.
DISTRIBUTION AND HABITATS. Restionaceae
are most common on oligotrophous soils in
southwestern Africa and south-western Australia.
Elsewhere, especially in tropical or more temper-
ate environments, the family is found in marshy or
swampy habitats, which are effectively also low-
nutrient environments (Linder 1991). The family
shows root specialisations to enhance nutrient
uptake: these finely divided roots with very long
root hairs have been reported from New Zealand,
Australia and South Africa (Lamont 1982). It
has generally been assumed that mycorrhizas play
no part in the nutrition of Restionaceae, and
Mitchell and Read {1987) failed to find mycor-
rhizal associations with the roots of the African
Restionaceae, but Meney et al. {1993) recorded
their presence for a brief period during winter in
the roots of the West Australian Alexgeorgea
nitens and Lyginia sp. This suggests that there
may be brief periods during which a mycorrhizal
association may be important.
The majority of the species in the family occur in
fire-prone environments, and show distinctive ad-
aptations to cope with fire. The contrast between
sprouting and non-sprouting biology has been
explored for the Australian species. Pate et al.
{1991) and Bell and Pate (1993) were able to dem-
onstrate distinctly different growth forms and pat-
terns of resource allocation in the two strategies.
Sprouting species tend to have starch deposits in
the rhizomes, while non-sprouting species gener-
ally invest in a larger reproductive capacity. The
changes between sprouting and non-sprouting
appear to occur readily, and closely related species
may have alternative strategies (e.g. Bell and Pate
1993; Linder 1995a,b).
In theW estern Cape, South Africa, Restionaceae
are frequently locally dominant (Taylor 1978), and
range in altitude from sea level to 2100 m (Linder
et al. 1993). In Australia, the family is less abun-
dant. Swamp species tend to be locally dominant,
both in Australasia and in tropical Africa.
432 Restionaceae

PALAEOBOTANY. Hochuli (1979) presented evi- KEY TO THE GENERA

dence from fossil pollen data that Restionaceae
1. Anthers with 2 cells 2
may have been widespread in the northern hemi- - Anthers with 1 cell each 3
sphere during the Tertiary. The pollen, placed 2. Stamen filaments fused into a tube; culm margins not
in the pollen form-genus Milfordia, is known scabrid 2. Lyginia
from across Eurasia and North America. In addi- - Stamen filaments free; culm margins scabrid
tion, Hochuli (1979) interprets the Tertiary fossil 1. Hopkinsia
3. Plants from Africa or Madagascar; culms with protective
Rhizocaulon as restionaceous. However, Linder cells around substomatal cavities 4
(1987) showed that the Eurasian and North - Plants from Australasia, Malesia or S America; culms
American Milfordia pollen could equally well ap- without protective cells 33
ply to Flagellariaceae, Joinvilleaceae or Anarthria- 4. Style solitary 5
ceae, and that there was therefore no convinc- - Styluli 2-3 8
5. Fruit dehiscent 4. Ischyrolepis
ing reason why it should be accepted to be - Fruit indehiscent 6
restionaceous. 6. Perianth chartaceous; male spikelets nearly globose
Similar pollen is also recorded from the South- 3. Staberoha
ern Hemisphere, and from later in the Tertiary - Perianth cartilaginous; male spikelets oblong 7
7. Male spikelets erect; sheath apices persistent
and appears more likely to be Restionaceae. 11. Calopsis
Scholtz (1985) recorded Restionaceae from - Male spikelets pendent; sheath apices soon decaying
Namaqualand, southern Africa, from the Pale- 12. Thamnochortus
ocene, while the first Australian records are from 8. Styluli 2 9
the Eocene (Martin 1973; Stover and Partridge - Styluli 3 24
9. Ovary dehiscent 10
1973). By the late Tertiary Restionaceae are - Ovary indehiscent 11
recorded from all southern continents (South 10. Styluli fused at the base, or, if free, then the bracts longer
Africa: Thiergart et al. 1963; Coetzee and Rogers than the flowers 4. Ischyrolepis
1982; Coetzee 1986; Australia: Martin 1973; Stover - Styluli free, bracts shorter than the flowers
6. Chondropetalum
and Partridge 1973; New Zealand: Mcintyre 1965;
11. Culms simple 12
Mcintyre 1968). - Culms branching 17
The identity of the northern hemisphere pollen 12. Male flowers in spikelets 13
assigned to Milfordia hungarica and Milfordia - Male flowers racemose 16
incertae sensu Linder (1987) remains a problem, 13. Male spikelets pendent; nut enclosed by the perianth
3. Staberoha
which could have bearing on the evolution of the
- Male spikelets erect; perianth shorter than the nut, or, if as
southern flora. large, then hyaline 14
14. Nut flattened on one side; female spikelets usually with
ECONOMIC IMPORTANCE. Restionaceae are not several flowers 15. Cannomois
widely used. In South Africa a small thatching - Nut round in cross-section 15
15. Male bracts acute; female flowers always with a single
industry uses Thamnochortus insignis (Rourke flower 20. Hypodiscus
1974; Linder 1990). In the past few years - Male bracts rounded; female spikelets with several flowers
Restionaceae have also become popular horticular 16. Nevillea
subjects in southern Africa - of particular in- 16. Male flower tepals valvate, lanceolate 14. Ceratocaryum
terest are Chondropetalum tectorum and Elegia - Male flower tepals linear 21. Willdenowia
17. Female spikelets with several flowers 18
capensis. Some species have contributed to the - Female spikelets with a solitary flowers 21
formation of peat deposits in Australia that are 18. Female bracts hyaline-chartaceous 17. Hydrophilus
now being exploited, despite the rarity of such - Female bracts coriaceous to osseous 19
deposits there. 19. Bracts shorter than the perianth 6. Chondropetalum
- Bracts longer than the perianth 20
20. Nut less than 5mm long 11. Calopsis
CuRRENT TAXONOMY. The taxonomy of the - Nut more than 10mm long 15. Cannomois
Australasian Restionaceae is currently under 21. Bracts shorter than the perianth and nut 5. Elegia
revision as a result of work by B.J. Briggs and - Bracts taller than the perianth and the nut 22
L.A.S. Johnson (1998a), leading to a drastic divi- 22. Male bracts and perianth lobes similar, linear and hyaline;
sion of the currently recognised genera. Briggs male flowers racemose 21. Willdenowia
- Male bracts not linear, larger than the tepals; male flowers
and Johnson (1998b) also proposed to separate in spikelets 23
Lyginia and Hopkinsia into separate families: the 23. Female bracts osseous; male flowers in conelike spikelets
current treatment is conservative in retaining 19. Mastersiella
them in the Restionaceae. - Female bracts chartaceous; male flowers in a few-flowered
spikelets 18. Anthocortus
 Restionaceae 433

24. Sheaths usually deciduous; bracts shorter than the flowers, - Culm with 5 or more cavities in the pith; male spikelets
or if longer, then hyaline and lacerated 25 many-flowered 29. Harperia
- Sheaths persistent; bracts usually cartilaginous, often 46. Leaf lamina reflexed from the sheath; culms not striate,
overtopping the flowers 27 glabrous, branched 26. Empodisma
25. Ovary unilocular, indehiscent 5. Elegia - Lamina absent or not reflexed; culms mostly striate,
- Ovary 3-locular, usually at least some dehiscent 26 glabrous or hairy, simple or branched 47
26. Culms branching; bracts shorter than the flowers, 47. Female flowers subterranean except for styluli and stigmas
cartilaginous or bony 7. Dovea which emerge above ground level; rhizome and culm bases
- Culms simple; bracts longer than the flowers, hyaline sheathed by glossy scales; rhizomes long and slender
8. Askidiosperma 38. Alexgeorgea
27. Culms flattened; sheaths green with a stout mucro; seeds - Female flowers not subterranean; rhizome and culm bases
white with a fragile surface 9. Platycaulos sheathed by dull or glossy scales; rhizomes long or short
- Culms terete, or if flattened, then not with the above or plant caespitose 48
characters 28 48. Pericarp of nut woody 50. Hypolaena
28. Culms with a simple central axis, and with sterile and/or - Pericarp of nut thin-textured 49
fertile branches clustered at the nodes; male spikelets 49. Female flowers in distinct spikelets, spikelets not com-
usually pendent 13. Rhodocoma pound, with no bracts between the flower and subtending
- Culms usually branching; if simple, then without branches glume; flowers dorsiventrally flattened with winged lateral
clustered at the nodes; male spikelets usually erect 29 tepals 49. Leptocarpus
29. Fruit a capsule; if a nut, then the bracts narrowly - Female flowers in large clusters without distinct spikelet
lanceolate, concolorous 30 structure or in compound spikelets with small bracts
- Fruit a unilocular nut 32 between the flower and subtending glume; flowers not
30. Culms branching 10. Restio dorsiventrally flattened (sometimes laterally flattened) 50
- Culms simple 31 50. Male flowers with a swollen base; females with clusters
31. Bracts shorter than the flowers; spikelets numerous at of 3-5 flowers crowded into larger aggregations; culms
several nodes 6. Chondropetalum simple below the infloresence 51
- Bracts at least as long as the flowers; spikelets few to - Male flowers without a swollen base, female spikelets
several 10. Restio distinct or females crowded into large aggregations;
32. Culms branching 11. Calopsis culms simple or branched 52
- Culms simple 3. Staberoha 51. Fruiting perianth with awns or slender tepals much longer
33. Flowers not in spikelets, culms not striate 34 than the fruit 53. Chaetanthus
- Flowers in spikelets; culms striate or not striate 37 - Fruiting perianth less than 2 times as long as fruit
34. Ovary 1-locular; styluli 3 35 55. Apodasmia
- Ovary 3-locular or if 1-locular, then style 1 36 52. Female flowers in clusters of 3-5 crowded into larger ag-
35. Rhizomatous, the rhizome pubescent; culms simple gregations (without conspicuous spikelet structure); male
25. Winifredia inflorescences appearing similar to the females and with
- Caespitose, the base glabrous; culms branched exserted anthers or of distinct pendulous spikelets with
24. Calorophus non-exserted anthers 54. Dapsilanthus
36. Culm pith cavity circular; stomates scattered on culms; - Female flowers in distinct spikelets (compound spikelets);
leaf sheaths striate; ovary 3-locular 22. Lepyrodia male and female inflorescences very different in appear-
- Culm pith cavity angular; stomates arranged in horizontal ance; male spikelets pendulous and anthers not exserted
bands on culms; leaf sheaths not striate; ovary 1-3-locular 53
23. Sporadanthus 53. Female spikelets (compound spikelets) very slender, 1-2-
37. Fruit indehiscent 38 flowered; culms much branched 52. Stenotalis
- Fruit dehiscent 54 - Female spikelets (compound spikelets) not very slender,
38. Style 1 39 several- or many-flowered; culms branched or simple
- Styluli 2 or 3 46 51. Meeboldina
39. Monoecious with male spikelets terminating culms, female 54.Chlorenchyma not interrupted by pillar cells, girders or
spikelets in lower axils 27. Coleocarya enlarged epidermal cells; culm usually not striate 55
- Dioecious 40 - Chlorenchyma interrupted by pillar cells, girders or
40. Culms simple 41 enlarged epidermal cells; culm usually striate 58
- Culms branching 45 55. Flower attached to glume; capsule falling with glume and
41. Culm sheaths lacking or culms with 1-3 persistent sheaths perianth attached; female flowers with 4 tepals; seeds
42 smooth 34. Baloskion
- Culm sheaths more than 3 44 - Flower not attached to glume; capsule not falling at seed
42. Culms densely pubescent; male spikelets 2-7-flowered dispersal; female flowers with 6 tepals; seeds striate with
32. Kulinia longitudinal lines of convex cells 56
- Culms glabrous; male spikelets many-flowered 43 56. Female spikelets with > 5 flowers; culms simple or spar-
43. Culms strongly flattened 31. Catacolea ingly branched; chlorenchyma 2-3 cells thick 36. Acion
- Culms not flattened 30. Onychosepalum - Female spikelets !-several-flowered; chlorenchyma 1-cell
44. Culm sheaths caducous; culm with 1 central cavity or culm thick 57
solid 33. Lepidobolus 57. Sheaths lax; culms repeatedly branched 35. Guringalia
- Culm sheaths persistent; culm with 5 or more cavities in - Sheaths appressed; culms simple below the inflorescence
the pith 29. Harperia 37. Saropsis
45. Culm with 1 central cavity; male spikelets !-many- 58. Female spikelets with 1 flower 59
flowered 28. Desmocladus - Female spikelets with more than 1 flower 60
434 Restionaceae

59. Culms dimorphic with fertile culms erect but sterile culms curved awn. Inflorescence of compound flower
more branched and flexuose; style-branches 2; plants clusters or reduced to single flowers, the two sexes
without common and conspicuous galls 46. Loxocarya
- Culms all much branched; style-branches mostly 3; dissimilar; tepals bony. Male flowers with the
commonly with large galls that resemble much enlarged outer lateral segments conduplicate; stamen fila-
spikelets 47. Taraxis ments connate into a stout tapering columnar
60. Female spikelet structure not conspicuous, the glumes tube; anthers 2-lobed, dehiscing by 2 parallel slits;
scarcely overlapping, not forming a compact spikelet; ovary 3-locular, style with 3 stout bushy branches.
culms repeatedly branched 48. Tyrbastes
- Female spikelet structure conspicuous (multi-glumed Fruit a woody, 3-locular capsule. Seeds ovoid
spikelets with overlapping glumes), culms simple or or globose, encircled by a narrow hyaline
branched 61 flange, the surface with fine short spines or hooks
61. Culms flattened 62 arising from the corners of minute concavities.
- Culms terete 64
62. Male flowers with 2 stamens; anthers exserted; female
Three spp. SW of Western Australia. In well-
flowers with 4 tepals 41. Eurychorda drained or seasonally wet sites in oligotrophic
- Male flowers with 3 stamens; anthers not exserted; female kwongan soils in regions of moderate to relatively
flowers with 6 tepals 63 low rainfall.
63. Ovary 3-locular; styluli 3; seed surface of subangular or
slightly lobed cells in an irregular pattern 42. Platychorda
- Ovary 2-locular; styluli 2; seed surface of rhombic cells in 3. Staberoha Kunth
longitudinal lines 43. Tremulina
64. Male and female spikelets more than 10mm long 65 Staberoha Kunth, Enum. Pl. 3: 442 (1841}; Pillans, Trans. R.
- Spikelets less than 10mm long 66 Soc. S. Afr. 16: 384-390 (1928}; Linder, Bothalia 15: 396-397
65. Spikelets ovoid, more than 3 mm diam.; glumes rigid and (1995}.
black 44. Melanostachya
- Spikelets cylindrical, less than 3 mm diam.; glumes Caespitose, with simple culms to 80 em tall;
scarious-membranous 40. Dielsia
66 Chlorenchyma interrupted by pillar cells; (in all except C. sheaths persistent. Male inflorescence of several
microcodon) male spikelets erect and anthers exserted globose, usually pendulous spikelets, all bracts
39. Chordifex fertile. Female inflorescence of 1-4 stiffly erect,
- Chlorenchyma interrupted by enlarged epidermal cells; tapering spikelets, with all bracts fertile and much
male spikelets pendulous and anthers not exserted taller than the flowers. Flowers keeled or winged,
45. Cytogonidium
perianth papery to membranous. Styluli 2-3(1),
ovary unilocular, indehiscent, nut shed with the
1. Hopkinsia W. Fitzg. perianth. About nine spp. endemic to the south-
Hopkinsia W. Fitzg., J.W. Aust. Nat. Hist. Soc. 1: 33 (1904}.
western Cape Province, Africa.

Caespitose or rhizomatous, culms unbranched, 4. Ischyrolepis Steud.

flattened on one side and minutely scabrous on
the angles, sheaths lax, with a short mucro. Male Ischyrolepis Steud., Syn., Pl. Glum. 2: 249 (1855}; Linder,
Bothalia 15: 397-418 (1985}.
and female inflorescences similar; flowers in the
axil of a membranous bract, singly or several Caespitose, rhizomatous or mat-forming; culms
together at the upper nodes of the culms. Tepals branching, rarely simple or with verticellated
glabrous, hyaline; stamens 2-lobed, dehiscing branches, 0.1-2m tall; sheaths persistent. Male in-
by 2 separate slits; ovary 1-locular; style stout, florescence of 1 to many spikelets. Female inflo-
bushy. Fruit a nut, the outer pericarp thin, some-
rescence usually similar but with larger and fewer
what fleshy, shed with the bracts and perianth spikelets. Perianth cartilaginous to osseous, outer
attached. Seed surface patterned with convex
lateral tepals conduplicate, glabrous or villous,
isodiametric angular cells. Two spp. SW of
other tepals fiat. Stylebranches 2, fused at the base
Western Australia, in deep, seasonally moist,
into a persistent peg; ovary 1-2locular, dehiscent.
sandy soils near watercourses in regions of rather About 48 spp, endemic to S Africa, common in the
low rainfall.
southwestern and southern Cape Province.

2. Lyginia R. Br. Fig. 101

5. Elegia L.
Lyginia R. Br., Prodr.: 248 (1910}.
Elegia L., Mantissa Pl. Altera: 162 (1771); Pillans, Trans. R.
Soc. S. Afr. 16: 313-342 (1928}; Linder, Bothalia 15: 418-427
Caespitose to rhizomatous, culms unbranched, (1985}.
terete; sheaths tightly convoluted with a rigid Lamprocaulos Mast. (1878).
 Restionaceae
Caespitose or rhizomatous; culms simple or
branched, rarely with verticillate branches;
sheaths usually caducous. Male and female inflo-
rescences usually similar, much branched with
numerous small, often dark-coloured spikelets or
solitary flowers, spathes prominent, but males
often more branched and exposed; bracts ge-
nerally shorter than the cartilaginous, glabrous,
persistent perianth, tepals all similar, styluli 2-3,
ovary unilocular, indehiscent. About 35 spp.,
endemic to the southwestern and southern Cape
Province of S Africa.
6. Chondropetalum Rottb.
Chondropetalum Rottb., Program.: 11 (1772); Pillans, Trans.
R. Soc. S. Afr. 16: 297-313 (1928); Linder, Bothalia 15: 427-
430 (1985).
Caespitose or rhizomatous; culms simple, rarely
branching; sheaths caducous, rarely persistent.
Male inflorescences from several nodes, usually
branching, flowers aggregated into spikelets;
bracts bony, often dark-coloured, shorter than the
perianth. Female inflorescence similar, but with
larger and fewer flowers and spikelets. Perianth
cartilaginous to osseous, glabrous, persistent;
tepals all similar, or whorls of different lengths,
styluli 3(2), ovary 3-locular, usually dehiscent.
About 12 spp., endemic to the southwestern and
southern Cape Province.
7. Dovea Kunth
Dovea Kunth, Enum. Pl. 3: 457 (1841); Linder, Bothalia 15:431
(1985).
Caespitose, to 1.5 m tall; culms branching; sheaths
caducous. Male inflorescence much branched with
numerous flowers loosely aggregated into spike-
lets, bracts shorter than the perianth. Female
inflorescence with globular, 1-flowered spikelets,
perianth glabrous, tepals all similar, styluli 3,
ovary 3-locular, dehiscent; seed with a corky bark.
Only one sp., D. macrocarpa Kunth, southwestern
Cape, S. Africa.
8. Askidiosperma Steud.
Askidiosperma Steud., Syn. Pl. Glum. 2: 257 (1855); Linder,
Bothalia 15: 431-434 (1985).
Caespitose, culms simple; sheaths caducous.
Male inflorescence with simple branches, flowers
weakly aggregated into spikelets; all bracts fertile,
membranous, much taller than the flowers and
435
soon lacerated. Female inflorescence similar, flow-
ers usually larger, perianth glabrous, persistent,
tepals all the same, but whorls of different lengths.
Styluli 3, ovary 3-locular, occasionally only 1-2
locules fertile, usually dehiscent. About 11 spp.,
restricted to the mountains of the southwestern
Cape Province, S Africa, on dry slopes and damp
hollows.
9. Platycaulos Linder
Platycaulos Linder, Bothalia 15:64 (1984); Linder, Bothalia 15:
434-437 (1985).
Caespitose to mat-forming; culms branching,
more or less compressed; sheaths green, persis-
tent, with narrow hyaline margins and large
stout awns. Male inflorescence of several
racemose spikelets. Female inflorescence similar,
often with large or solitary spikelets, bracts all
fertile; perianth with outer lateral tepals con-
duplicate, glabrous or villous; styluli 3; ovary 1-2
locular, dehiscent, seed surface white and finely
ornamented. About eight spp., endemic to the
southwestern and southern Cape Province, S
Africa, generally found in seepages and wet
habitats. Dubiously distinct from Restio.
10. Restio Rottb.
Restio Rottb., Program.: 9 (1772); Pillans, Trans. R. Soc. S. Afr.
16: 210-297 (1928); Linder, Bothalia 15: 437-464 (1985).
Craspedolepis Steud. (1855).
Caespitose, rhizomatous or mat-forming, culms
branching or rarely simple; sheaths persistent.
Male flowers aggregated into spikelets, bracts
usually cartilaginous. Female inflorescence more
or less similar to male, but spikelets usually fewer
and larger. Perianth cartilaginous to bony, outer
lateral tepals glabrous or villous-carinate. Styluli
3, ovary 1-3 locular, usually dehiscent, seed vari-
ously ornamented or shiny. About 93 spp., com-
mon in the south-western and southern Cape,
reaching northwards to Zaire and Madagascar, in
diverse habitats.
11. Calopsis Beauv.
Calopsis Beauv. ex Desv. in Ann. Sci. Nat. 13: 44 (1828); Linder,
Bothalia 15: 464-471 (1985).
Leptocarpus sensu Pillans, Trans. R. Soc. S. Afr. 16: 342 (1928).
Caespitose; culms branching; sheaths persistent.
Male flowers aggregated into spikelets. Female
inflorescence similar, racemose or paniculate, the
436 Restionaceae
spikelets with many to few flowers, tepals
chartaceous to cartilaginous, the outer lateral
tepals glabrous, winged or villous-carinate. Styluli
3( -2,-1), ovary unilocular, indehiscent. About 23
spp., West and South Cape, as well as Natal. Dubi-
ously distinct from Restio.
12. Thamnochortus Berg.
Thamnochortus Berg., Descr. Plant. Cap. 353 (1767); Pillans,
Trans. R. Soc. S. Afr. 16: 358-384 (1928); Linder, Bothalia 15:
471-478 (1985).
Caespitose, or rhizomatous; fertile culms usually
simple, often with clusters of sterile culms devel-
oping from the nodes; sheath persistent, the upper
2/3 soon decaying. Male inflorescence paniculate,
spikelets lorate to linear, nodding. Female inflo-
rescence erect, paniculate or racemose, all bracts
fertile, obscuring the flowers. Perianth persistent,
cartilaginous, glabrous, outer lateral tepals often
strongly keeled to winged. Style solitary, ovary
unilocular, indehiscent, shed with the perianth.
About 34 spp., widespread in the southwestern
and southern Cape Province of S Africa. T. insignis
Mast. is widely used for thatching.
13. Rhodocoma Nees
Rhodocoma Nees in Lind!., Nat. Syst. Bot., edn. 2: 450 (1836);
Linder & Vlok in Plant. Syst. Evol. 175: 139-160 (1991).
Caespitose; fertile culms simple, usually with clus-
ters of sterile branches produced at the nodes;
sheaths persistent. Male inflorescence paniculate,
spikelets usually pendent. Female inflorescence
paniculate to racemose, erect. Perianth cartilagi-
nous, glabrous, tepals all the same. Styluli 3; ovary
3-locular, dehiscent. Seven spp. southern Cape
Province, S Africa.
14. Ceratocaryum Nees
Ceratocaryum Nees in Lind!., Nat. Syst. Bot., ed. 2:451 (1836);
Linder, Bothalia 15: 479-480 (1985); Linder inS. Afr. J. Bot.
61: 222-225 (1995).
Caespitose; culms simple; sheaths persistent, up-
per margins decaying or lacerate. Male inflores-
cence dense, spikelets indistinct or not formed,
paniculate, flowers with lanceolate, strawlike
tepals and bracts. Female inflorescence of several
racemose 1-flowered spikelets with numerous
sterile practs. Perianth membranous, smaller than
the nut, stipitate or sessile. Styluli 2, stout; ovary
unilocular; fruit a woody nut. About six spp.,
endemic to the southwestern and southern Cape
Province of S Africa.
15. Cannomois Beauv. ex Desv.
Cannomois Beauv. ex Desv. in Ann. Sc. Nat. 13: 43 (1828);
Pillans, Trans. R. Soc. S. Afr. 16: 410-419 (1928); Linder,
Bothalia 15: 480-482 (1985).
Caespitose to rhizomatous; culms simple or
branching; sheaths persistent. Male inflorescence
paniculate, with numerous small, several-flowered
spikelets. Female inflorescence of 1-several race-
mose spindle-shaped spikelets; bracts osseous;
flowers several per spikelet; perianth membra-
nous, as long as or shorter than the nuts. Styluli 2,
ovary unilocular; fruit a shiny black, somewhat
flattened, very woody nut. About seven spp., wide-
spread in the southern and southwestern Cape
Province of S Africa.
16. Nevillea Esterhuysen & Linder
Nevillea Esterhuysen & Linder, Bothalia 15: 66 (1984); Linder,
Bothalia 15: 482-484 (1985).
Caespitose; culms simple; sheaths persistent. Male
inflorescence of a few conelike spikelets with
numerous fertile small rounded imbricate bracts.
Female inflorescence racemose, with several 3-5
flowered spikelets, very different from the male.
Tepals chartaceous, glabrous, the outer laterals
conduplicate. Style branches 2; fused at base,
ovary bilocular, fruit 1-2 locular, indehiscent.
Two spp., mountains of the southwestern Cape.
17. Hydrophilus Linder
Hydrophilus Linder, Bothalia 15: 66 (1984); Linder, Bothalia
15: 484 (1985).
Caespitose with spreading subterranean stolons;
culms branching; sheaths persistent. Male inflo-
rescence of several spikelets with imbricate,
lanceolate, silvery spathes and bracts. Female
inflorescence similar, all bracts fertile; tepals
glabrous, the outer lateral tepals conduplicate.
Style branches 2, shortly fused at the base; ovary
unilocular, fruit a small soft nut. Only one sp., H.
rattrayi (Pillans) Linder, Cape Province of S
Africa, along streams in the arid margins of the
Cape Floristic Region, dubiously distinct from
Anthochortus.
 Restionaceae 437

18. Anthochortus Nees 21. Willdenowia Thunb.

Anthochortus Nees in Lind!, Nat. Syst. Bot., ed. 2: 451 (1836); Willdenowia Thunb. inK. Svenska Vetensk. Akad. Hand!. 11:
Linder, Bothalia 15: 484-487 (1985). 28 (1790); Pillans, Trans. R. Soc. S. Afr. 16: 419-431 (1928);
Phyllocomos Mast. (1900). Linder, Bothalia 15: 493-497 (1985).
Hypolaena sensu Pillans (1928). Nematanthus Nees (1830).

Mat-forming to caespitose, stoloniferous plants; Caespitose; culms usually branching, smooth or

culms branching; sheaths persistent, often with striate; sheaths closely convoluted, persistent,
fiat blades. Male inflorescence of several racemose with decaying apices. Male inflorescence panicu-
spikelets with many sterile bracts. Female in- late; flowers not aggregated into spikelets, the
florescence of several obscure single-flowered perianth and bracts filiform. Female inflorescence
spikelets with several sterile bracts, perianth racemose, spikelets 1-fiowered; perianth membra-
membranous or lacking, flowers sessile or pe- nous, sessile or stipitate; styluli 2; ovary unilocu-
dicellate. Styluli 2, often fused below; ovary lar; fruit a woody nut. About 12 spp., endemic to
unilocular; fruit a variously ornamented, woody the Cape Province of S Africa.
nut. About 15 spp., damp habitats and seepages,
endemic to the Western and Southern Cape of S
22. Lepyrodia R. Br.
Africa.
Lepyrodia R. Br., Prodr.: 274 (1810).

19. Mastersiella Gilg-Benedict

Mastersiella Gilg-Benedict in Nat. Pflanzenfam. edn. 2; 15A: 25
(1930); Linder, Bothalia 15: 487-488 (1985).
Hypolaena sensu Pillans (1928).
Caespitose; culms branching, spreading from a
central knot, without a rhizome; sheaths persis-
tent. Male inflorescence of several spindle-shaped
many-flowered spikelets. Female inflorescence a
single, 1-flowered spikelet with several cartilagi-
nous, sterile bracts. Perianth membranous; styluli
2, ovary unilocular; fruit a woody nut. Three spp.,
common in the Southern and Western Cape of S
Africa.
20. Hypodiscus Nees Fig. 102
Hypodiscus Nees in Lind!., Nat. Syst. Bot., ed. 2: 450 (1836);
Pillans, Trans. R. Soc. S. Afr. 16: 399-410 (1928); Linder,
Bothalia 15: 488-493 (1985).
Caespitose, rarely rhizomatous; culms simple,
striate or smooth; sheaths persistent. Male
inflorescence of 1-several racemose, many-
flowered spikelets. Female inflorescence of 1-
several racemose, 1-flowered spikelets, with
many sterile bracts. Perianth membranous or
lacking, sessile or pedicellate. Style branches 2,
often fused below, ovary unilocular; fruit a vari-
ously ornamented, woody nut, often with a fleshy
pedicel and with a variously ornamented cap.
About 15 spp., endemic to the Cape Floristic
Region of S Africa.
Dioecious, rarely monoecious or hermaphrodite,
caespitose or rhizomatous. Culms terete or
slightly compressed, smooth or minutely rugose.
Flowers not in spikelets, clustered on the culms,
males and females similar in dioecious species.
Flowers with (1- )2 bracts, tepals often rigid.
Female flowers with 3-locular ovary, styluli 3, free
or very shortly connate at the base, usually arising
close together on the ovary summit. Fruit a cap-
sule. 2n = 14. Twenty two spp., southeast and
southwest Australia, mostly in moist sites.
23. Sporadanthus F. Muell.
Sporadanthus F. Muell., Trans. N. Z. Institute 6: 389 (1874).
Dioecious, rhizomatous or caespitose. Culms
smooth or minutely rugose, with persistent
sheaths, mostly closely appressed. Inflorescence of
many flowers, not in spikelets, similar in males
and females. Flowers with 1 or 2 bracts. Female
flowers with 3- or 1-locular ovary, styluli 3 or 1,
arising from the outer corners of the ovary sum-
mit or when 1, not central on ovary, the bases
persistent; fruit a capsule or nut. 2n = 18. Seven
spp.; six southeast and southwest Australia, one
New Zealand, in moist sites. Only the New Zealand
species has been validly named under Spora-
danthus; four of the Australian species were
named under Lepyrodia and two are undescribed.
24. Calorophus Labill.
Calorophus Labill., Nov. Hoi!. Sp. Pl. 2: 78, t. 228 (1806).
438 Restionaceae

Dioecious, caespitose. Culms with transverse lines odes, male and female spikelets sometimes on the
of stomates usually visible on culm surface, same culm; rhizomatous. Culms simple or occa-
branched, trailing, forming tangled masses, with sionally branched, slightly tuberculate, with sev-
many persistent appressed sheaths. Inflorescence eral persistent lax sheaths. Male spikelets with
of solitary flowers each with 2 bracts and 6 rigid 8-16 flowers, all bracts fertile, outer tepals keeled.
tepals. Male flowers singly or in pairs in the upper Female spikelets 1-flowered, with 1-2 sterile lower
axils. Female flowers solitary, in upper axils or bracts, tepals 5, membranous, ovary 1-locular,
subterminal, with 3-locular ovary, only 1 loculus style 1. Fruit a bony nut, somewhat striate, shed
developing to maturity, styluli 3, slightly or widely with persistent perianth and bract, stylar beak
separated on ovary summit. Fruit a globular nut usually not persistent. 2n = 22. Only one sp., C.
with irregular longitudinal ridges indicating the gracilis S.T. Blake, southern Queensland and
positions of the nondeveloped carpels. Two spp., northern New South Wales.
Tasmania, one also Otway Ranges in Victoria, in
moist sites. 28. Desmocladus Nees
Desmocladus Nees in Lehm. Plant Preiss. 2: 56 (1846}.
25. Winifredia L.A.S. Johnson & B.G. Briggs
Dioecious, rarely parthenocarpic; caespitose or
Winifredia L.A.S. Johnson & B.G. Briggs, Telopea 2: 737
(1986}. rhizomatous, culms terete or flattened, sometimes
pubescent, smooth or striate or tuberculate,
Dioecious, rhizomatous. Culms simple with per- branched or with short lateral verticillate branch-
sistent sheaths. Inflorescences slender, males and lets. Spikelets solitary or in clusters. Male spikelets
females similar, with very short lateral few- 1-many-flowered, bracts numerous, all fertile or
flowered branches at each of several upper nodes, lower bracts sterile, flowers with (1-)5(-6) tepals,
flowers each with 2 bracts, not in spikelets, tepals 2 outer tepals keeled. Female spikelets 1-several
6. Female flowers with ovary 1-locular, 3-angled, flowered, usually with sterile lower bracts, tepals
with thickened pale ridges at the angles (perhaps 0-5, ovary 1-locular, style 1. Fruit an ovoid nut,
fruit dehiscent), styluli 3. Mature fruit and seed with a conical stylar beak, shed with persistent
not known. Only one sp., W. sola L.A.S. Johnson & pedicel and tepals if present. Sixteen spp., south-
B.G. Briggs, southwest of Tasmania, in a region of ern Western Australia, one sp. extending to South
very high rainfall. Australia, mostly in regions of rather low rainfall.

26. Empodisma L.A.S. Johnson & D.F. Cutler
Empodisma L.A.S. Johnson & D.F. Cutler, Kew Bull. 28: 383
(1974}.
Dioecious, caespitose or rhizomatous. Culms thin
and wiry, glabrous, branched and often flexuose,
with persistent sheaths. Male spikelets several to-
gether or rarely solitary, 1- or several-flowered.
Female spikelets solitary, axillary, 1-flowered.
Male flowers with 6 membranous tepals. Female
flowers with 4 or 6 tepals, ovary 1-locular, styluli 2
or 3. Fruit an ovoid or obovoid nut shed with
perianth and bract. 2n = 24. Two spp., one sp., in
southwest of Western Australia and one in east
Australia and New Zealand, in moist sites.
29. Harperia W. Fitzg.
Harperia W. Fitzg., J.W. Aust. Nat. Hist. Soc. 1: 34 (1904).
Caespitose or rhizomatous, culms terete, some-
times pubescent. Male and female inflorescences
similar, spikelets sessile, bracts especially in
females with a rigid, dark mucro or apical spine.
Male spikelets many-flowered, lower bracts often
sterile, tepals 5 or 6, membranous, 2 outer tepals
keeled. Female spikelets 1- or several-flowered,
bracts few, lower bracts sterile, tepals 4-6, similar
to males, ovary 1-locular, style 1. Fruit a nut, shed
with persistent pedicel and perianth, stylar beak
usually not persistent. Four spp., southern West-
ern Australia, mostly in seasonally dry regions.

27. Coleocarya S.T. Blake 30. Onychosepalum Steudel

Coleocarya S.T. Blake, Proc. R. Soc. Queensl. 54: 75 (1943}. Onychosepalum Steudel, Synopsis Plant Glum. 2: 249 (1855).

Monoecious; male spikelets solitary and terminal Caespitose, with glossy, glabrous cataphylls.
on culms, female spikelets in axils oflower intern- Culms simple, 1-few internodes. Male and female
 Restionaceae
spikelets similar, solitary and terminal or axillary
at several upper nodes, several- or many-flowered,
lower bracts sometimes sterile; bracts auriculate
with a dark slender mucro. Flowers with 2-5
membranous tepals. Female flowers with 1-locular
ovary, style 1. Fruit a smooth nut, shed with per-
sistent perianth. Three spp., west of Western
Australia, in moist sites in seasonally dry regions.
31. Catacolea B.G. Briggs & L.A.S. Johnson
Catacolea B.G. Briggs & L.A.S. Johnson, Telopea 7: 346 (1998).
Caespitose, with glossy dark brown to straw-
coloured basal sheaths; culms simple, strongly
flattened, straight or flexuose, of 1-2 internodes.
Male inflorescence terminal, of 1-3 shortly
pedunculate, pendulous many-flowered spikelets;
bracts acuminate and scarious, without a mucro
or awn. Female inflorescence of 1 terminal spikelet
with 1-3 flowers and several sterile lower bracts;
tepals membranous, (4) 5 (6); ovary 1-locular,
style 1. Fruit a nut; shed with persistent pedicel,
perianth, and stylar beak. Only one rare sp., west
of Western Australia, in a seasonally dry region.
32. Kulinia B.G. Briggs & L.A.S. Johnson
Kulinia B.G. Briggs & L.A.S. Johnson, Telopea 7: 349 (1998).
Caespitose, the base densely long-villous; culms
simple, often curled, densely pubescent, with 1-3
persistent sheaths. Male inflorescence a solitary
terminal spikelet, 2-7 flowered. Female inflores-
cence of 2-3 1-flowered spikelets. Bracts with a
dark mucro. Male flowers with 2-5 membranous
tepals; female flowers lacking tepals, ovary
unilocular, style single, fruit a thin-walled small
nut shed with attached pedicel. Only one rare
sp., south of Western Australia in a seasonally dry
region.
33. Lepidobolus Nees
Lepidobolus Nees in Lehmann, Plant. Preiss. 2: 66 (1846).
Caespitose or rhizomatous, the base long-villous;
culms simple, terete or quadrangular, smooth or
rugose or striate, sheaths caducous and lax. Male
and female spikelets terminal or axillary in upper
axils or rarely the females in lower axils. Male
spikelets large, many-flowered, with few or many
sterile bracts, tepals 5 (6), 2 outer tepals keeled.
Female spikelets similar to males but !-many-
flowered; bracts with rigid, black mucro; tepals
439
1-5, ovary 1-locular with 1 style. Fruit a nut; shed
with persistent perianth, pedicel and stylar beak.
2n = 24. Eight spp.; seven spp. south of Western
Australia, one sp. east of South Australia and in
western Victoria, mostly in seasonally dry regions.
34. Baloskion Raf.
Baloskion Raf., Fl. Tell. 4: 32 (1838).
Restio Rottb. p.p.
Caespitose or rhizomatous; culms simple below
the inflorescence or with numerous sterile lateral
branches; sheaths persistent. Male and female
spikelets similar, many-flowered, usually with sev-
eral sterile lower bracts. Outer tepals keeled, male
flowers with 6 tepals, female flowers with 4 tepals,
ovary 2-locular, styluli 2. Fruit a thin-walled cap-
sule, shed with persistent perianth and still at-
tached to the glume but eventually dehiscing. Seed
surface smooth, with elongated or isodiametric
cells. 2n = 14, 22, 24. Eight spp., E Australia, in
moist sites.
35. Guringalia B.G. Briggs & L.A.S. Johnson
Guringalia B. G. Briggs & L.A.S. Johnson, Telopea 7:353 (1998).
Restio Rottb. p.p.
Rhizomatous plants; culms repeatedly branched,
erect or sinuous, sheaths persistent, lax. Male
spikelets several, with 3-8 flowers and 7-8 sterile
lower bracts; female spikelets with 1 flower and 7-
10 sterile lower bracts. Tepals 6; males outer tepals
villous-carinate; females with sparsely pubescent
outer tepals, ovary 2-locular but often 1 loculus
aborted, styluli 2. Fruit a firm-walled capsule.
Seeds striate with longitudinal lines of convex
cells. 2n = 14. Only one sp., Sydney region of E
Australia, in well-drained or seasonally moist
sites.
36. Acion B.G. Briggs & L.A.S. Johnson
Acion B.G. Briggs & L.A.S. Johnson, Telopea 7: 353 (1998).
Restio Rottb. p.p.
Rhizomatous plants; culms unbranched or spar-
ingly branched, erect, sheaths persistent, lax.
Spikelets with 6-17 flowers and 5-20 sterile lower
bracts. Tepals 6; males with outer tepals villous-
carinate; anthers fully or partially exserted; fe-
males with glabrous tepals, ovary loculi and styluli
2. Fruit a firm-walled capsule. Seeds striate with
longitudinal lines of convex cells. 2n = 32. Two
440
spp., previously known as Restio monocephalus
R. Br. and R. hookeri D.I. Morris, Tasmania, in
well-drained or seasonally wet sites.
37. Saropsis B.G. Briggs & L.A.S. Johnson
Saropsis B.G. Briggs & L.A.S. Johnson, Telopea 7: 355 (1998).
Restio Rottb. p.p.
Shortly rhizomatous; culms unbranched below
the inflorescence but with numerous elongated
inflorescence branches, erect, sheaths persistent,
appressed. Spikelets narrow, males ca. 7-fiowered,
all bracts fertile; females 1-several flowered and
with several sterile lower bracts. Tepals 6; males
glabrous, 2 outer tepals keeled; females with usu-
ally ciliate tepals, ovary loculi and styluli 2. Fruit a
firm-walled capsule. Seeds striate with longitudi-
nal lines of convex cells. 2n = 14. Only one sp., S.
fastigiata (R. Br.) B.G. Briggs & L.A.S. Johnson,
eastern Australia in central coastal New South
Wales and also southeast Queensland; in season-
ally moist, sandy soils.
38. Alexgeorgea Carlquist
Alexgeorgea Carlquist, Aust. J. Bot. 24: 282 (1976).
Long-rhizomatous, cataphylls glossy; culms
sometimes pubescent, striate, with persistent
appressed sheaths, with short flattened lateral
branches or terete and repeatedly branched. Male
spikelets solitary and terminal, several-flowered.
Females solitary, sessile on the rhizomes, so that
the flowers are largely below ground, single-
flowered, with glossy bracts and very long styles,
with only the 3 long style branches and tips of
bracts above ground level; the 1-locular ovary
maturing below ground into a large (10-15mm
long), globular or obconical glossy nut. 2n = ca.
44. Three spp., southwest ofWestern Australia, in
mesic and seasonally dry regions, in well-drained
or peaty sites.
39. Chordifex B.G. Briggs & L.A.S. Johnson
Chordifex B.G. Briggs & L.A.S. Johnson, Telopea 7: 356 (1998).
Restio Rottb. p.p.
Caespitose or rhizomatous; culms terete, striate,
with persistent sheaths. Male and female inflores-
cences similar, the flowers in erect or pendulous
spikelets, with several or many flowers and several
or many sterile lower bracts. Tepals 5 or 6, 2 outer
tepals keeled; ovary loculi and styluli 2 or rarely 1.
Restionaceae
Fruit a capsule. Seeds striate with longitudinal
lines of convex cells (sometimes imprinted with a
further striate or irregular pattern) or the lines
exaggerated into sharp ridges of radially elongated
cells. Eighteen spp., south of Western Australia;
mostly in mesic regions.
40. Dielsia Gilg
Dielsia Gilg in Diels & Pritzel, Bot. Jahrb. Syst. 35: 88 (1904).
Rhizomatous, with stout villous rhizomes; culms
much branched, upper branches often flexuose
and sometimes forming dense trailing masses, ter-
ete, striate, with persistent appressed sheaths.
Male and female inflorescences similar, of solitary,
terminal, erect, elongated, cylindrical, many-
flowered spikelets. Tepals 6, outer 2 keeled, the
keel densely red-brown pubescent. Ovary 2-locu-
lar, styluli 2. Fruit a capsule. Seed surface smooth
with longitudinal lines of transversely elongated
rectangular cells. 2n = 22. Only one sp., southwest
of Western Australia, in seasonally wet sites in a
region of moderate rainfall.
41. Eurychorda B.G. Briggs & L.A.S. Johnson
Eurychorda B.G. Briggs & L.A.S. Johnson, Telopea 7: 357
(1998).
Restio Rottb. p.p.
Caespitose; culms simple, erect, striate, strongly
compressed, edges of culms minutely scabrid,
with persistent appressed herbaceous sheaths with
obtusely angled auriculate apices. Male and
female inflorescences similar with erect spikelets
on slender peduncles, solitary or a few together at
several upper nodes. Male spikelets 30-50-
flowered; females 5-13-fiowered; bracts aristate.
Tepals 4, outer tepals keeled, sparsely pubescent.
Stamens 2; ovary bilocular, styluli 2. Fruit a cap-
sule. Seeds with irregular pattern of subangular or
slightly lobed, fiat or slightly convex, isodiametric
cells. 2n = 24. Only one sp., eastern Australia from
southern Queensland to Tasmania, in wet or
seasonally wet sites.
42. Platychorda B.G. Briggs & L.A.S. John2son
Platychorda B.G. Briggs & L.A.S. Johnson, Telopea 7: 359
(1998).
Restio Rottb. p.p.
Caespitose; culms unbranched, erect, striate,
compressed but without scabrid edges, sheaths
persistent, appressed; apices gradually tapering,
 Restionaceae 441

auriculate. Male and female inflorescences simi- sheaths. Male spikelets pendulous; bracts spread-
lar, spikelets erect or the males somewhat pendu- ing lanceolate, acute; 30-50-flowered. Female
lous; several-flowered. Tepals 6, outer 2 keeled. spikelets erect, ovoid; bracts broad ovate, aristate;
Anthers not exserted; ovary 3-locular, styluli 3. 5-10-flowered. Tepals 5, outer 2 keeled and
Fruit a capsule. Seeds with irregular pattern of females with dense red-brown hairs along the keel;
subangular or slightly lobed, flat or slightly con- anthers not exserted; ovary bilocular, styluli 2.
vex, isodiametric cells. Two spp. southwest of Fruit a capsule. Seeds broad ellipsoid, surface cov-
Western Australia, in wet or seasonally wet sites. ered with convex cells in irregular longitudinal
lines (not separated by lines of flat cells as in
43. Tremulina B.G. Briggs & L.A.S. Johnson Chordifex). Only one sp., C.leptocarpoides (Benth.)
B. G. Briggs & L.A.S. Johnson, southwest ofWestern
Tremulina B.G. Briggs & L.A.S. Johnson, Telopea 7:361 (1998). Australia; in moist or seasonally moist sites.
Restio Rottb. p.p.

Caespitose; culms erect, strongly compressed or 46. Loxocarya R. Br.

terete, without scabrid edges, striate, sheaths per-
sistent, appressed, apices blunt, auriculate. Male
and female spikelets on slender lateral branches,
erect or the males pendulous; males 10-50-
flowered; females 1-8-flowered; both with several
sterile lower bracts; bracts not auriculate. Tepals
6, outer 2 keeled; anthers not exserted; ovary
bilocular, styluli 2. Fruit a capsule. Seeds with lon-
gitudinallines of flat, rectangular or rhombic cells
which are transverse or oblique to the long axis.
Two spp., southwest of Western Australia, in wet
or seasonally wet sites.
44. Melanostachya B.G. Briggs & L.A.S. Johnson
Melanostachya B.G. Briggs & L.A.S. Johnson, Telopea 7: 361
(1998).
Restio Rottb. p.p.
Dioecious, caespitose; culms terete, simple or
occasionally with 1-2 branches, striate, with
persistent appressed sheaths. Male and female
inflorescences similar, spikelets large, erect, 10-
20-flowered; bracts rigid, dark brown or black,
obtuse or with a short rigid acute tip. Tepals 6,
outer 2 rufous-villous carinate; ovary bilocular,
styluli 2. Fruit a capsule. Seeds broad ellipsoid,
with longitudinally elongated cells with sinuous
walls. Only one s:p., M. ustulatus (F. Muell ex
Ewart & Sharman) B.G. Briggs & L.A.S. Johnson,
southwest of Western Australia; in wet or season-
ally wet sites.
45. Cytogonidium B.G. Briggs & L.A.S. Johnson
Cytogonidium B.G. Briggs & L.A.S. Johnson, Telopea 7: 362
(1998).
Restio Rottb. p.p.
Loxocarya R. Br., Prodr. 249 (1810).
Megalotheca F. Muell, Fragm. Phytogr. Aust. 8: 98 (1873).
Rhizomatous; culms striate, fertile culms taller
and less branched than shorter, much-branched
and strongly flexuose vegetative culms. Inflores-
cences with several or many spikelets in a usually
branched inflorescence; male spikelets many-
flowered, females i-S-flowered. Tepals 4 or 6,
membranous, outer tepals keeled; ovary 1- or 2-
locular; styluli bushy, 1 or 2, the lower part con-
nate. Fruit a woody capsule, broadest near apex.
Seed oblong, with a longitudinal furrow encircling
the seed or on one side only; seed surface with
slightly convex, lobed cells. Five spp., south of
Western Australia, mostly in seasonally dry
regions, one sp. in a region of moderately high
rainfall.
47. Taraxis B.G. Briggs & L.A.S. Johnson
Taraxis B.G. Briggs & L.A.S. Johnson, Telopea 7: 363 (1998).
Culms stout near base, much branched, upper
branches slender, flexuose, forming tangled
masses, striate, glabrous except in sheath axils,
sheaths, appressed, persistent. Male inflores-
cences of small clusters of spikelets at several
upper nodes. Female spikelets solitary at several
upper nodes. Male spikelets 4-13-flowered, female
spikelets 1-flowered. Male flowers with 5(6) tepals,
2 outer tepals keeled. Mature female flowers, fruit
and seeds not known; tepals 3, style branches 3,
fused at base; spikelets usually transformed into
galls. Only one sp., southwest of Western Austra-
lia, in a region of high rainfall.

48. Tyrbastes B.G. Briggs & L.A.S. Johnson

Caespitose; culms terete, simple or sparsely
branched, striate, with persistent appressed Tyrbastes B.G. Briggs & L.A.S. Johnson, Telopea 7: 365 (1998).
442 Restionaceae

Culms slender, erect or trailing, flexuose and with thickened fleshy stalk, perianth, bracts and
much-branched, often forming tangled masses, subtending glume or the complete compound
striate, glabrous except in sheath axils, sheaths spikelet is dispersed. 2n = 24. Eight spp., south of
appressed, persistent. Inflorescences with several Western Australia, one of them also extending to
or many branches, with spikelets singly at the up- southeastern Australia including Tasmania, in
per nodes. Male spikelets more numerous than mesic and seasonally dry sites.
females, 4-10(24)-flowered, bracts mostly fertile.
Female spikelets 1-several-flowered, often with 1-
2 sterile lower bracts. Male flowers with 6 tepals, 51. Meeboldina Suessenguth
2 outer tepals keeled, inner tepals flat. Female Meeboldina Suessenguth, Boissiera 7: 20 (1943).
flowers with 6 tepals, ovary 2( -3)-locular, style Leptocarpus p.p.
short, style branches 2( -3). Fruit a thin-walled
capsule, seed with slightly convex, slightly lobed Culms striate, usually with appressed fan hairs.
cells in indistinct rows. Only one sp., southwest of Male spikelets usually numerous, many-flowered,
Western Australia, in a region of high rainfall. pendulous; apparent female spikelets multiply
compound; each 1-flowered true spikelet consists
only of a single flower stalk bearing 2 bracts, and
49. Leptocarpus R. Br. Fig. 103
this assemblage falls with the fruit. Male flowers:
Leptocarpus R. Br., Prodr. 250 (1810), nom. cons. tepals 5-6, stamens (2)3, not exserted. Female
Schoenodum Labill. (1806). flowers: tepals 6, usually keeled, often ciliate;
ovary 1-locular, style branches 3. Fruit a small,
Culms simple or branched, striate, covered with a thin-walled nut, shed with enclosing glume and
layer of appressed fan hairs. Inflorescence: males persistent perianth which may be modified to bear
and females very different, male spikelets usually finely hairy awns, hairs or lateral wings. 2n = 22.
numerous, many-flowered, pendulous; female Ten spp., south of Western Australia, in perma-
flowers in few or numerous simple spikelets with nently or seasonally wet sites.
several flowers and several to many rigid, shortly
awned bracts. Flowers with (4-)6 tepals; anthers
not exserted; female flowers dorsiventrally flat- 52. Stenotalis B.G. Briggs & L.A.S. Johnson
tened, tepals 6, outer lateral tepals winged; ovary Stenotalis B.G. Briggs & L.A.S. Johnson, Telopea 7: 368 (1998).
1-locular, style branches 3. Fruit a small, thin- Hypolaena p.p.
walled nut, shed with enclosing glume and persis-
tent perianth. 2n = 22. Three spp., south of Caespitose with ascending rhizomes. Culms much
Western Australia, of which L. tenax occurs also in branched, striate, with appressed fan hairs. Male
eastern Australia from southeast Queensland to spikelets usually numerous, several to many-
Tasmania and South Australia. (The application of flowered, pendulous; female flowers in very
the name Leptocarpus adopted here will require narrow spikelets with 1-2 flowers but which are
that a new conserved lectotype be designated. compound with 1 bract on the flower stalk. Male
Such conservation will be proposed in the inter- flowers: tepals 5, stamens (2)3, not exserted.
ests of nomenclatural stability.) Female flowers: tepals 6, ovary 1-locular, style
branches 3. Fruit a small, thin-walled nut, shed
50. Hypolaena R. Br. with enclosing glume and persistent perianth.
Only one sp., St. ramosissima (Gilg) B.G. Briggs &
Hypolaena R. Br., Prodr. 251 (1810). L.A.S. Johnson, south ofWestern Australia, in sea-
sonally moist sites.
Culms branching, glabrous or pubescent with
appressed or spreading fan hairs. Male spikelets
numerous, pedicellate and pendulous or sessile on 53. Chaetanthus R. Br.
short curved axillary branchlets, !-many- Chaetanthus R. Br., Prodr. 251 (1810).
flowered; flowers with (5- )6 tepals, outer 2-3 Prionosepalum Steudel, Synopsis Plant. Glum. 2: 266 (1855).
keeled; anthers mostly not exserted. Female Leptocarpus p.p.
spikelets compound but 1-flowered, with 1-2
bracts on the stalk of the flower; flowers with 6 Culms simple, striate, with appressed fan hairs.
tepals; ovary 1-locular, style (2- )3-branched. Fruit Male spikelets pendulous, on slender branches at
a nut, with hard pericarp, smooth or pitted; shed upper nodes; flowers with a swollen base, 5 mem-
 Restionaceae
branous tepals; anthers not exserted. Female inflo-
rescence of crowded clusters of spikelets each of
3-5 flowers, the clusters in compact heads, termi-
nal or at several upper nodes; tepals 6; outer tepals
rigid, acute or acuminate; ovary 1-locular; style
branches 3, basally connate. Fruit a small, thin-
walled nut, shed with enclosing glume and per-
sistent perianth which bears long hairs and
elongated hairy awns. 2n = 22. Three spp. south
of Western Australia, in seasonally wet sites in a
region of high rainfall.
54. Dapsilanthus B.G. Briggs & L.A.S. Johnson
Dapsilanthus B.G. Briggs & L.A.S. Johnson, Telopea 7: 369
(1998).
Leptocarpus p.p.
Culms branched, striate, with fan hairs. Inflores-
cences branched; males variable; females com-
pound, with clusters (true spikelets) of 3-5 very
small flowers each with 1 bract on the flower stalk
crowded in larger aggregations. Tepals 4-6, hya-
line or scarious, acute or acuminate; outer tepals
keeled; stamens 2 or 3, anthers mostly exserted;
ovary 1-locular; style branches 3, fused at base.
Fruit a small, thin-walled nut, shed with enclosing
glume, bract and persistent perianth. Four spp.;
three in seasonally wet sites in northern Australia
and monsoonal areas of southern New Guinea
(two of these also in the Aru Islands), as well as
one species in southeast Asia (Malaysia, Cambo-
dia, Thailand and the southeastern Chinese island
ofHainan).
55. Apodasmia B.G. Briggs & L.A.S. Johnson
ApodasmiaB.G. Briggs & L.A.S. Johnson, Telopea 7:371 (1998).
Leptocarpus p.p.
Dioecious, rhizomatous. Culms unbranched, stri-
ate, glabrous to densely pubescent; with fan hairs.
Male spikelets erect or pendulous, with partly ex-
serted anthers. Female inflorescences, with clus-
ters (true spikelets) of 3-5 very small flowers each
with 1 bract on the flower stalk crowded in larger
aggregations. Tepals hyaline or scarious, acute or
acuminate; outer tepals keeled. Male flowers with
a globular swollen base, 4-6 tepals and 3 stamens,
anthers mostly exserted. Female flowers with 5-6
rigid tepals; ovary 1-locular; style branches 3,
fused at base. Fruit a small, thin-walled nut, shed
with enclosing glume, bract and persistent peri-
anth. Four spp., one each in the south of Western
and eastern Australia, New Zealand and Chile.
443
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Restionaceen im Vergleich mit Gramineen und verwandten
Familien. Diss. Bot. 94.
Krupko, S. 1962. Embryological and cytological investigations
in Hypodiscus aristatus Nees (Restionaceae). J. S. Afr. Bot.
28: 21-44.
Krupko, S. 1963. Macrosporogenesis and embryo sac develop-
ment in Chondropetalum hookerianum (Mast.) Pillans
(Restionaceae). Acta. Soc. Bot. Pol. 32: 17-190.
Krupko, S. 1966. Some loose embryological and cytological
observations on members of Restionaceae family. Bull. Soc.
Amis Sci. Lett. Poznan Ser. D 7: 59-67.
Ladd, P.G. 1977. Pollen morphology of some members of the
Restionaceae and related families, with notes on the fossil
record. Grana 16: 1-14.
Lamont, B. 1982. Mechanisms for enhancing nutrient uptake
in plants, with particular reference to Mediterranean South
Africa and Western Australia. Bot. Rev. 48: 597-689.
Linder, H.P. 1984. A phylogenetic classification of the genera
of the African Restionaceae. Bothalia 15: 11-76.
Linder, H.P. 1987. The evolutionary history of the Poales/
Restionales- a hypothesis. Kew Bull. 42: 297-318.
Linder, H.P. 1990. A review of the African Restionaceae.
Contrib. Bolus Herb. 14: 209-264.
Linder, H.P. 1991. A review of the southern African Restiona-
ceae. Contrib. Bolus Herb. 13: 209-264.
Linder, H.P. 1992a. The gynoecia of Australian Restionaceae:
morphology, anatomy and systematic implications. Aust.
Syst. Bot. 5: 227-245.
Linder, H.P. 1992b. The structure and evolution of the female
flower of the African Restionaceae. Bot. J. Linn. Soc. 109:
401-425.
Linder, H.P. 1995a. Ceratocaryum pulchrum, a new restioid
from the Bredasdorp plains. S. Afr. J. Bot. 61: 222-225.
Linder, H.P. 1995b. Restio mlanjiensis, a new species of
Restionaceae from south-central Africa. Kew Bull. 50: 623-
625.
Linder, H.P., Ferguson, I.K. 1985. On the pollen morphology
and phylogeny of the Restionales and Poales. Grana 24:
65-76.
Linder, H.P., Rudall, P.J. 1993. The megagametophyte in
Anarthria (Anarthriaceae, Poales) and its implications for
the phylogeny of the Poales. Am. J. Bot. 80: 1455-1464.
Linder, H.P., Vlok, J.H., McDonald, D.J., Oliver, E.G.H.,
Boucher, C., Van Wyk, B.-E., Schutte, A. 1993. The high
altitude flora and vegetation of the Cape Floristic Region,
South Africa. Opera Bot. 121: 247-261.
Malmanche, L.-A. 1919. Contribution a !'etude anatomique
des eriocaulonacees et des families voisines: restiacees,
centrolepidacees, xyridacees, philydracees, mayacacees.
These Fac. Sci., Paris. Paris: Imprimeries Girault.
Manning, J.C., Linder, H.P. 1990. A cladistic analysis of pat-
terns of endothecial thickenings in the Poales/Restionales.
Am. J. Bot. 77: 196-210.
Martin, H.A. 1973. The palynology of some Tertiary and Pleis-
tocene deposits, Lachlan River valley, New South Wales,
Aust. J. Bot. Suppl. 6: 1-57.
Masters, M.T. 1878. Restiaceae. In: De Candolle, A., De
Candolle, C. (eds.) Monographiae Phanerogamarum 1.
Masson: Paris, pp. 218-398.
Mcintyre, D.J. 1965. Some new pollen species from New
Zealand Tertiary deposits. N. Z. J. Bot. 3: 204-214.
Mcintyre, D.J. 1968. Further new pollen species from New
Zealand Tertiary and uppermost Cretaceous deposits. N. Z.
J. Bot. 6: 177-204.
Meney, K.A., Pate, J.S. (eds.) 1998. Australian rushes- biology,
identification and conservation of Restionaceae and allied
families. Perth: University of Western Australia Press.
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phology, anatomy, phenology and reproductive biology of
Alexgeorgea spp. (Restionaceae) from south-western West-
ern Australia. Aust. J. Bot. 38: 523-541.
Meney, K.A., Dixon, K.W., Scheltema, M., Pate, J.S. 1993. Oc-
currence of vesicular mycorrhizal fungi in dryland species
of Restionaceae and Cyperaceae from SW Western Austra-
lia. Aust. J. Bot. 41: 733-37.
Mlodzianowski, F. 1964. The structure and the later stages
of development in the embryo sac of Thamnochortus
fruticosus Berg (Restionaceae). Bull. Soc. Amis Sci. Lett.
Poznan Ser. D 4: 3-11.
Noel, A.R.A. 1959. The stem anatomy of Restio triticeus Rottb.,
Bobartia indica L., and Cadaba juncea (Sparm.) Harv. J.S.
Afr. Bot. 25: 357-370.
Pate, J.S., Meney, K.A., Dixon, K.W. 1991. Contrasting growth
and morphological characteristics of fire-sensitive (obligate
seeder) and fire-resistant (resprouter) species of Restio-
naceae (S. Hemisphere restiads) from south-western
Australia. Aust. J. Bot. 39: 505-525.
Rourke, J.P. 1974. On restios and roofs. Veld Flora 4: 57-59.
Rudall, P.J., Linder, H.P. 1988. Megagametophyte and nucellus
in Restionaceae and Flagellariaceae. Am. J. Bot. 75: 1777-
1786.
Scholtz, A. 1985. The palynology of the upper lacustrine sedi-
ments of the Arnot Pipe, Banke, Namaqualand. Ann. S. Afr.
Mus. 95: 1-109.
 Restionaceae/Ruppiaceae
Stover, L.E., Partridge, A.D. 1973. Tertiary and Late Creta-
ceous spores and pollen from the Gippsland Basin, south-
eastern Australia. Proc. R. Soc. Victoria 85: 237-286.
Taylor, H. C. 1978. Capensis. In: Werger, M.J.A. (eds.) Biogeog-
raphy and ecology of southern Africa. The Hague: Junk.
Thiergart, F., Frantz, U., Raukopf, K. 1963. Palynologische
Untersuchungen von TertHirkohlen und einer Oberflachen-
probe nahe Knysna, Stidarika. Adv. Front. Plant Sci. 4: 151-
178.
Dberfeld, M. 1925. Beitrage zur Kenntniss des sexuellen
Dimorphismus der Restionaceae. Bot. Jahrb. Syst. 60: 176-
206.
Van Greuning, J.V., Vander Schijff, H.P. 1973. Chlorenchyma
and sclerenchyma in the culms of Willdenowia Thunb. and
Hypodiscus. Ber. Dtsch. Bot. Ges. 86: 537-550.
Van Greuning, J.V., Vander Schijff, H.P. 1974. External mor-
phology of the genera Hypodiscus Nees and Willdenowia
Thunb. and key to genera and species based on external
morphological properties. Kirkia 9: 331-347.
Williams, C.A., Harborne, J.B., Greenham, J., Briggs, B.G.,
Johnson, L.A.S. 1997. Flavonoid evidence and the classifica-
tion of the Anarthriaceae within the Poales. Phytochemistry
45: 1189-1196.
Williams, C.A., Harborne, J.B., Greenham, J., Briggs,
B.G., Johnson, L.A.S. (in press) Flavonoid patterns and the
revised classification of Australian Restionaceae.
Phytochemistry.
445
Ruppiaceae
R.R. HAYNES, L.B. HOLM-NIELSEN,
and D.H. LEs
Ruppiaceae Horan. ex Hutch., Fam. Fl. Pl. 2: 48 (1934), nom.
cons.
Annual or rarely perennial submersed glabrous
herbs growing in fresh or brackish waters. Roots
fibrous, few, non-septate, from lower nodes of
stems. Stems slender, branched or unbranched,
terete, often dimorphic, the lower stems rhizoma-
tous, the upper erect, leafy, the tips not modified
into turions. Leaves alternate to subopposite, di-
vided into blade and stipular sheath encircling the
stem; blade entire below, serrulate above, 1-
veined, thin, linear; infravaginal scales present;
turions rare, with 1 internode, the leaves undiffer-
entiated. Inflorescence a 1-few-fiowered capitate
axillary or terminal spike, at first enclosed by 2
subopposite foliage leaves. Peduncle elongating at
an thesis, elevating the inflorescence to or near the
water surface, or remaining short, occasionally
becoming twisted in fruit, then pulling the devel-
oping fruit below the water surface. Flowers per-
fect, hypogynous, actinomorphic; tepals absent.
Stamens 2, sessile; anthers extrorse, with two
bisporangiate thecae dehiscing by longitudinal
slits; connective broad. Carpels (2)4( -16), distinct,
each forming a pistil with a sessile, peltate,
lobed stigma and a single lateral, more or less
pendulous, campylotropous, bitegmic, and cras-
sinucellar ovule. Fruits long-stipitate or sessile
drupes, the outer soft layers decaying, the inner
sclerotic layers persistent and opening via a small
lid at germination. Seeds exalbuminous, embryo
with enlarged hypocotyl.
Subcosmopolitan, the single genus variously
regarded as a single variable species or as com-
prising about ten species.
VEGETATIVE MORPHOLOGY (mostly after Tomlin-
son 1982). Ruppia is an aquatic herb growing en-
tirely submersed (Fig. 106) sometimes surviving
the winter by means of a buried rhizome, but more
often the plant is annual. The roots are un-
branched and arise singly at the nodes. Turions
are described for the Australian R. tuberosa, where
they are distal swellings of the shoots consisting of
a single thickened, starch-filled internode.
446 Ruppiaceae

plasmatic trichoblasts that are distinct in ap-

pearance, but not in size, from adjacent cells of
the rhizodermis. Well-developed air lacunae are
present in the mesophyll of the leaf and cortex of
the stem and root. The epidermis contains chloro-
phyll and is the chief photosynthetic tissue of the
plant. Stomata are absent. The vascular system in
the leaf is reduced to a single median vein, directly
continuous with the vascular system of the stem,
together with 2 obscure lateral veins in the leaf
margin. The central xylem in the leaf, stem, and
root are largely represented by a xylem lacuna.
Tannin cells are common elsewhere in the plant
body; vessels, silica bodies, and calcium oxalate
are lacking.
The sieve-tube plastids have cuneate protein
bodies but no starch or filamental protein.

INFLORESCENCE AND FLOWER STRUCTURE. The

inflorescence terminates the main axis and vegeta-
tive growth continues for a short time from the
axils of the foliage leaves below the inflorescence.
Fig. 106A-J. Ruppiaceae. Ruppia maritima. A Habit. B Tip of
These renewal shoots in turn terminate as inflo-
shoot with sheathing leaves (stipules adnate) enclosing inflo- rescences, resulting in the sympodial pattern of
rescence, above, and stipular sheath (partially cut away), two growth.
young leaves, and peduncle of inflorescence, below. C Inflo- The inflorescence consists of 2 perianthless
rescence, from above, pollen already shed from stamens of flowers (rarely more) subopposite to each other.
lower flower. D Lowet flower, side view, two carpels and two
half-anthers removed. E Inflorescence, flower to left with de- Each flower has 2 median sessile stamens, which
veloping carpels and elongating gynophores, flower to right are bithecate and tetrasporangiate, and open with
with unfertilized carpels. F Carpel, vertical section, diagram- longitudinal slits. The connective is broad and has
matic. G Pollen grain. H Mature drupe. I Endocarp, side view, a dorsiventral outgrowth. The gynoecium consists
lid to left. J Same, adaxial view, with lid removed, lid to right. of usually 4 ascidiate carpels each of which bears a
(Haynes 1978)
single ovule (Fig. 106E).
The interpretation of the outgrowth of the sta-
men connective has been a matter of debate;
The leaves are distichously arranged and sepa- Gamerro (1968) has given a summary of this
rated by long internodes, which are suppressed in controversy. At face value, the outgrowth seems
the distal portions of the shoot. The leaves have to be completely homologous with the appendage
an open, tubular sheath sometimes interpreted on the connective of Potamogeton. Poluszny
as "adnate stipules" (Fig. 106B, although true and Sattler (1974) have shown, however, that
stipules of course are absent from monocotyle- this enation develops after stamen inception, un-
dons). The blade has a conspicuous middle vein like in Potamogeton, but equally as in Groen-
and reaches a length of !Scm and width of about landia.
2 mm. An apical pore is not developed. The
sequence of leaves on new branches is always EMBRYOLOGY. Anther wall formation is appar-
commenced by a bicarinate prophyll. A pair of ently unknown. The tapetum cells are pre-
intravaginal scales (squamules) is present in the dominantly !-nucleate. The walls eventually
axils of the leaves. The squamules are non- disintegrate and a periplasmodium is formed. Mi-
vasculated, but their cells are densely cyto- crosporogenesis is of the Successive type. The tet-
plasmatic with conspicuous nuclei. rads are usually isobilateral. The mature grains
are then 3-celled. In the ovule, the micropyle is
VEGETATIVE ANATOMY (Tomlinson 1982). The formed by both integuments. The archesporia! cell
anatomy of Ruppia is much reduced in relation to cuts off a primary parietal cell that gives rise up to
its aquatic life style. The root hairs are often long- 7 parietal layers below the nucellar epidermis. The
persistent; they are produced from densely cyto- chalaza! megaspore of a linear or T -shaped tetrad
 Ruppiaceae
develops into a Polygonum-type embryo sac.
Endosperm formation is Helobial (Nuclear after
Graves 1908).
Embryo formation follows the Caryophyllad
type. The ripe seeds lack endosperm and peri-
sperm. The embryo is cylindrical, straight curved,
or coiled, and macropodous.
POLLEN MORPHOLOGY. The pollen is shed in
monads, which are elongate-arcuate and inaper-
turate (Fig. 106G). The exine is reticulate, the
reticulum being thinned at both ends of the grain
and at the distal convex part (Erdtman 1952). The
curved shape of the grains is related to their ability
to float by means of holding air bubbles (Gamerro
1968).
KARYOLOGY. Reese (1962) reported somatic
chromosome numbers of 20 for Ruppia maritima
and 40 for R. spiralis. The karyotype comprises
nine small and one large chromosomes. Chromo-
some numbers based on x = 8 and 12 likewise
have been reported (Fedorov 1969).
PoLLINATION. Although it has been claimed that
pollination of Ruppia is underwater there are
many observations documenting that it takes
place on the surface of the water. The most de-
tailed observations on the process of pollination
are due to Gamerro (1968), who studied Ruppia
cirrhosa. Flowering begins with the opening of the
leaf sheath surrounding the developing inflores-
cence, followed by peduncle elongation. When the
peduncle lies horizontally just below the surface,
with a thin film of water surrounding it, pollen
release takes place. The opening of the anther
locule is indicated by the appearance of air
bubbles, which carry the abscised thecae upwards.
When reaching the surface, the thecae open
violently, spreading the pollen over the water
surface. The grains adhere end to end, forming
chains, and are oriented with the concave cavity
upward. Bubbles trapped in the cavity and exine
reticulation keep chains afloat.
Following pollen release, growth of the peduncle
resumes, and the inflorescence, now reduced to 2
apocarpous gynoecia, is brought to the water sur-
face. Pollen is carried to the stigmas by wave,
wind, or current. After fertilization is accom-
plished, the whole inflorescence is drawn beneath
the surface of the water by spiral contraction of
the peduncles.
FRUIT AND SEED. The endocarp of the fruit is
heavily sclerified, but perforated in the placental
447
regions by 2 ducts of pitted parenchyma, visible
externally as 2 depressions.
The seed coat consists of a thin, 2-layered testa
and a tegmen of which only the outer epidermis
persists, which is remarkable for the size of
its cellulosic cells filled with profusely ramified
and fused cellulosic corniform protuberances
(Gamerro 1968).
DISPERSAL. The fruits, after being released, float
on the water for a period of time varying from less
than a week to several months. They then sink and
become covered by the substrate. According to
Graves (1908), while covered by the substrate, the
exocarp and mesocarp decay, leaving the en-
docarp intact. The seed coat remains permanently
attached to the endocarp.
PHYTOCHEMISTRY. Saponins and alkaloids are
absent in the family. Les and Haynes (1995) report
leucoanthocyanins, flavonols, and 0-glycosyl fla-
vones in the family.
AFFINITIES. In most systems of classifica-
tion Ruppia is considered as closely related to
Potamogeton, and indeed similarities exist in the
structure of shoots and inflorescences, and
Poluszny and Sattler (1974) demonstrated even a
basic agreement in flower development, particu-
larly as to stamens and carpels. Differences with
Potamogeton exist, however, in the nuclear en-
dosperm development, the absence of a perianth,
the number of stamens, and the ontogeny of the
stamen appendage, which in Ruppia develops
as an outgrowth of the connective at a late stage
of floral development. These differences may
warrant family status for Ruppia distinct from
Potamogetonaceae. However, recent investiga-
tions using molecular data (Les et al. 1993; Les and
Haynes 1995) have failed to corroborate a close
relationship between Ruppia and Potamogetona-
ceae. These studies indicated that Ruppiaceae
are actually related to the marine families
Cymodoceaceae and Posidoniaceae but are not
closely allied to Potamogetonaceae.
The phenetic similarity of Cymodoceaceae,
Posidoniaceae, and Ruppiaceae (and their dis-
similarity from Potamogetonaceae) was earlier
recognized by Clifford (1970). Apart from mor-
phological features, it may be significant that
Ruppia is highly salt-tolerant and can withstand
salinities up to 70 000 ppm total dissolved salts
(Sainty and Jacobs 1994). Because Ruppia is found
more typically in brackish water than in marine
habitats, its relationship to other "seagrasses" has
448
not been given adequate consideration. Yet the
bisexual condition of floral organs in Ruppia,
together with a tolerance to a wide range of sa-
linities, may indicate a phylogenetic position
intermediate between a bisexual, freshwater an-
cestor and the Posidoniaceae (bisexual; exclu-
sively marine) and Cymodoceaceae (unisexual;
exclusively marine). All members of these three
families are water-pollinated and share adapta-
tions such as elongate pollen. Recalling past
comparisons with leaves of Potamogetonaceae,
similarity is also evident between leaves of Ruppia
and the filiform leaves of Syringodium (Cymo-
doceaceae).
DISTRIBUTION AND HABITATS. The Ruppiaceae
are subcosmopolitan. Most species occur in tem-
perate regions, especially in the northern hemi-
sphere. All species grow in fresh to brackish
waters, but none is marine. Those of freshwater
habitats often occur in waters with high concen-
trations of sulfur.
PALEOBOTANY. The earliest records of the family
are from the Paleocene (Muller 1981). Friis (1985
reports ?Limnocarpus sp. from the Middle Mi-
ocene in Denmark. She gives an illustration of
the endocarp, which resembles the endocarp of
Ruppia, and indicates that the specimen may rep-
resent either Potamogetonaceae or Ruppiaceae.
Daghlian (1981) makes no mention ofRuppiaceae.
A single genus:
Ruppia L.
Ruppia L., Sp. Pl. 1: 127 (1753).
Characters of the family. Seven or eight spp.; R.
maritima L. nearly cosmopolitan, R. cirrhosa
(Petagna) Grande in Europe and Nand S America;
three spp. endemic to Australia and New Zealand,
and one sp. to Central America/Caribbean. The
differences among the species are summarized in
Novelo (1991).
Selected Bibliography
Clifford, H.T. 1970. Monocotyledon classification with special
reference to the origin of grasses (Poaceae). In: Robson,
N.K.B., Cutler, D.F., Gregory, M. (eds.) New research in
plant anatomy. London: Academic Press pp. 25-34.
Daghlian, C.P. 1981. A review of the fossil record of monocoty-
ledons. Bot. Rev. (Lancaster) 47: 517-555.
Dahlgren, R.T.M., Clifford, H.T. 1982. See general references.
Ruppiaceae
Dahlgren, R., Rasmussen, F.N. 1983. Monocotyledon evolu-
tion. Evol. Biol. 16: 255-395.
Davis, G.L. 1966. See general references.
Erdtman, G. 1952. See general references.
Fedorov, A.A. 1969. See general references.
Friis, E.M. 1985. Angiosperm fruits and seeds from the Middle
Miocene of Jutland (Denmark). K. Dan. Vidensk. Selsk.
Biol. Skr. 24(3): 1-165.
Gamerro, J.C. 1968. Observaciones sobre Ia biologia floral y
morfologia de Ia potamogetonacea Ruppia cirrhosa (Petag.)
Grande(= "R. spiralis" L. ex Dum.). Darwiniana 14: 575-
608.
Graves, A.H. 1908. The morphology of Ruppia maritima.
Conn. Acad. Arts Sci. 14: 59-170.
Haynes, R.R. 1978. The Potamogetonaceae in the Southeastern
United States. J. Arnold Arbor. 59: 170-191.
Jacobs, S.W.L., Brock, M.A. 1982. A revision of the genus
Ruppia (Potamogetonaceae) in Australia. Aquat. Bot. 14:
325-337.
Les, D.H., Haynes, R.R. 1995. Systematics of subclass
Alismatidae: a synthesis of approaches. In: Rudall, P.J.,
Cribb, P.J., Cutler, D.F., Humphries, C.J. (eds.) Monocotyle-
dons: systematics and evolution. Royal Botanic Gardens,
Kew pp. 353-377.
Les, D.H., Garvin, D.K., Wimpee, C.F. 1993. Phylogenetic stud-
ies in the monocot subclass Alismatidae: evidence for a
reappraisal of the aquatic order Najadales. Mol. Phylog.
Evol. 2: 304-314.
Muller, J. 1981. Fossil pollen records of extant angiosperms.
Bot. Rev. (Lancaster) 47: 1-142.
Murbeck, S. 1902. Ober die Embryologie von Ruppia rostellata
Koch. Koninkl. Sven. Vetensk. Ak. Handl. 36(5): 1-21.
Novelo, R.A. 1991. Ruppia didymia (Potamogetonaceae) en
Mexico y las Antillas. An. Inst. Biol. Univ. Nac. Auton. Mex.
Ser. Bot. 62: 173-180.
Poluszny, U., Sattler, R. 1974. Floral development of Ruppia
maritima var. maritima. Can. J. Bot. 52: 1607-1612.
Preston, C.D. 1995. Pondweeds of Great Britain and Ireland.
BSBI Handbook no. 8. London: Botanical Society of the
British Isles.
Reese, G. 1962. Zur intragenerischen Gliederung der Gattung
Ruppia L. Z. Bot. 50: 237-264.
Fig. 106 Sainty, G.R., Jacobs, S.W.L. 1994. Waterplants in Australia.
Darlinghurst: Sainty and Associates.
Schwanitz, G. 1967. Untersuchungen zur postmeiotischen
Mikrosporogenese. I. Morphogenese des Ruppia-Pollens.
Pollen Spores 9: 9-48.
Takhtajan, A.L. 1982. See general references.
Tomlinson, P.B. 1982. Anatomy of the monocotyledons VII.
Helobiae (Alismatidae). Oxford: Clarendon Press.
 Scheuchzeriaceae 449

Scheuchzeriaceae lacunae in the mesophyll, root cortex, and

throughout the ground tissue of the axis. Hypo-
R.R. HAYNES, D.H. LES, dermal files of crystals line the vascular bundles of
and L.B. HOLM-NIELSEN the leaf. Long vessels with scalariform perforation
plates occur in the roots. Tannin cells are present
(Tomlinson 1982).

FLORAL MORPHOLOGY. Developmental studies

provide no indication of spiral inception of floral
Scheuchzeriaceae F. Rudolphi, Syst. Orb. Veg.: 28 (1830), nom. organs; instead, floral organs of each whorl arise
cons.

A glabrous rhizomatous perennial. Rhizome hori-

zontal, sympodial, covered with persistent fibrous
remains of older leaves. Adventitious roots in-
serted just below the node. Aerial stem erect, to ca.
35 em. Leaves basal and cauline, alternate, disti-
chous, linear, compressed-terete, several-veined,
with sheathing base, the apex obtuse with a
distinct pore; sheath open, marginally hyaline,
fibrous remains often persisting, with 2 delicate,
obtuse auricles; infravaginal hairs numerous.
Inflorescence a simple bracteate raceme with a
terminal flower. Flowers perfect, hypogynous,
trimerous and pentacyclic. Tepals distinct, all
similar, lanceolate-elliptic. Stamens distinct; an-
thers bithecate, 4-sporangiate, extrorse, dehiscing
by longitudinal slits, with short connective tip.
Carpels 3(-6), only basally connate, with decur-
rent, ventral, sessile, dry stigmas. Ovules (1- )2
per locule, basal-axile, erect, anatropous. Fruits
adaxially dehiscing follicles. Seeds 1(-2), oblong,
exalbuminous.
Only one species, Scheuchzeria palustris L., from
the arctic and temperate regions of the Northern
Hemisphere.

VEGETATIVE MORPHOLOGY. Scheuchzeria is a

rushlike, glabrous herb of Sphagnum bogs. The
branched roots are developed from the rhizome
just below the nodes. The blades are more or less
terete below, becoming flattened above, and taper
to a rounded apex, which is terminate adaxially by
a conspicuous pore. The sheath is wider than the
blade and has a hyaline margin that extends up-
wards as 2 small rounded auricles (Fig. 107). In the
axis of the sheaths are numerous hairs that sur-
round the nodes. These are obviously homologous
to the infravaginal squamules found in related
families.

VEGETATIVE ANATOMY. The sieve-tube plastids

contain cuneate protein bodies and lack starch or Fig. 107A-E. Scheuchzeriaceae. Scheuchzeria palustris. A
filamental protein. The anatomy is reduced in re- Flowering plant. B Fruiting plant. C Flower. D Fruit. E Seeds.
lation to the wet habitat, with well-developed air (Takhtajan 1982)
450 Scheuchzeriaceae

almost simultaneously in acropetal sequence AFFINITIES. The phylogenetic position of

(Posluszny in Tomlinson 1982: 233). Scheuchzeria is usually believed to be close to the
Juncaginaceae (including Lilaea). An expanded
EMBRYOLOGY. The walls of the uninucleate tapetal concept of Scheuchzeriaceae that include Lilaea
cells disintegrate and a periplasmodium is formed and other genera ofJ uncaginaceae was suggested
that intrudes between the pollen tetrads. Mi- by Buchenau (1903). The relationship between
crosporogenesis has not been studied (Davis 1966). Scheuchzeriaceae and Juncaginaceae is supported
The pollen is shed as dyads in the 3-celled stage. by the restricted occurrence of the cynogenic glu-
The ovules are anatropous and bitegmic, the mi- coside triglochinin in these families.
cropyle is formed by the inner integument, and a Characteristics of Scheuchzeriaceae that differ
parietal cell is formed and gives rise to 4 layers of from Juncaginaceae include pollen dyads, infra-
parietal tissue. The megaspore tetrad is linear. vaginal hairs, tetracytic stomata, follicular fruits, 2
Embryo sac formation follows the Polygonum ovules per carpel, and subterete leaves with a con-
type, and endosperm formation is Helobial. The spicuous apical pore. Because of these (and other)
micropylar chamber becomes coenocytic, but later distinctive features (Tomlinson 1982), the major-
cellular, while the nucleus of the chalaza! chamber ity of authors have recognized Scheuchzeriaceae
does not divide or exhibit any haustoria! activity as a monotypic family.
(Stenar 1935). Embryo development follows the Early classifications (e.g., Lindley 1853; Bessey
Caryophyllad type, in which the basal cell does 1915; Skottsberg 1940) placed Scheuchzeriaceae in
not divide but enlarges and forms the terminal cell the Alismatales. Hutchinson (1959) also followed
of the short suspensor. The ripe seeds are exal- this convention and considered the family to be
buminous. The embryo is chlorophyllous and has derived from the Alismataceae. Most contempo-
a small plumule and a single rounded cotyledon. rary authors (e.g., Cronquist 1981; Dahlgren et al.
1985) have placed the Scheuchzeriaceae in the
PoLLEN MORPHOLOGY. The pollen grains are dis- Zosterales ("Najadales").
persed in dyads (Erdtman 1952). They are globose, Molecular data (Les and Haynes 1995; D.H. Les,
inaperturate, and the exine is reticulate and unpubl.) do not support the merger of Scheu-
tenuimurate. The reticulum is continuous be- chzeriaceae and Juncaginaceae. Despite their
tween the 2 cells. shared occurrence of triglochinin, these families
are not phylogenetic sister groups, but are more
KARYOLOGY. The basic chromosome number is distantly related. The significant morphological
n = 11 (Love and Love 1961). and anatomical divergence of these families
(Tomlinson 1982) is paralleled by a considerable
PoLLINATION. This is probably by wind. level of molecular divergence. Morphological
and molecular data support the retention of
SEED. There is an obvious raphe arising at the Scheuchzeriaceae and depict a more distant
micropyle and running nearly the entire length relationship between these families than many
of the seed. The testa is glossy, smooth, and authors have assumed. Cladograms of the
unornamented. Alismatidae constructed from molecular data
clearly place the Scheuchzeriaceae among
PHYTOCHEMISTRY. Silica bodies, saponins, and families of the Zosterales, with no indication
alkaloids are absent in the family. The plants of a close relationship to families of the
are especially rich in the cyanogenic glucoside Alismatales. In these analyses. Scheuchzeriaceae
triglochinin. Flavonoids are unknown in the follow Aponogetonaceae as the most basal mem-
family (McClure 1970). The nonlignified cell bers of the Zosterales.
walls contain no ferculate.
Only one genus:
DISTRIBUTION AND HABITATS. Scheuchzeria
grows in temperate to subarctic regions in the Fig. 107
Scheuchzeria L.
northern hemisphere. In North America it ranges
in the south to N California and New Jersey, in Scheuchzeria L. Sp. Pl.: 338 (1753).
Europe to Italy and Jugoslavia, and in Asia to Japan.
It grows in transitional and ombrogenous Description as for family.
Sphagnum bogs, especially in bogs extending over
postglacial kettle lakes.
 Scheuchzeriaceae/Strelitziaceae
Selected Bibliography
Bessey, C.E. 1915. The phylogenetic taxonomy of flowering
plants. Ann. Mo. Bot. Gard. 2: 109-164.
Buchenau, F. 1903. Scheuchzeriaceae. Pflanzenreich IV. 14: 1-
20. Leipzig: W. Engelmann.
Cronquist, A. 1981. See general references.
Dahlgren, R.M.T. eta!. 1985. See general references.
Davis, G.L. 1966. See general references.
Erdtman, G. 1952. See general references.
Hutchinson, J. 1959. The families of flowering plants, Vol. 2,
2nd edn. Monocotyledons. Oxford: Clarendon Press.
Les, D.H., Haynes, R.R. 1995. Systematics of subclass
Alismatidae: a synthesis of approaches. In: Rudall, P.J.,
Cribb, P.J., Cutler, D.F., Humphries, C.J. (eds.) Monocotyle-
dons: systematics and evolution. Royal Botanic Gardens,
Kew pp. 353-377.
Lindley, J. 1853. The vegetable kingdom, 3rd edn. London:
Bradbury and Evans.
Love, A., Love, D. 1961. Chromosome numbers of central and
northwest European plant species. Opera Bot. 5: 1-581.
McClure, J.W. 1970. Secondary constituents of aquatic an-
giosperms, In: Harborne, J.B. (ed.) Phytochemical phylog-
eny. London: Academic Press pp. 233-268.
Skottsberg C.J.F. 1940. Vaxternes liv popular vetenskaplig
handbok Vol. 5. Stockholm: Aktiebolaget Familjeboken.
Stenar, H. 1935. Embryologische Beobachtungen iiber
Scheuchzeria palustris L. Bot. Not. 1935: 78-86.
Takhtajan, A.L. 1982. See general references.
Tomlinson, P.B. 1982. Anatomy of the monocotyledons VII.
Helobiae (Alismatidae). Oxford: Clarendon Press.
451
Strelitziaceae
L. ANDERSSON
Strelitziaceae (K. Schum.) Hutch., Fam. Fl. Pl. 2: 72 (1934),
nom, cons.
Musaceae subfam. Strelitzioideae K. Schum. (1900).
Arborescent suckering plants or acaulescent plants
with dichotomously branched stem, suckers axil-
lary when present. Leaves alternate, distichously
arranged, differentiated into sheath, petiole (often
indistinct or lacking), and blade; sheath without
distinct ligule; blade entire, with coarse midrib and
closely set, parallel, slightly sigmoid lateral veins
fusing near margin; lateral veins interconnected by
tertiary cross-veins. Inflorescence a terminal or lat-
eral thyrse, bearing cincinnate flower clusters in
axils of spathaceous bracts on an indeterminate
main axis. Flowers perfect, trimerous and 5-cyclic,
heterochlamydeous, zygomorphic, subtended by
carinate bracteoles; stamens 5 or 6, no staminodes
present; anthers basifixed, elongate, 2-thecate, 4-
sporangiate. Ovary inferior, 3-locular, with deeply
sunken septal nectaries; ovules anatropous, nu-
merous on axile placentae; style filiform, often
dilated distally. Fruit a dehiscent capsule; seeds
inoperculate, arillate, with a 1-layered, starchless
perisperm, with copious starchy endosperm.
Three genera and six or seven species; tropical
America, southern Africa and Madagascar.
VEGETATIVE MORPHOLOGY. Various aspects of
vegetative morphology have been described by
Tomlinson (1962).
The plants generally have a sympodial rhizome
with branches terminating in aerial shoots, and
with renewal branches arising near the base of the
aerial shoot. Vegetative buds are axillary (Fisher
1978). Dichotomous branching has been demon-
strated in Strelitzia reginae (Fisher 1976). Rhi-
zome branches are usually short and massive,
becoming cormlike at soil surface. Leaves are pro-
duced from the "corm" tip. The short rhizome
branches typically give the plants a caespitose
habit. Phenakospermum has rhizome branches
developed as runners. Starch is commonly stored
in the ground parenchyma of the rhizome
(Tomlinson 1959), but no specialised storage
structures are known.
452 Strelitziaceae
Fig. 108A-D. Strelitziaceae. Ravena/a madagascariensis.
A Flowering plant. B Partial inflorescence, the spathaceous
bract half cut away, floral bracts removed, showing one open
flower after anthesis and a second mature, but unopened
flower. C Ovary, transversal section. D Stigma with fingerlike
protuberances surrounding the receptive area. (Kress et a!.
1994)
Ravenala, Phenakospermum and Strelitzia nico-
lai have an elongate, woody aerial stem, which
bears the leaves in a fanlike cluster on top
(Fig 108). In other species of Strelitzia the stem is
short and cormlike. Even though the stems are
woody in texture, no secondary growth has been
demonstrated.
The leaf is usually differentiated into sheath,
petiole and blade, although a distinct petiole is
sometime missing. All divisions of the leaf are
derived from the proximal (Unterblatt) part of the
primordium, the distal (Oberblatt) part giving rise
only to a precursory tip, which is usually not iden-
tifiable in the mature leaf (Kaplan 1973).
The ptyxis is supervolute. In Phenakospermum
the mature leaf appears torn off at tip (although
the shedding of the leaf tip does not seem to have
been observed). The leaf blade in all genera has a
massive midrib from which closely set, parallel
secondary veins diverge. These have a slightly
sigmoid course and fuse near the leaf margin.
They are interconnected by parallel tertiary veins,
which are perpendicular to the secondary ones.
VEGETATIVE ANATOMY. Vegetative anatomy was
described in detail by Tomlinson (1959, 1960). The
stomatal apparatus was described by Tomlinson
(1974) and Olatunji (1980).
Root stele with wide vessels and phloem strands
scattered throughout the stelar ground tissue. Rhi-
zomes with the central cylinder demarcated by
an endodermis. Hairs absent from aerial parts.
Arborescent stem with a peripheral mechanical
layer with congested vascular bundles with mas-
sive fibre sheaths; central cylinder not demarcated
by an endodermis. Leaf axis with a main arc of
vascular strands which are continuous with the
major lateral veins of the lamina, several arcs of
smaller and shorter strands, some entirely fibrous,
present on the abaxial side of the main arc; with
one separate system of air canals on each side of
the main arc of vascular bundles, each system
often with several arcs of air canals. Lamina: epi-
dermal cells with anticlinal walls variable but pre-
dominantly non-sinous; stomata polycytic, guard
cells symmetrical in cross-section; hypodermis
2- or more-layered; mesophyll homogeneous and
composed of compact or lobed cells; vascular
strands with a sheath of fibres on either side, all
veins connected to both sides. Epicuticular wax
rodlets (Strelitzia wax) present (Frolich and
Barthlott 1988). Silica cells associated with vascu-
lar bundles in all parts of the plant except the
roots, silica bodies druselike or spherical. Raphide
sacs present in the ground tissue of all organs.
Tannin present, in morphologically unspecialised
cells in all parts of the plant.
INFLORESCENCE. The inflorescence is lateral in
Ravenala and Strelitzia, but terminal in Phenako-
spermum. The main axis is indeterminate. It
bears distichous, persistent, spathaceous bracts
(spathes), in the axils of which partial inflores-
cences arise. The flowers are bracteolate.
The partial inflorescences have been variously
interpreted. The classical (and still most com-
monly accepted) interpretation, proposed by
 Strelitziaceae
Eichler (1875), is that it is a cincinnus. The whole
inflorescence is then a thyrse with monochasial
partial inflorescences (see also under Heli-
coniaceae ).
FLORAL STRUCTURE AND ANATOMY. The flowers
have a basically trimerous, pentacyclic organiza-
tion, with 2 whorls of tepals, 2 of stamens, and 1 of
carpels. The flowers are oriented with the un-
paired sepal in the posterior position. The outer
tepals are free and equal or subequal, whereas the
inner are basally fused for a short distance, the 2
anterior being distinctly larger than the posterior
inner tepal. In Strelitzia the 2 anterior inner tepals
are connivent, to form a sagittate structure, which
centrally forms a groove enclosing filaments and
style. The sagittate structure is brightly coloured
by special carotenoids and obviously has an ad-
vertising function.
Ravenala has a complete set of 6 fertile stamens,
whereas the other genera have 5. The missing
stamen belongs to the inner whorl. It is not repre-
sented by a staminode.
The gynoecium is formed by 3 carpels. The
ovary is inferior and trilocular, with numerous
ovules on axile placentae. It has a conspicuous
solid prolongation above the locules. The septal
nectaries are deeply sunken, reaching as far down
as the ovarian transmitting tissue (Kronestedt and
Walles 1986).
EMBRYOLOGY. The anthers are tetrasporangiate.
The tapetum of Strelitzia is secretory and multi-
layered. Microspore formation is successive and
tetrads are isobilateral, linear or T-shaped, in
approximately equal proportions. Pollen grains
are 1-celled at dispersal due to disintegration of
the vegetative cell (van de Venter 1976a).
The ovule is anatropous, with 2 integuments.
The micropyle is formed by both integuments.
The nucellus is crassinucellate. Megasporogenesis
follows the Polygonum type. No parietal cell
seems to be formed. A circular thickening, the so-
called Mikropylarkragen, is formed by periclinal
divisions in the outer integument. The nucellar
epidermis becomes palisadelike at the maturity of
the embryo sac. The aril is formed partly from the
outer integument near the micropyle, partly from
the funicle opposite the micropyle. Endosperm
formation is Nuclear (Mauritzon 1936).
The mature embryo is club-shaped in Strelitzia
(van de Venter 1976b).
POLLEN MORPHOLOGY. Pollen grains are radio-
symmetric and isopolar, without a structurally
453
distinct aperture. The exine is endexinous in na-
ture, very thin and in Phenakospermum discon-
tinuous. It is made up of several layers which may
(Phenakospermum) slough off during develop-
ment. The intine is thick and 2-layered. The
exintine is granular and the endintine is chan-
nelled, appearing radially striate in the light
microscope. In Strelitzia reginae grains are con-
nected together within a pollen sac by cellular,
non-sporopolleninous threads that are not con-
nected with the exine (Hesse and Waha 1983).
KARYOLOGY. Chromosome numbers of 2n = 14
and 2n = 22 have been reported for Strelitzia and
2n = 22 for Ravenala (Bisson et al. 1968). Other
numbers for these genera seem unreliable. One
pair of chromosomes has satellites (Cheesman
and Larter 1935).
PoLLINATION. Early observations of bird pollina-
tion in Ravenala (Scott Elliot 1890) have not been
confirmed. Kress et al. (1994) instead gave a de-
tailed report on the pollinator relationship with
the ruffed lemur. The specialisations of Ravenala
for visitation by large non-flying animals include
the easy access of the inflorescences to arboreal
animals; the large size of the flowers enclosed in
tough, protective bracts requiring a strong polli-
nator to open; and the production of copious,
sucrose-rich nectar. In contrast, Strelitzia nicolai
and S. reginae are pollinated by sunbirds (Frost
and Frost 1981). The birds perch on the anther
sheath and stigma, and thus transfer pollen pri-
marily with their feet. Nectar-feeding bats of the
Phyllostomatidae appear to be the primary polli-
nators of Phenakospermum in South America
(W.J. Kress and D. Stone, unpubl.).
FRUIT AND SEED. The fruit is a woody, trilocular,
loculicidal capsule, opening at maturity to expose
the seeds and their brightly coloured arils. The aril
is made up of threads and orange in Strelitzia,
blue in Ravenala, and red in Phenakospermum.
Seed anatomy of Strelitzia and Ravenala was de-
scribed by Desler (1994). The mature seeds lack
hypostase and operculum. The testa has a con-
spicuous outer cuticle. It is made up almost
entirely by tissues derived from the outer integu-
ment; the inner integument is represented only by
compressed remainders and a cuticle. The inner
epidermis of the outer integument consists of
large cells with the inner tangential walls con-
spicuously thickened (U-celllayer). The nucellus
is represented by a single layer of cells, which
may store lipids but no starch. Starch is stored in
454
the endosperm in the form of simple, fusiform
grains.
DISPERSAL. The seeds are obviously dispersed by
birds, which are attracted by the brightly coloured
arils.
RELATIONSHIPS. Strelitzia, Ravenala, and Phena-
kospermum have traditionally been treated as a
tribe of the Musaceae (e.g. Schumann 1900;
Winkler 1930). Hutchinson (1934) placed them
together with Heliconia in a separate family, the
Strelitziaceae. Nakai (1941), finally, segregated
Heliconia, which he referred to a separate family.
Nakai's view has subsequently become predomi-
nant (e.g. Tomlinson 1962; Cronquist 1981;
Dahlgren et al. 1985) and is also supported by
cladistic analyses (Kress 1995).
In a phylogenetic analysis of the Zingiberales
based on morphological characters (Kress 1990)
the Strelitziaceae are basal to the other members
of the "banana group" (excepting Musaceae), in
which the four "ginger families" are nested.
DISTRIBUTION. The Strelitziaceae have a relictual
distribution with Ravenala endemic to Madagas-
car, Phenakospermum widely distributed in north-
ern S America, and Strelitzia distributed along the
SE coast of S Africa.
KEY TO THE GENERA
1. Petals very unequal: the 2 paired (anterior) ones connivent
into a large sagittate structure with central groove enclosing
stamens and style; the posterior petal small and scalelike
- Petals ± equal in shape but the posterior smaller
2. Inflorescence lateral; stamens 6
- Inflorescence terminal; stamens 5 2. Phenakospermum
1. Ravenala Adanson
Ravena/a Adanson, Fam. Pl. 2: 67 (1763), Lane, Mitt. Bot.
Staatssamml. Miinch. 13: 114-131 (1955), rev.
Urania Schreb. (1893).
Ravena/a subgen. Urania (Schreb.) Baker (1893).
Arborescent, suckering plants, with woody trunk
apically with leaves in fanlike cluster; leaf tips
acuminate. Inflorescence axillary. Outer tepals
equal, free; inner equal in shape but posterior one
smaller than the lateral; stamens 6; each locule
with 2 rows of ovules. Seeds depressed; aril blue.
Only one sp., R. madagascariensis Sonn., endemic
to Madagascar, now widely cultivated as orna-
mental in the tropics.
Strelitziaceae
2. Phenakospermum Endl.
Phenakospermum End!., Gen. Pl. 3: 229 (1837).
Ravena/a subgen. Phenakospermum (End!.) Baker (1893).
Arborescent, suckering plants; leaves in fanlike
cluster on top of trunk; leaf tip truncate due to
pseudocalyptrate bud opening. Inflorescence ter-
minal. Outer tepals equal, inner shorter, fused at
base, subequal in shape, but posterior one smaller
than lateral; stamens 5, staminode lacking; each
locule with 4-many rows of ovules. Seed elongated
along micropyle-chalaza axis; aril bright red. Only
one sp., P. guyannense (Rich.) Endl. ex Miq.,
widely but patchily distributed in tropical S
America.
3. Strelitzia Ait.
Strelitzia Ait., Hort. Kew. 1: 285 (1789).
Plant either arborescent and suckering or acaules-
cent with dichotomously branching, cormlike
stems. Inflorescence lateral. Outer tepals unequal,
the median smaller than the lateral paired ones;
inner tepals strongly unequal, the paired ones
large, brightly coloured and connivent into a sag-
ittate structure centrally with a groove enclosing
style and stamens, the unpaired (posterior) small;
stamens 5, staminode lacking; locules with 2 rows
of ovules. Aril orange. Four or five spp., mainly
along the East coast of S Africa, from the Cape
Province into Transvaal. S. regineae Ait. and S.
nicolai Regel & Koch widely cultivated as orna-
mentals, the former also grown on a large scale for
3. Strelitzia cut flowers (Bird of Paradise).
2
1. Ravena/a
Selected Bibliography
Fig. 108
Bisson, S., Guillemet, S., Hamel, J.-L. 1968. Contribution a
!'etude caryo-taxonomique des Scitaminees. Mem. Mus.
Hist. Nat. II. B. 18: 59-145, pl. 1-19.
Chessman, E.E., Larter, L.N.H. 1935. Genetical and cytological
studies of Musa. III. Chromosome numbers in the Musa-
ceae. J. Genet. 30: 31-52.
Cronquist, A. 1981. See general references.
Dahlgren, R. et a!. 1985. See general references.
Desler, B. 1994. Vergleichende Untersuchungen der Anatomie
und Morphologie von Scitamineen-Samen. Unpublished
Diploma Thesis. Kaiserslautern: Faculty of Biology, Univer-
sity of Kaiserslautern.
Eichler, A.W. 1875. Bliithendiagramme construiert and
erHiutert Vol. 1. Leipzig: Engelmann.
Fisher, J.B. 1976. Development of dichotomous branching and
axillary buds in Strelitzia (Monocotyledonae). Can. J. Bot.
54: 578-592.
 Strelitziaceae/Thurniaceae
Fisher, J.B. 1978. Leaf-opposed buds in Musa: their develop-
ment and a comparison with allied monocotyledons. Am. J.
Bot. 65: 784-791.
Frolich, D., Barthlott, W. 1988. See general references.
Frost, S.K., Frost, P.G.H.1981. Sunbird pollination ofStrelitzia
nicolai. Oecologia 49: 379-384.
Hesse, M., Waha, M. 1983. The fine structure of the pollen wall
in Strelitzia reginae (Musaceae). Plant. Syst. Evol. 141: 285-
298.
Hutchinson, J. 1934. Families of flowering plants, 2nd edn.,
Vol. 2. Oxford: Clarendon Press.
Kaplan, D.R. 1973. The problem of leaf morphology and
evolution in monocotyledons. Q. Rev. Bioi. 48: 437-457.
Kress, W.J. 1990. The phylogeny and classification of the
Zingiberales. Ann. Mo. Bot. Gard. 77: 698-721.
Kress, W.J., Schatz, G.E., Andrianifahanana, M., Morland, H.S.
1994. Pollination of Ravena/a madagascariensis (Strelitzia-
ceae) by lemurs in Madagascar: evidence for an archaic
coevolutionary system? Am. J. Bot. 81: 542-551.
Kronestedt, E., Walles, B. 1986. Anatomy of the Strelitzia
reginae flower (Strelitziaceae). Nord. J. Bot. 6: 307-320.
Mauritzon, J. 1936. Samenbau und Embryologie einiger
Scitamineen. Lunds Univ. Arsskr. N.F. Adv. 2, 31: 1-31.
Nakai, T. 1941. Notulae ad plantas orientalis (XVI). Jpn. J. Bot.
17: 189-203.
Olatunji, O.A. 1980. The structure and development of sto-
mata in some Zingiberales. Notes R. Bot. Gard. Edinb. 38:
499-515.
Schumann, K. 1900. Musaceae. In: Engler A. (ed.) Das
Pflanzenreich IV.46: 1-45. Leipzig: Engelmann.
Scott-Elliot, G.F. 1980. Note on the fertilization of Musa,
Strelitzia reginae, and Ravena/a madagascariensis. Ann.
Bot. 4: 259-264.
Tomlinson, P.B. 1959. An anatomical approach to the classifi-
cation of the Musaceae. J. Linn. Soc. Bot. 55: 779-809.
Tomlinson, P.B. 1960. The anatomy of Phenakospermum
(Musaceae). J. Arnold Arbor. 41: 287-297.
Tomlinson, P.B. 1962. Phylogeny of the Scitamineae - mor-
phological and anatomical considerations. Evolution 16:
192-213.
Tomlinson, P.B. 1974. Development of the stomatal complex
as a taxonomic character in the monocotyledons. Taxon 23:
109-128.
Venter, H.A. Van de. 1976a. Microsporogenesis and -gameto-
genesis in Strelitzia reginae Ait. J.S. Afr. Bot. 42: 25-31.
Venter, H.A. Van de 1976b. Notes on the morphology of the
embryo and seedling of Strelitzia reginae Ait. J.S. Afr. Bot.
42: 63-69.
Winkler, H. 1930. Musaceae. In: Engler, A., Prantl, K. (eds.)
Die natiirlichen Pflanzenfamilien, 2nd edn. 1Sa. pp. 505-
541. Leipzig: W. Engelmann.
455
Thurniaceae
K. KUBITZKI
Thurniaceae Engl., Syllabus ed. 5: 94 (1907}, nom. cons.
Coarse perennial herbs from an upright roostock.
Leaves tristichously arranged, simple, parallel-
veined, all basal, with sheathing base and elongate,
leathery, flat or (in cross-section) V-shaped,
sometimes prickly margined blade. Inflorescences
of one or more dense, racemose, globose to ellip-
soid heads subtended by spreading, leafy bracts
on an otherwise leafless, bluntly triangular or
quadrangular peduncle; flowers perfect, regular;
pedicel short, swollen, spreadingly puberulous,
grading into the perigone; tepals persistent,
chaffy, 3 + 3 equal, spotted; stamens distinct, 3 +
3, with the filaments somewhat adnate to the base
of the tepals; anthers basifixed, tetrasporangiate,
introrse, opening by longitudinal slits; pollen
borne in tetrads, scabrate and ulcerate; gynoecium
of 3 carpels united to form a compound, superior,
trilocular pistil with 3 or more erect, anatropous,
crassinucellar, ascending ovules; ovary tapering
distally, scarcely produced into a style, with 3
elongated stylodia, stigmas dry. Fruit a loculicidal
capsule, commonly with 3 spindle-shaped seeds
supplied at each end with a hispid process, 1 in
each locule; embryo small, cylindric, monocotyle-
donous, embedded in copious, mealy, starchy
endosperm.
Comprising a single genus with three species
centred in the Guayana region of northern S
America, extending into the Amazon region;
growing in flooded habitats on sandy soil.
ANATOMY (Cutler 1965, 1969). On the leaves, the
stomata are confined to intercostal zones and are
paracytic, rarely tetracytic. A 1-2-layered hypo-
dermis is developed beneath both leaf surfaces
partly as sclerenchyma, partly as parenchyma,
with cells of the latter sometimes protruding be-
tween the epidermal cells. The leaf blades are pro-
vided with numerous air canals which at intervals
are traversed by diaphragms. The leaf vascular
bundles are arranged in vertical pairs composed
of a larger adaxial and a smaller abaxial bundle.
The latter is inverted so that the phloem portions
of both bundles face each other, a trait considered
456 Thurniaceae
as very peculiar by Cutler; it is different from an
arrangement to be expected in unifacialleaves.
Vessels with slanting end walls and scalariform
perforation are present in all vegetative parts.
Silica is present in sandlike particles in paren-
chyma cells and 1-several small, spheroidal or
nodular silica bodies per cell in some epidermal
cells. Tannin cells are scattered; crystals of
calcium oxalate are missing.
EMBRYOLOGY AND SEED (from Thimm 1985).
The division of the pollen mother cell is simul-
taneous. Mature pollen is 3-celled. The ovule is
bitegmic, crassinucellar and has a zig-zag micro-
pyle. In the seed coat, the outer integument is
composed of 1-3 layers of sclerenchymatic fibres,
which extend into the terminal processes of the
seeds, and 1-2 inner parenchymatic layers. The
tegmen is represented by 2 cell layers densely filled
with tannin, the outer small and compressed, the
inner large.
Stevenson and Loconte (1995) (confirmed by
D.W. Stevenson, pers. comm.) reported for
Thurnia the presence of an aril and diffuse
centromeres.
AFFINITIES. At one time, the Thurniaceae had
been included in the Juncaceae, with which they
agree in several traits, most notably the tetrad pol-
len (Fig. 109), but from which they differ in the
possession of silica bodies which are absent from
the latter family but occur - in different shape - in
Fig. 109. Thurniaceae. Thurnia polycephala. Pollen tetrad
with finely areolate, microspinulose exine sculpture, SEM
X2400. (Photo E. Grafstrom, Palynol. Laboratory, Stockholm)
I
P·
.' ~
A !
.,lj
c D
Fig.110A-D. Thurniaceae. A, B Thurnia jenmanii. A Capitate
inflorescence. B Flower. C, D Th. sphaerocephala. C Capsule.
D Seed. (Takhtajan 1982)
the Cyperaceae. Thus it would appear attractive
to view Thurniaceae as linking Juncaceae with
Cyperaceae (all three with diffuse centromeres!),
as suggested, among others, by Dahlgren et al.
(1985) and Huber (1991). In a cladistic analysis of
the Cyperales, which was based on structural data,
Simpson (1995) found Thurnia embedded in the
Juncaceae. It must be borne in mind, however,
that the little, that up to the present is known of
Thurnia, makes it distinct from the Juncaceae.
Only one genus:
Thurnia Hook. f. Fig. 109, 110
Thurnia Hook. f. in Hook., Ic. Pl.: t. 1407-1408 (1883).
Decription as for family.
Selected Bibliography
Cheadle, V.I., Kosakai, H. 1974. Vessels in Juncales. I.
Juncaceae and Thurniaceae. Phytomorphology 23: 80- 87.
Cutler, D.P. 1965. Vegetative anatomy of Thurniaceae. Kew
Bull. 19: 431-441.
Cutler, D.P. 1969. Juncales. In: Metcalfe, C.R. (ed.) Anatomy of
the monocotyledons, IV. Oxford: Clarendon Press, pp. 78-
86.
Dahlgren, R. et al. 1985. See general references.
Huber, H. 1991. See general references.
Pilger, R. 1930. Thurniaceae. In: E.-P., Natiirliche Pflanzen-
familien, 2nd edn., vol. 15a. Leipzig: W. Engelmann, pp. 58-
59.
Simpson, D. 1995. Relationships within Cyperales. In: Rudall,
P.J., Cribb, D.F., Cutler, D.P., Humphries, C.J. (eds.) Mono-
 Thurniaceae/Typhaceae 457

cotyledons; systematics and evolution. Royal Botanic Typhaceae

Gardens, Kew, pp. 497-509.
Stevenson, D.W., Loconte, H. 1995. Cladistic analysis of K. KUBITZKI
monocot families. In: Rudall, P.J., Cribb, D.F., Cutler, D.F.,
Humphries, C.J. (eds.) Monocotyledons: systematics and
evolution. Royal Botanic Gardens, Kew, pp. 543-578.
Takhtajan, A.L. 1982. See general references.
Thimm, U. 1985. Zur Embryologie, Bliiten- und Fruch-
tanatomie der isolierten Juncales-Gattungen Prionia and
Thurnia. Diploma Thesis. Bochum: University of Bochum,
Division of Biology.
Typhaceae Juss., Gen. Pl.: 25 (1789), nom. cons.
Sparganiaceae F. Rudolphi (1830).

Glabrous, perennial, herbaceous amphiphytes

from scaly, sympodial, starchy rhizomes, emergent
from shallow water or growing on wet soil, some-
times wholly submerse and floating. Leaves alter-
nate, distichous, with strongly sheathing base and
an elongate, parallel-veined blade, basal in sterile
plants and forming an erect false stem, on
flowering plants also dispersed along the stem,
rarely stem bearing only hypsophylls. Inflorescence
racemose, the numerous monoecious, hypogynous
flowers grouped into dense, complex globular
heads or very dense cylindric spikes; perianth of
1-several inconspicuous tepals or more or less
numerous slender bristles; stamens (1-)3(-8), the
filaments distinct or connate; anthers basifixed,
tetrasporangiate, dithecal; gynoecium pseudo-
monomerous, with a single fertile carpel, less often
with 2-3 ± fully developed carpels connate to form
a compound ovary, the stigmas or branches of the
common style accordingly 1-3; ovule solitary (in
each locule), pendulous from near the top oflocule,
apotropous, anatropous, bitegmic, crassinucellar.
Fruit small, dry, indehiscent or dorsally dehiscent;
seed with a straight embryo that has a single termi-
nal cotyledon and a lateral plumule; embryo sur-
rounded by copious, mealy endosperm and thin
perisperm, the endosperm containing starch, pro-
tein, and oil. x = 15.
Two genera with 16-30 spp. of wide distribution
in the Northern (and one genus also Southern)
Hemisphere.

VEGETATIVE MORPHOLOGY. The sympodial rhi-

zomes, which bend upward to form sterile leaf
clusters or upright stems, produce 1-numerous
axillary buds continuing the shoot system. Vari-
ous shoot generations can be produced every year,
although usually only the first shoot generation
comes to flower, while the subsequent ones pro-
duce sterile leaf clusters or finally overwintering
buds.
Some Sparganium spp. are floating-leaved and
have flaccid, flat, or weakly keeled leaves of
458 Typhaceae

dorsiventral internal structure. Other typically

emergent species produce robust, stiff, strongly
keeled, but not dorsiventral leaves. Kaul (1972,
1976) suggested that the floating-leaved species
are neotenous forms derived from emergent
species, and that Sparganium is a genus in
transition from the emergent to the floating habit;
Cook and Nicholls (1986) are of the opposite
opinion.

VEGETATIVE ANATOMY. The leaves in both gen-

era possess a prominent system of aerenchy-
matous channels. These are compartmentalized
by diaphragms which are 1-3 cell layers thick,
interspersed with large intercellular spaces.
Cross-sections of the leaves are diagnostic for
most species. Stomata are paracytic. The leaf epi-
dermis contains myriophyllin cells filled with tan-
nin and, on the adaxial side of the leaf sheaths of
Typha, mucilage cells. Raphide bundles are com-
mon, and oxalate styloids are known from Typha.
In the rhizomes of both genera, starch is stored in
large quantities. For further details see Solereder
and Meyer (1933). Typha has epicuticular wax
crystals of the Strelitzia type (Frolich and Barthlott
1988).

INFLORESCENCE AND FLORAL MORPHOLOGY.

Following earlier suggestions (i.a. Celakovsky
1885, 1891), it is now generally accepted that the
carpellate inflorescences of Sparganium and
Typha are highly contracted branching systems.
Developmental studies have provided evidence
for the compound nature of the inflorescences in
both genera. While the capitula of Sparganium are
di- or even tricephalia (Miiller-Doblies 1969), the
flowers in the carpellate spikes Typha are believed
to originate on axes of at least the 4th degree
(Miiller-Doblies 1969). The stigmas have a Dry
surface, which is papillate in Sparganium, non-
papillate in Typha. For details of the bewildering
variation in the flowers of both genera and the Fig. lllA-F. Typhaceae. A, B Sparganium erectum. A Habit,
homology of the hairs surrounding staminate and the stamina! heads distal to the pistillate. B Fruit. C-F S.
emersum. C Habit. D Staminate flower. E Pistillate flower.
carpellate flowers of Typha, see Miiller-Doblies F Ovary, longitudinal section to show pendulous ovule.
(1969, 1970). (Takhtajan 1982}

EMBRYOLOGY. Microsporogenesis is successive

and leads to isobilateral, decussate, linear, and
T-shaped tetrads. Both genera agree in having
apotropous and anatropous, crassinucellate
ovules with a primary tapetal cell, an embryo
sac developing according to the Polygonum type,
Helobial endosperm development, and (needing
confirmation in Typha) a secretory tapetum with 8
nuclei per cell (Miiller-Doblies 1969, 1970).
POLLEN MORPHOLOGY. Pollen grains are spheroi-
dal monads or tetrads (only Typha) with a distal
ulcus. Tetrads have evolved various times inde-
pendently in Typha (Punt 1975). As Nicholls and
Cook (1986) have demonstrated, in Typha, the
dense aggregation of pistillate flowers permits
every pollen grain in a pollen tetrad to pollinate
1 pistil (Fig. 113).
 Typhaceae
Fig. 112A-J. Typhaceae. A-H Typha latifolia. A Fruiting
shoot. B Rhizome and sheathing leaf bases. C Three staminate
flowers, the three filaments of each flower connate. D Cluster
of four carpellate flowers. E Three carpellate flowers, many
trichomes omitted to show stipitate ovaries. F Infructescence
shedding fruits. G Mature fruit with numerous hairs on the
stipe, the style persistent. H Ovary in longitudinal section to
show pendulous ovule. I, J T. angustifolia. I Inflorescence,
showing gap between distal staminate and lower carpellate
portions. J Portion of carpellate inflorescence to show three
flowers, simple and glandular trichomes, and a spathulate
sterile flower. (Thieret and Luken 1996, Generic Flora of
the southeastern United States)
KARYOLOGY. Nearly invariably, the known
counts in both genera are 2n = 30, with rare and
questionable instances of tri- or polyploidy.
REPRODUCTION. Both genera are strictly pro-
togynous. Ample hybridization is known from
Typha and has been studied in European
(Krattinger 1978} and N American (Smith 1987)
populations.
459
FRUIT, SEED, AND DISPERSAL. The ripe fruits of
Sparganium drop into water, where they can float
for at least a year. Eventually, the spongy exocarp
may rot off, leaving only the endocarp. Dispersal is
mainly hydrochorous, supplemented by epi- and
endozoochory (by water birds). At the distal end
of the endocarp there is a pore through which the
embryo emerges in germination. The inner open-
ing of the pore is blocked by a micropylar plug
formed by an enlargement of the integuments.
Germination can be delayed for several years after
maturity, apparently by the retaining influence of
the micropylar plug. After the decay or removal of
the plug, and given sufficient moisture, germina-
tion readily occurs.
Typha is well adapted to anemochory by its
light-weight fruits (0.02-0.003 mg, Finlayson et al.
1983} and perigone scales transformed into hairs
in order to facilitate dispersal by the wind. Conse-
quently, the rates of seed production are enor-
mous. A single carpellate spike of T. latifolia can
produce an estimated 220 000 seeds; one of T.
dominguensis 682 000 (Yeo 1964; Finlayson et al.
1983).
The differences between the two genera are
clearly related to the switch from hydrochory in
Sparganium to anemochory in Typha. Thus, the
persistent perigone has been dissolved into fine,
uniform hairs without vascular bundles; starch in
the endosperm has partially been replaced by fatty
oil to diminish the weight of the diaspores; the
mesocarp and endocarp has been strongly re-
duced, and the inflorescences strongly condensed.
PHYTOCHEMISTRY. Common cinnamic acids
and flavonoids including proanthocyanidin
were found in both genera, including in Spar-
ganium prodelphinidin and myricetin, the latter
compound being relatively rare in monocots
(Hegnauer 1963, 1986}. The unlignified cell walls
are fluorescent under UV and contain bound
ferulate (Harris and Hartley 1980).
AFFINITIES. Evidence provided by Miiller-
Doblies (1969, 1970} make out a strong case for
combining Sparganium and Typha in a single
family. The former association of this family with
the Pandanaceae is not confirmed, however. Dif-
ferent molecular (Duvall et al. 1993; Chase et al.
1995; Davis 1995) and morphological/molecular
(Linder and Kellog 1995) evidence (including the
possession of Strelitzia wax) consistently place the
Typhaceae in the "commelinoid clade". A close
relationship to Poales seems at present the best-
supported hypothesis (Linder and Kellogg 1995}.
460
A B
Fig. 113A-C. Typhaceae. Typha latifolia. A Pollen tetrad, two
grains germinating (X 1200). B Pollen tubes growing over and
between stigmas, the tubes of one tetrad often reaching differ-
ent stigmas ( X60). C Pollen tubes crossing from pollinated
stigma to unpollinated stigma ( X30). (SEM photo Nicholls
and Cook)
PALAEOBOTANY. The fossil record ofboth genera
dates back into the Upper Cretaceous (Herendeen
and Crane 1995), although this is questioned by
Cook and Nicholls (1987). Fruits of Sparganium
from the Paleocene have 5-locular fruits, and
during the Tertiary and Pleistocene a gradual
reduction to the presently predominant 1-locular
condition can be observed. In extant 2-3-locular
fruits, which occasionally occur, generally only 1
locule contains a functional ovule.
EcoLOGY AND UsEs. Typha is important in its
role as a member of the reed belt, which in our
time is continually gaining in importance as con-
tributing to the biological cleaning of polluted
waters. The stems of Typha and Sparganium pro-
vide shelter and a substratum for developing
arthropods. Virtually every part of Typha is edible
for some animals and the young staminate spikes,
the rhizomes, and even pollen of Typha (which
can be gathered in major quantities) have been
used for human consumption. The fruits of Spar-
ganium are important food for water birds, and
the fruits, stems, and leaves are eaten by mammals
such as muskrat and moose.
KEY TO THE GENERA
l. Partial inflorescences headlike; drupe with a persistent
perigone and airfilled mesocarp 1. Sparganium
- Inflorescence spikelike, erect, with a lower pistillate portion
and several upper staminate portions; fruits with a tuft of
perigone hairs, finally dehiscent 2. Typha
Typhaceae
c
1. Sparganium L. Fig. 111
Sparganium L., Sp. Pl.: 971 (1753); Cook & Nicholls, Bot. Helv.
96: 213-267 (1986), and 97: 1- 44 (1987), monogr.
Marsh or aquatic perennials with sheathing leaves,
simple or branched stems, flowers in dense, glo-
bose heads on the upper part of the stem or on the
branches, the staminate heads distal to the carpel-
late; stamens 1-8; tepals 1-6; ovary 1-locular or
very rarely 2-3-locular; fruit with spongy meso-
carp and hard endo carp with a distinct micropylar
plug; seeds with membranous testa and starchy
endosperm. Fourteen spp., temperate and arctic
regions of Northern Hemisphere, mountains of
Sumatra and New Guinea, and SE Australia and
New Zealand.
2. Typha L. Fig. 112, 113
Typha L., Sp. Pl.: 971 (1 753); Kronfeld, Verh. Zoo!.-Bot. Ges.
Wien 39: 89-190 (1889), monogr.
Sometimes vast (T. elephantina Roxb.) marsh
perennials with sheathing leaves, simple flowering
stems, minute imperfect flowers in dense,
cylindrical, terminal spikelike inflorescences,
carpellate and staminate spikes on the same
axes with the staminate distal to the carpellate;
perianth lacking or represented by scales or
trichomes; staminate flowers with 1- 3( - 8)
monadelphous stamens; carpellate flowers uni-
carpellate, with a single 1-ovulate ovary, the stipe
bearing many elongated trichomes producing the
"fluff' of mature spikes; fruit minute, 1-seeded,
wind-dispersed. Eight-13 spp., widely distributed
in temperate and tropical regions of all continents,
often forming extensive monospecific stands.
Kronfeld's monograph now outdated.
 Typhaceae/Xyridaceae
Selected Bibliography
Celakovsky, L. 1885. Uber die Inflorescenz von Typha. Flora
68: 617-630.
Celakovsky, L. 1891. Uber die Verwandtschaft von Typha und
Sparganium. bsterr. Bot. Z. 41: 117-121, 154-160, 195-199,
224-228,266-272.
Chase, M.W. et al. 1995. See general references.
Cook, C.D.K., Nicholls, M.S. 1986/87. A monographic study of
the genus Sparganium (Sparganiaceae). Bot. Helv. 96: 213-
267, 97: 1-44.
Davis, J.I. 1995. A phylogenetic structure for the mono-
coyledons, as inferred from chloroplast DNA restriction site
variation, and a comparison of measures of clade support.
Syst. Bot. 20: 503-527.
Duvall, M.R. et al. 1993. Phylogenetic hypotheses for the
monocotyledons reconstructed from rbcL sequence data.
Ann. Mo. Bot. Gard. 80: 607-619.
Finlayson, C.M., Roberts, J., Chick, A.J., Sale, P.J.M. 1983. The
biology of Australian weeds II. Typha dominguensis Pers.
and Typha orienta/is Presl. J. Aust. Inst. Agric. Sci. 49:3-10.
Frolich, D., Barthlott, W. 1988. See general references.
Harris, P.J., Hartley, R.D. 1980. See general references.
Hegnauer, R. 1963, 1986. See general references.
Herendeen, P.S., Crane, P.R. 1995. The fossil history of the
monocotyledons. In: Rudall, P.J., Cribb, P.J., Cutler, D.F.,
Humphries, C.J. (eds.) Monocotyledons: systematics and
evolution. Royal Botanic Gardens, Kew, pp. 1-21.
Kaul, R.B. 1972. Adaptive leaf architecture in emergent and
floating Sparganium. Am. J. Bot. 59: 270-278.
Kaul, R.B. 1976. Anatomical observations on floating leaves.
Aquat. Bot. 2: 215-234.
Krattinger, K. 1978. Biosystematische Untersuchungen
innerhalb der Gattung Typha L. Mitt. Bot. Mus. Univ.
Ziirich 298. 270 pp.
Linder, P.H., Kellogg, E.A. 1995. Phylogenetic patterns in the
commelinoid clade. In: Rudall, P.J., Cribb, P.J., Cutler, D.F.,
Humphries, C.J. (eds.) Monocotyledons: systematics and
evolution. Royal Botanic Gardens, Kew, pp. 473-496.
Miiller-Doblies, D. 1970. Uber die Verwandtschaft von Typha
und Sparganium im Infloreszenz- und Bliitenbau. Bot.
Jahrb. Syst. 89: 451-562.
Miiller-Doblies, U. 1969. Uber die Bliitenstande und Bliiten
sowie zur Embryologie von Sparganium. Bot. Jahrb. Syst.
89: 359-450.
Nicholls, M.S., Cook, C.D.K. 1986. The function of pollen
tetrads in Typha (Typhaceae). VerOff. Geobot. Inst. ETH,
Stiftung Riibel, Ziirich 97: 112-119.
Punt, W. 1975. The Northwest European pollen flora: 5.
Sparganiaceae and Typhaceae. Rev. Palaeobot. Palynol. 19:
75-88.
Smith, S.G. 1987. Typha: its taxonomy and the ecological
significance of hybrids. Arch. Hydrobiol. 27: 129-138.
Solereder, H., Mayer, F.J. 1933. Systematische Anatomie der
Monokotylen. Heft I. Pandanales-Helobiae-Triuridales. 1.
Teil. Typhaceae-Scheuchzeriaceae. Berlin: Borntraeger.
Thieret, J.W. 1982. The Sparganiaceae in the southeastern
United States. J. Arnold Arbor. 63: 341-355.
Thieret, J.W., Luken, J.O. 1996. The Typhaceae in the south-
eastern United States. Harvard Pap. Bot. 8: 27-56.
Yeo, R.R. 1964. Life history of the common cattail. Weeds. 12:
284-288.
461
Xyridaceae
R.KRAL
Xyridaceae C. Agardh, Aphor. Bot.: 158 (1823), nom. cons.
Rosulate to caulescent, low and delicate to tall and
coarse, perennial to annual, terrestrial (rarely
aquatic) herbs, mostly heliophytes of high-
hydroperiod acidic soils. Roots mostly slender,
diffuse-fibrous, with root hairs. Axis sympodial or
monopodia!. Leaves alternate, distichous or spiral,
ligulate or eligulate, the bases broad, open, sheath-
ing, sheaths frequently equitant and keeled; blades
laterally to dorsiventrally compressed, less often
terete, angulate, or variously canaliculate. In-
florescence lateral or terminal, scapose (rarely
subsessile), the scapes 1-few from axils of scape
sheaths or inner leaves, naked or with distant to
approximate pairs of bracteal leaves, each scape
bearing apically 1(-2) imbricate-bracted spikes or
heads or a panicle of spikes. Flowers perfect, 1-
many, solitary and subsessile in axils of chaffy,
leathery or scarious bracts, or (Achlyphila) pedi-
cellate in spathe axils; perianth in 2 differentiated
whorls; sepals (2-)3, the anterior (inner) 1 a
reduced scale, or subequal to the rest, or (Xyris)
membranous and wrapped around the corolla,
abcissing as the flower opens, the outer 2
subopposite, connivent to connate, chaffy, mostly
navicular, often keeled, persistent around the ripe
fruit; petals 3, equal or unequal, distinct to united,
then salverform or bilabiate, mostly narrowed
to claws or to a narrow tube; stamens 3( -6),
oppositipetalous, borne upon the petals or hypo-
gynous; anthers tetrasporangiate, basifixed, usu-
ally bilocular at anthesis, introrsely or laterally
dehiscent, dehiscing longitudinally; staminodia
(1- )3, hypogynous to borne upon the petals, fila-
mentous to bibrachiate and penicillate-hairy;
gynoecium 3-carpellate, superior, the ovary 1-
locular to completely or incompletely trilocular;
placentation marginal, parietal, basal, free-central
or axile (all conditions found in Xyris, in all other
genera axile); style terminal, proximally terete to
angled, slender, appendaged or exappendiculate,
apically 3-branched to U-shaped stigmas, or vari-
ously dilated to infundibular or laminar or
subcapitate, usually papillate or fringed stigmatic
zones. Fruit capsular, mostly loculicidal; seeds
462 Xyridaceae

usually numerous, rarely 1, mostly with strong

longitudinal ridges and finer cross-lines, translu-
cent or farinose-opaque, the embryo small, situ-
ated at base of or lateral to an abundant mealy
(sometimes oily) endosperm.
An essentially pantropic family of five genera
and over 300 species, most of them in Xyris, the
only genus to occur in both New and Old Worlds,
and to range into the north-temperate zone; the
remaining four genera confined to S America.

VEGETATIVE MORPHOLOGY. The roots are fi-

brous, diffuse, mostly filiform, less often fleshy.
The rootstocks are commonly rhizomes, rarely
bulbous; stems are usually short, less often
elongate, sometimes caudiciform, sometimes
stout and subligneous, mostly with contracted
internodes.
The leaves are equitant and distichous in
Achlyphila and in most Xyris, are polystichous
in Abolboda, Aratitiyopea, Orectanthe and some
Xyris; in all Abolboda, in Orectanthe sceptrum, in
Aratitiyopea and in most Xyris a strong rosette is
formed around a contracted stem or apically on
vegetative shoots of a season. In more elongate-
stemmed examples in Orectanthe they are in high
spiral, or in Achlyphila and some Xyris, form long,
frondlike plates of imbricate-based leaves and no
rosette is evident. The leaf sheaths are open in all
genera. In Achlyphila and most Xyris the blades
are unifacial; in the other genera the blade is dor-
siventrally compressed, often lingulate, variously Fig. 114A-G. Xyridaceae. Abolboda ciliata. A Habit. B Spike.
tapering uninterruptedly from broad sheaths. In C Spread corolla with stamens and staminodes. D Stamen. E
those Xyris in which the blade is not unifacial it Pistil, note appendaged style. F Capsule (left), adaxial view of
capsule valve (right). G Seed. (Kral1992)
is typically narrowly linear to filiform, terete,
or angulate, variously canaliculate, particularly
adaxially. In all genera save Xyris, even in most of
these, the transition from sheath to blade is unin- The roots have an exodermis of one or more
terrupted, but in some Xyris there is a ligule or compact, lignosuberized layers under a larger-
the scarious sheath border breaks into auricles at celled piliferous layer (or this lacking in Xyris
convergence of sheath and blade. brevifolia); root hairs mostly arising toward one
end of an epidermal cell, sometimes in irregular
VEGETATIVE ANATOMY. Anatomical studies have vertical series. The middle cortex is frequently
been done on all genera except the recently de- narrow, sometimes a single layer of cells
scribed Aratitiyopea, these excellently detailed (Abolboda, Xyris brevifolia); in Xyris it ranges
and summarized by Tomlinson (1969), whose from radial plates of cells separated by wide inter-
source particularly for much information on cellular spaces to more uniform cortex. Some-
Abolboda, Achlyphila, and Orectanthe was times horizontal irregular plates of lobed or
Carlquist (1960). Features shared by members of stellate cortical cells cut off from ends oflong cells
the Xyridaceae are: vessels with simple perfora- to form irregular transverse diaphragms or an
tion plates in roots, stems and leaves, and irregular reticulum. The endodermis incon-
amphivasal vascular bundles in rhizome and/or spicuous, of large and uniformly thickened cells
stems. Most have irregular plates of stellate cells in (this thickening specific in degree and orienta-
the root cortex and a more or less uniform distri- tion), usually of 1 layer but sometimes truly
bution of phloem through the root stele. multiseriate. Stele usually narrow in comparison
 Xyridaceae 463

to cortex. The pericycle consists of small, thin- might be a chlorenchymatous layer, including
walled cells, rarely or often interrupted by adja- palisade.
cent vessels. The stele is diarch to polyarch with 2 The mesophyll of most Xyris is uniform, and the
or more peripheral vessels. Protoxylem is absent cells adjacent to substomatal chambers are often
or present as isolated narrow elements. Phloem L-shaped. In a few species the mesophyll may be of
elements are mostly not grouped in peripheral elongate, lobed cells with lobes connivent at the
phloem strands, but are scattered irregularly tip and in thick or terete-bladed species the central
throughout the central ground tissue. mesophyll may be of large colorless cells which
The rhizome or caudex, except in Achlyphila may break to produce 1 or more longitudinal air
has short internodes, and in stocky-stemmed canals. In Achlyphila the mesophyll has a periph-
Abolboda, Orectanthe, it is subligneous. The eral chlorenchymatous layer and a superficial
surface layers are often suberized in Abolboda palisade that surrounds the colourless central
and Orectanthe, sometimes so in Xyris, and in region. In some Abolboda the mesophyll usually
Achlyphila they are thin-walled. lacks distinct adaxial palisade (except in A. ciliata,
Leaf anatomy has been the most studied (see A. grandis, and some A. macrostachya), and it
Arber 1922; Solereder and Meyer 1929; and is compact, mostly unlobed with some having
Tomlinson 1969 for summaries). Epicuticular air canals, in A. linearifolia these alternating
waxes are absent (Frolich and Barthlott 1988), but pectinately with vascular bundles. In Orectanthe
in one Xyris the leaf epidermis was found to be the mesophyll is of lobed, elongate cells and air
covered by a secretion deposited in droplets that canals are lacking.
could not be dissolved by organic solvents. Since Xyris is by far the largest genus and has the
In Xyris the leaf blade epidermis is typically uni- broadest geographic range and spread of habitat,
form and the cells are longitudinally oriented and it is not suprising that its foliar anatomy is the
rectangular. There are many species in which most diverse, particularly as to number and
outer walls are produced into papillae, tubercles, arrangement of foliar venation and vascular
etc., in yet others the end walls are distinctly bundles. Veins range from few (1)-many, from 1
thickened and are variously cutinized. Epidermal series with alternate bundles oriented in opposite
sculpture may be useful taxonomically. In directions to 2 or more series with bundles all
Achlyphila epidermal cell ends sometimes overlap similarly oriented; bundles may be compound or
at their narrowed ends, thus in cross-section the simple. Such variation ranges across what are
epidermis may appear biseriate; walls are much thought to be sections. Achlyphila, nearestXyris in
thickened, anticlinally conspicuously pitted and leaf character, still may be distinguished by its
with transverse protuberances on end walls giving complete bundle sheaths around bundles, vascu-
the blade an irregular epidermal outline. While lar tissues without protoxylem lacunae and meta-
the epidermis in some Abolboda (i.e. A. scabrida) xylem having irregular series and lacking the 2
may be produced into rugulosities or be papillose obvious wide vessels. Abolboda and Orectanthe
or scabrid, hairs are produced only in Xyris. rarely have the compound vascular bundles so
Hairs in Xyris may be uniseriate-glandular, frequent in many Xyris. Tracheids with broad
short-unicellular and unbranched, or rarely annular-helical thickenings have been found in
branched (X. tenella, leaf sheath). In several spe- leaf tips of some Abolboda.
cies long hairs may be produced at sheath margins Calcium oxalate as small crystals has been
and many have ciliate or ciliate-scabrid blades. In reported in the leaf mesophyll (Solereder and
some species the inner surface ofleaf sheaths may Meyer 1929) and the leaf epidermal wall
produce hairs or have secretory function. (Tomlinson 1969) in Xyris. Raphides are lacking.
In Achlyphila and most Xyris stomata are found Tannin is reported, mostly restricted to the epi-
on both sides of the lamina; in the other genera dermis, sometimes found in all cells or scattered
they are strictly abaxial. In all genera studied, irregularly. Starch is found in the rhizomes or
guard cells are not sunken and are similar in con- larger stems and commonly in ground paren-
figuration as are lateral subsidiary cells. Terminal chyma of leaf bases, the grains reported as com-
subsidiary cells are not developed except in some pound in Xyris, simple, large, ellipsoid, or
Abolboda. irregularly rounded in Abolboda.
The hypodermis is well developed in Orectanthe
and most Abolboda; in Xyris it is lacking, though INFLORESCENCE. In all Xyridaceae but Achlyphila
in some there may be a distinct palisade of the inflorescence is a closed, chaffy-bracted,
anticlinally elongate cells, and in Achlyphila this conelike spike or a head which in a few species is
464 Xyridaceae

sessile, but which in most is atop a variously

elongate naked or bracteate (some Abolboda,
Achlyphila) scape. The flowers are sessile or
subsessile in axils of chaffy, loosely to tightly im-
bricate, spirally or decussately arranged bracts. In
Achlyphila, however, the inflorescence is open,
with flowers pedicellate in axils ofleaflike spathes.
The morphology of inflorescence bracts of
spikes and heads is typically comparable to the
sheaths of the foliage leaves. Outer and lower
(sterile) ones may be strongly bladed or leaflike, as
in involucrate Xyris (i.e., X. involucrata, X.
mertensiana, X. nigricans, X. pallidula), or in the
capitate Aratitiyopea gradate to surrounding foli-
age leaves. In most the bracts are chaffy, some
medially longitudinally herbaceous, but all typi-
cally with broad, chaffy, or even scarious borders
and most lacking blades, and in most the bracts
are navicular to relatively broad, convex or keeled,
or lingulate, the outer !-several ones sterile,
mostly keeled, shorter, equal to, or longer than the
fertile, grading into them. Costae (nerves) range
from a single strong median one, or this bordered
by few to several, generally less distinct laterals; in
most the median zone is thickest and is sometimes Fig. llSA-E. Xyridaceae. Flower structure in Xyris. A Ventral
evidently herbaceous. In Xyris this herbaceous and cross-sectional view of sepals in bud. B Same of petals.
zone, mostly sharply differentiated in texture and C Staminode, showing yokelike blade and moniliform
color, running the length of the bract or more staminodial beard hairs. D Pistil, showing style base, tubular
often terminal or subterminal is referred to as style branches, and stigmas. E Diagrammatic representation
of flower. 1 Disarticulated dorsal sepal; 2 lateral sepals; 3 petal
dorsal area, and it may be specific in character. blades (claws foreshortened) with adnate stamens; 4 stamino-
Bract arrangement may be entirely spiral, entirely dia, claws foreshortened, showing flattened, bifurcate blades
distichous, or decussate at one level (usually and terminal tufts of trichomes; 5 cross-section of flower
proximally) and spiral upspike. towards base as oriented in bud; 6 stylar apex, showing
horseshoelike stigma pattern at about level of staminode tip.
(Krall988)
FLORAL STRUCTURE (Fig. 115). In all the genera
the flowers are perfect and the perianth is in 2
differentiated whorls. The calyx is regular or
nearly so in Achlyphila, Aratitiyopea, and connate at base, more often with broad blades and
Orectanthe, the 3 members distinct, chaffy, nav- clawed bases, and are mostly buttercup yellow,
icular or conduplicate; in Xyris and Abolboda it is less often white. In Aratitiyopea the petals are
distinctly irregular, in the former with the inner equal, connate, the corolla tubular-salverform,
(anterior) sepal membranous and caducous, white to pale purple. In Abolboda, they are
inclosing the rest of the flower in bud, the other 2 equal, connate, funnelform-tubular to spreading,
(outer) chaffy, navicular, keeled-carinate, free or broad, white to blue or lavender lobes, while in
connate, clasping the ripening fruit. In Abolboda Orectanthe the irregular gamopetalous yellowish
the anterior sepal is generally reduced, even lack- (rarely lavender) corolla is distinctly bilabiate.
ing, the laterals distinct, mostly navicular and Fertile stamens are 3 in all genera (with the
carinate-keeled, often similar to spike bracts. In exception of one Xyris, which has 6) and are
the other genera the sepals are strongly convex oppositipetalous. In Achlyphila they are hypogy-
except towards the apex, there often with pro- nous, and in the other four genera borne upon
nounced but entire keel. the petals with short or long filaments. In all
The corolla is distinctive in all genera and is genera the anthers are tetrasporangiate, dithecal,
always tripartite. In Achlyphila petals are equal are mostly emarginate apically, the bases often
and distinct, with blades yellowish, tapering to separated by wide thin connective and auriculate
base; in most Xyris the petals are equal, rarely to sagittate; they seem to be basifixed throughout.
 Xyridaceae 465

Staminodia are lacking in Achyphila, Arati-
tiyopea, and Orectanthe; in Abolboda they are
lacking or filamentous and often less than 3, in any
case borne upon the petals below corolla sinuses;
in Xyris they are the most elaborate and are usu-
ally hypogynous and long-clawed, the flat narrow
claws branching distally, the flattened branches
equipped with variably elongate, mostly penicil-
late hairs or in a few cases reduced, rarely lacking.
In all genera the gynoecium is tricarpellate, the
ovary superior. In Xyris, placentation ranges from
entirely axile to basal-axile, basal, parietal, or
marginal; in the other genera it is axile. In those
Xyris with basal or basal-central placentation, the
funiculus is often much longer than the mature
seed. Styles are terminal, elongate in all genera. In
Xyris the tubular style has 3 well-defined branches
distally, each branch tubular-conduplicate and
terminating in a U-shaped to shallowly infundi-
bular, papillate to fimbriate stigmatic zone; in
Achlyphila the stigmatic zone is nearly continuous
with the style, small and 3-lobed; in Aratitiyopea
the stylar apex is produced into 3 connivent,
bristly glandular lobes; in Abolboda it is mostly
infundibular with erose to fimbriate edges; in
Orectanthe it is typically distinctly bent with a
capitate-patelliform, papillate to fimbriate stigma.
Stylar appendages are lacking in Achlyphila
and Xyris, but well developed in the other three
genera and always 3 in number. In all these cases
there are 3 appendages on a style. In Abolboda the
(usually) triquetrous style has a lateral, ascending
to spreading or reflexed, stipitate or dilated-
bladed appendage on each of the broader side
wings and a 3rd, usually filamentous and reflexed,
from the adaxial, narrower wing; the broader
appendage blades are often auriculate at junction
with stipe. In Aratitiyopea, the appendages
are at the very style base, with erect, terete stipes
and flattened, sharply reflexed, flabelliform
blades. In Orectanthe, there is a similar basal
arrangement, the narrow, firm, terete appendages
erect around the style base, then sharply reflexed-
curved.
endosperm development are known (Tiemann
1985; Johri et al. 1992).
POLLEN MORPHOLOGY. Pollen grains sulcate (or
occasionally disulcate?) in Xyris and inaperturate
and spheroidal in Abolboda, Orectanthe, and
probably Achlyphila (Fig. 116). They can be very
large [most pollen grains of Abolboda studied by
Erdtman (1952) and Carlquist (1960) have a
diametre of over 100 [-Lm, those of the two species
of Orectanthe range from 140-165 [,tm, but those of
Achlyphila and Xyris have less than 50 [-Lm
(Erdtman 1952)] and then only a small number of
them is formed in each anther (Carlquist 1960).
This enormous size of the pollen grains seems to
be related to their function as reward for pollina-
tors. The exine is thin and not very resistant to
acetolysis; it is composed of 2 layers of different
stainabilty and is underlain by a thin intine. The
surface is smooth in Xyris and Achlyphila, but
ornamented by large spines or knobs which are
more resistant to acetolysis in the remaining three
genera.
KARYOLOGY. Thus far, studies of microsporogen-
esis in American Xyris have shown complements
of n = 9 and root tip work showing 2n = 18. In
African Xyris (X. capensis), however, sporophyte
counts have been 2n = 34. Fedorov (1969) adds 2n

EMBRYOLOGY. There is considerable vanat10n

with respect to embryological traits within the
family. Different kinds of endothecium and tape-
tum structure have been reported, but the division
of pollen mother cells is always successive. Mature
pollen grains are indicated as 2- or 3-nucleate. The
ovules are anatropous (Achlyphila and Alboboda),
atropous (Xyris), or slightly campylotropous Fig. 116. Xyridaceae. Xyris involucrata, distal face of pollen
(Orectanthe), bitegmic and slightly crassinucellar with U-shaped sulcus, SEM X2200. (Photo Palynological
(Tiemann 1985), and both Nuclear and Helobial Laboratory Stockholm)
466 Xyridaceae
= 26 and 32 for Xyris species. In Abolboda counts
have been registered as n = 8, 9, 13, 17.
POLLINATION. No nectar is produced by the flow-
ers of the Xyridaceae, and pollinating bees are at-
tracted by pollen or imitated pollen masses (Vogel
1981); only Orectanthe is ornithophilous. S.
Renner (pers. comm.) found Xyris sp. in Minas
Gerais (Brazil) buzzed and pollinated by females
of an anthophorid Exomalopsis sp.
FRUIT AND SEED. In all genera the fruit is a locu-
licidal capsule. The 3 valves separate to the fruit
base and the capsule walls are thickened or differ-
entially thickened in most species except those of
Xyris, where valves are usually thin.
Seeds in Achlyphila, Aratitiyopea, and Xyris are
typically symmetric, ranging from subglobose to
broadly ellipsoid or ovoid (in the former two),
additionally (in Xyris) to variously ellipsoid or
cylindric, frequently curved, often with the outer
integument produced to an apical coma of scales
or fimbriae. In Abolboda they are mostly thick-
ened and curved dorsally, are asymmetrically obo-
void, strongly ribbed on the thickened backs,
whilst in Orectanthe they are laterally compressed,
mitten-shaped, with a broad wing external to the
embryo on one side, more narrowly on the edge
opposite the thumb of the mitten. In Abolboda,
Achlyphila, Aratitiyopea, and in most Xyris they
are small, rarely over 1.2 mm. In some Xyris with
basal or basal-central placentation some may ex-
ceed 3 mm (including apiculus and apical coma)
and in Orectanthe they may range from 2-4 mm.
In all the embryo is basal or lateral, the endosperm
copious, mealy, starchy and proteinaceous, the
starch grains (in Abolboda) compound. Seed-coat
sculpture in Xyris and Abolboda is often
specifically distinct, being variously papillate,
longitudinally distinctively ribbed, variously
cross-ribbed, the outer integument variously
produced in some Xyris into a distinctive coma of
scales or hairs.
Both integuments take part in the composition
of the seed coat, but only the tegmic layers, which
are filled with a resinlike or tanninlike substance,
provide mechanical strength (Carlquist 1960).
DISPERSAL. Seeds are shaken or fall from dehisc-
ing capsules, particularly as aging bracts and
sepals spread away from the fruit or as old scapes
topple. These seeds are often buoyant, thus are left
in drifts on the surface or are mixed with small
sand particles and silt or homogenized peat in a
shallow patina. If the substrate is moist, germina-
tion can take place within 2 weeks (in Xyris) and
young seedlings can form a veritable turf on open
seepage wherever there are clearings. On the other
hand, since abundant moisture triggers germina-
tion, if conditions are dry, Xyris seeds can remain
dormant for indefinite periods. I have kept dry
seed without refrigeration for up to 10 years and
still had good germination in greenhouse fiats.
I have run no such tests on seeds of the other
genera.
PHYTOCHEMISTRY. Apart from common flavonol
and flavone glycosides and proanthocyanidin, the
anthraquinone chrysazin has been detected in a
Xyris; this substance class may be widespread in
Xyris as judged from the reddish sublimation on
herbarium sheets. Cyanogenesis was found in one
Xyris (Hegnauer 1963, 1986). The family was posi-
tive in Harris and Hartley's (1980) tests for cell
wall-bound ferulate.
RELATIONSHIPS WITHIN THE FAMILY. Only the
Guayana Highlands has examples of all the genera,
thus this ancient Precambrian land mass acts as
what must be a primary centre. Circumstantial
(morphologic) evidence points to 2 distinct clus-
ters of character states, 1 having Abolboda,
Aratitiyopea, and Orectanthe, the other contain-
ing Achlyphila and Xyris. The former group is dis-
tinguished by its polystichous, bifacial leaves in
which there is no abrupt transition from sheath to
blade, blades usually with a colorless hypodermis
adaxially and often abaxially, a high degree of
petal connation, spinose pollen, staminodia
reduced to filaments or lacking, styles in most
species asymmetric and appendaged, and placen-
tation exclusively axile. The latter group has obvi-
ously or obscurely distichous, equitant-sheathed,
often ligulate, unifacially bladed leaves in which
there is often an abrupt transition from sheath to
blade, and blades typically lacking the colorless
hypodermis, a low degree of petal connation or
none, with pollen lacking spines, staminodia typi-
cally present and bifid (lacking in the monotypic
Achlyphila), styles symmetric and unappendaged,
and placentation ranging from axile to basal, cen-
tral, parietal, or marginal. Such evidence could
lead to the suggestion that the 2 branches arose
from ancestral stock, presumably in the Guayana
Highlands, that was rhizomatous, perennial, with
polystichous, bifacial leaves, was scapose, had
flowers in bracteate racemes, had sepals and petals
distinct and equal, had 6 distinct, hypogynous
stamens and a superior tricarpellate ovary with
an axile placentation (Kral 1992).
 Xyridaceae 467

FAMILY AFFINITIES. Nakai (1943) was of the

opinion that Abolboda and Orectanthe (first de-
scribed as an Abolboda), on a basis of their dis-
tinctive leaf and stem anatomy, and distinctive
spinose pollen should be broken out of the
Xyridaceae as the Abolbodaceae; this lead was
followed temporarily by Takhtajan (1959), but
only prior to the discovery of Achlyphila. Most
phylogenists since (Takhtajan 1980; Cronquist
1981; Thorne 1967) treat all genera as Xyridaceae.
The family has now often been considered as
being most closely related to the Rapateaceae and
Eriocaulaceae in the Commelinales. By and large
this is confirmed by the recent cladistic and mo-
lecular analyses of Linder and Kellogg (1995),
who, however, additionally have brought the
Bromeliaceae into the play, while in the restriction
site analysis of Davis (1995), Xyris appears as
sister to Carex.

DISTRIBUTION AND HABITATS. The family is es-

sentially pantropic, with four of the five genera
confined to S America and of these three,
Achlyphila, Aratitiyopea, and Orectanthe are lim-
ited to northern S America along with most of the
20 Abolboda species, with only a few of those
reaching southward into S Brazil and only one
ranging to Paraguay. Only Xyris has representa-
tives in the Eastern as well as the Western Hemi-
sphere with extensions northward into temperate Fig. 117. Xyridaceae. Orectanthe sceptrum, growing on
America and eastern Asia (China, Taiwan). Nearly flooded peat together with Xyris sp. and Eriocaulaceae.
all species are heliophytic and inhabit acidic, Venezuela, Mt Roraima. (Photo K. Kubitzki)
moist to wet habitats (Fig. 117); however, a few are
true aquatics (i.e., X. aquatica).
4. Flowers distinctly pedicelled; sepals alike; staminodia lack-
ing; style unbranched, the stigma trilobed, capitate; petals
KEY TO THE GENERA lacking claws; stamens hypogynous I. Achlyphila
- Flowers subsessile or sessile; sepals unlike, the lateral ones
1. Petals gamopetalous; leaves polystichous; styles or ovary chaffy and keeled, the inner one membranous; staminodia
summit appendaged; pollen spinose or papillate; corolla usually present, mostly bibrachiate; style 3-branched, the
regular or irregular; sepals 2- 3 2 stigmas mostly U-shaped; petal blades on claws; stamens
- Petals usually distinct; leaves distichous, mostly equitant; borne on petals 2. Xyris
styles unappendaged; pollen lacking spines or other protu-
berances; corolla regular; sepals 3 4
2. Stylar appendages mostly well above style base on style;
GENERA OF THE XYRIDACEAE
style often triquetrous at base; capsule apex thickened;
flowers 1- several, the inflorescence sessile or on variously
elongate, opposite-bracted scapes or in panicles; sepals 1. Achlyphila Maguire & Wurdack
2(- 3) 3. Abolboda
- Stylar appendages at or around style base; style terete; cap- Achlyphila Maguire & Wurdack, Mem. N.Y. Bot. Gard. 10 (2):
sule apex not appreciably thickened; flowers many in dense, 12 (1960); Kral, Ann. Mo. Bot. Gard. 79: 874-877 (1992),
globose or hemispheric large heads; sepals 3 3 rev.
3. Corollas arching-spreading, irregular, yellow (rarely
redpurple); style apex bent outward, stigma capitate, Perennial from slender, branched rhizomes;
lateral-terminal, papillate; seeds winged, 2-4mm, irregular, leaves narrowly ascending, equitant-distichous,
mitten-shaped 5. Orectanthe
- Corollas erect, regular, salverform, white to purple; stigma
the blades linear-triangular, unifacial. Scapes
terminal on erect style tip, trilobed, the lobes pilose; seeds single, terminal, compressed; inflorescence usu-
wingless, ridged-and-pitted, symmetric 4. Aratitiyopea ally with 2 approximate, distichously arranged
468 Xyridaceae
leaflike spathes subtending 1-3 flattened-
pedicelled flowers, sepals 3, landform, chaffy; pet-
als 3, distinct, unclawed, dull yellow; stamens 3,
distinct from petals; staminodia lacking; style
erect, linear, unappendaged. Only one sp., A.
disticha Maguire & Wurdack, summit elevations
on Cerro Neblina along the Brazil-SW Venezuela
border (1800-2300 m).
2. Xyris L.
Xyris L., Sp. Pl.: 42 (1753); Smith & Downs in Fl. Brasilica 9 (2):
1-215 (1968); Kral, Sida 2: 177-260 (1966), spp. USA &
Canada; Kral, Ann. Mo. Bot. Gard, 75: 522-722 (1988), spp.
northern S America.
Usually perennials, or annual to short-lived
plants; usually rosulate (less often caulescent),
solitary or caespitose, sometimes rhizomatous or
bulbous herbs; axis sympodial; leaves mostly
equitant and distichous. Scapes naked, terete to
compressed, ending in a single conelike spike or a
head (rarely bispicate); outer bracts sometimes
involucra! but mostly chaffy, grading to 1 or more
fertile bracts; flowers subsessile, solitary in bract
axils; sepals 3, the 2 laterals mostly chaffy, the
inner membranous and infolding the corolla bud,
abcissing when flower spreads; petals 3, distinct,
mostly clawed, yellow( -white); stamens 3;
staminodia hypogynous, mostly long-clawed,
bibrachiate, rarely reduced or lacking; style
erect, unappendaged, conduplicate-tubular, 3-
branched, terminating in U-shaped or semi-
infundibular stigmas. n = 9, less often 17; 2n = 18,
rarely 34. Nearly 300 described spp., mostly in S
America, several in Africa, Australia, and Asia, in
America extending N to Canada and in Asia north
to China and Taiwan. Three sections have been
described, mostly based on differences in placen-
tation, but their distinction is risky at best, par-
ticularly if there is no geographic correlation as
was once believed; a study of Xyris of the Guayana
Highlands reveals examples of all three "sections"
from the same habitat in the savannas and tepuis
of that region.
3. Abolboda Kunth Fig. 114
Abolboda Kunth in Humb. & Bonpl., Pl. Aequinoct. 2: 25, pl.
114 (1809); Kral, Ann. Mo. Bot. Gard. 79: 820-874 (1992),
rev.
Cespitose or solitary, mostly glabrous perennials,
rarely annuals; leaves mostly basal. Inflorescences
subsessile to strongly scapose, scapes naked or
bracteate, terminating in a head or spike (rarely in
a panicle of spikes), the inflorescence bracts
mostly chaffy in spiral, the lower empty, the fertile
loosely or tightly imbricate, each subtending a
single flower; sepals 2( -3); corolla gamopetalous,
actinomorphic, salverform, white, blue or laven-
der; stamens 3; staminodia 0-1-3; style usually
appendaged, mostly triquetrous-winged. Twenty
neotropical spp., boggy, warm to cool habitats,
centring in the savannas of northern S America.
4. Aratitiyopea Steyerm. & Berry
Aratitiyopea Steyerm. & Berry, Ann. Mo. Bot. Gard. 71: 297
(1984); Kral, Ann. Mo. Bot. Gard. 79: 877-879 (1992), rev.
Robust, "Navia" -like perennials from a thick rhi-
zome; leaves polystichous, crowded toward shoot
apex, linear-lorate, lingulate, sharp-tipped, the
longest downstem, the upper grading into a large,
chaffy-involucrate, hemispheric head of many
bracts, the inner ones increasingly narrower and
directly subtending flowers; sepals 3, similar to
cephalar bracts but slightly longer; corolla
tubular-salverform, actinomorphic, 7-10 em,
white to pale purple; stamens 3; staminodia lack-
ing; ovary dorsiventrally compressed; style
straight, basally provided with stipitate, recurved,
broadly flabellate appendages. Only one sp., A.·
lopezii (L.B. Smith) Steyerm. & Berry, with two
varieties, rare on arenaceous rock outcrops in NW
Brazil, SW Venezuela, and SE Colombia, at 250-
1600 m. Highly ornamental, described earlier as
the bivarietal Navia lopezii by L.B. Smith (Bot.
Mus. Leafl. 15: 40. 1951; ibid 16: 195. 1954).
5. Orectanthe Maguire Fig. 117
Orectanthe Maguire, Mem. N.Y. Bot. Gard. 10:2-3. 1958; Kral,
Ann. Mo. Bot. Gard. 79: 879-885 (1992), rev.
Glabrous perennials; stems either short, stout,
producing dense basal rosettes or lax, decumbent,
branching to short, erect tips. Leaves polysti-
chous, clasping-based, sharp-tipped; scapes 1-
several, naked, variously elongate. Inflorescence a
large, terminal burrlike head; outer and cephalar
bracts shorter than sepals; sepals 3, chaffy; corolla
gamopetalous, bilabiate, pale yellow, rarely pur-
plish; stamens 3; staminodia lacking; placentation
axile; style appendages basal, linear, doubled
back. Two spp., northern S America on the
Roraima sandstones of the Guayana Highland
at medium to high elevations, usually in boggy
pockets amongst rocks and in full sun.
 Xyridaceae 469

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beziehungen der Farinosae. Ein Beitrag zum System der
Monocotyledonen. Willdenowia 2: 639-768.
Hamann, U. 1962. Weiteres tiber Merkmalsbestand und
Verwandtschaftsbeziehungen der "Farinosae." Ibid. 3:
169-297.
Harris, P.J., Hartley, R.D. 1980. See general references.
Hegnauer, R. 1963, 1986. See general references.
Idrobo, J.M. 1954. Xiridaceas de Colombia. Caldasia 6 (20):
184-260.
Johri, B.M. eta!. 1992. See general references.
Kral, R. 1966. Xyris (Xyridaceae) of the continental United
States and Canada. Sida 2: 177-260.
Kral, R. 1983. The Xyridaceae in the southeastern United
States. J. Arnold Arbor. 64: 421-429.
Kral, R. 1988. The genus Xyris (Xyridaceae) in Venezuela and
contiguous northern South America. Ann. Mo. Bot. Gard.
75: 522-722.
Kral, R. 1992. A treatment of American Xyridaceae exclusive
of Xyris. Ann. Mo. Bot. Gard. 79: 819-885.
Linder, H.P., Kellogg, E.A. 1995. Phylogenetic patterns in the
Commelinid clade. In: Rudall, P.J., Cribb, P.J., Cutler, D.F.,
Humphries, C.J. (eds.) Monocotyledons: systematics and
evolution, Vol. 2. London: Royal Botanic Gardens, Kew,
pp. 473-496.
Maguire, B. 1958. Xyridaceae. In: Maguire, B., Wurdack, J.J.,
eta!. (eds.) Botany of the Guayana Highland, part III. Mem.
N.Y. Bot. Gard. 10 (1): 1-19. [Orectanthe described]
Maguire, B., Wurdack, J.J. 1960. Xyridaceae. In: Maguire, B.,
Wurdack, J.J., eta!. (eds.) Botany of the Guayana Highland,
part IV. Mem. N.Y. Bot. Gard. 10 (2): 11-15. [Achlyphila
described.]
Malme, G.O.A. 1930. Xyridaceae. In: Engler, A., Prantl, K.
(eds.) Natiirliche Pfianzenfamilien, 2nd edn., 15a: 35-38,
figs 11-15. Leipzig: W. Engelmann.
Nakai, T. 1943. Ordines, familiae, tribi, genera, sectiones, spe-
cies, varietates, formae et combinationes novae a Prof.
Nakai-Takenoshin adhuc et navis edita. Tokyo.
Solereder, H., Meyer, F.J. 1929. Xyridaceae. Systematische
Anatomie der Monokotyledonen 4. Berlin: Borntraeger,
pp. 36-50.
Suessenguth, K., Beyerle, R. 1935. Dber die Xyridaceengattung
Abolboda Humb. et Bonpl. Bot. Jahrb. Syst. 67: 132-142.
Takhtajan, A. 1959. Die Evolution der Angiospermen. Jena:
G. Fischer.
Takhtajan, A. 1980. Outline of the classification of flowering
plants (Magnoliophyta). Bot. Rev. 46: 225-359.
470
Zannichelliaceae
R.R. HAYNES, D.H. LES,
and L.B. HOLM-NIELSEN
Zannichelliaceae Dumort., Anal. Pam. Pl. 59, 61 (1829), nom.
cons.
Entirely submersed glabrous annual or perennial
herbs of fresh or brackish waters. Roots fibrous,
1-7, nonseptate, borne at lower nodes. Stems
slender, dimorphic, the lower often stoloniferous,
the upper erect and leafy, without teeth, turions, or
tubers. Leaves alternate, opposite, or pseu-
dowhorled, scalelike, without vascular tissue, or
foliaceous, linear, 1- or rarely 3-veined, subterete,
sessile, with basal sheath, the sheath adnate to
or free from blade; infravaginal scales present,
membranous. Inflorescence axillary, with 2-several
flowers. Flowers unisexual, monoecious, more
rarely dioecious; staminate short-pedicellate, peri-
anth mostly absent, rarely minute and 3-lobed,
stamen 1, connective extending into a blunt tip,
anther 2-12-loculed, dehiscing by longitudinal
slits, pollen often contained in gelatinous matrix;
carpellate short-pedicellate, often enclosed by
membranous spathelike envelope, perianth a
small cuplike sheath, or of 3 separate segments,
carpels 1-8, separate, short-stipitate, 1-locular,
ovule solitary, bitegmic, ventral-pendulous, anat-
ropous, placentation apical, styluli short to long,
stigma enlarged, funnel-shaped, feathery, or
pelatate. Fruit drupaceous, with membranous exo-
carp, fleshy mesocarp, and stony endocarp. Seed
solitary; embryo curved; endosperm Helobial in
development, absent in mature seed.
A family of near cosmopolitain distribution with
4 genera and 10-12 species.
VEGETATIVE MoRPHOLOGY. (Posluszny and
Tomlinson 1977). The plants grow in clones in
shallow, generally brackish, coastal waters or in
inland freshwater lakes.
The roots are all shoot-borne, unbranched, aris-
ing from the rhizome or nodes of the stem. The
rhizomes are creeping and sympodially branched.
The stems are erect, richly branched and leafy.
The leaves are distichous. The rhizome scales are
membranaceous, and without blades. The foliage
leaves are sheathing, with open, membranous,
overlapping sheaths, the sheaths extended and
Zannichelliaceae
ligulelike or auriculate. The blades are linear,
inserted just below the sheath, and the apices
rounded, pointed, truncate or toothed, without
an apical pore. Pairs of inconspicuous, filiform
squamules occur laterally at the nodes.
VEGETATIVE ANATOMY. All four genera have
been studied anatomically (review by Tomlinson
1982). The anatomy of the Zannichelliaceae is
simplified by reduction.
The unbranched roots have a thin-walled epi-
dermis of large cells and conspicuous root hairs
that arise from short trichoblasts. The outer cortex
is compacted into an exodermis of 1 or 2 layers of
narrow, slightly lignified, thick-walled cells. The
cortex is lacunose, the endodermis is uniserate
and thin-walled, and the stele is narrow, sur-
rounding a metaxylem lacuna.
The stems are nearly without mechanical tissue
and have a thin-walled epidermis. The cortex is
wide and lacunose, the endodermis uniseriate and
thin-walled, and the stele is narrow. The vascular
bundles supporting lateral organs diverge directly
from the stele, and there is no cortical vascular
system.
The leaf base is sheathing and colorless with
biseriate margins that include fibrous strands in
Althenia and Lepilaena. The scale leaves are
equivalent to bladeless sheaths.
The blade is glabrous with a uniform, thin-
walled, chlorophyllous epidermis. The epidermis
is mostly without stomata, although stomata oc-
cur in apices of leaf blades of certain Zannichellia
races. The mesophyll is lacunose of with lacunae
on each side of the midvein (Pseudalthenia and
Zannichellia). The vascular system is reduced to
a single median vascular bundle surrounded by
a uniseriate endodermis. The leaf blades have
submerginal fibers.
INFLORESCENCE STRUCTURE. The plants are mo-
noecious or rarely dioecious, with complex, sym-
podial inflorescence of reduced flowers subtended
by reduced bractlike leaves. The inflorescences
usually have 1 staminate flower terminating the
first order meristem, and often have several car-
pellate flowers terminal on branches of higher
orders (Fig. 118).
FLORAL STRUCTURE. The staminate flowers are
short-pedicellate, reduced to 1 stamen and are
provided with or without a short 3-lobed scalelike
perianth. The anther consists of 1 or more
bisporangiate units, with a short connective
appendage in Lepilaena. Anther dehiscence is by
 Zannichelliaceae
Fig.llSA- 1. Zannichelliaceae. Zannichellia palustris. A Fruit-
ing shoot. B Node with staminate and carpellate flowers, base
ofleaf (to right), portion of stem, and base of two branches. C
Staminate flower (a single stamen) and carpellate flower with
two carpels. D Anther shedding pollen. E Cross-section of
anther. F Nearly mature fruits of a four-carpellate flower, one
carpel undeveloped. G Endocarp with tip of style still covered
by outer part of pericarp. H Fruit, the ovary opened to show
embryo. I Embryo. (Haynes and Holm-Nielsen 1987)
longitudinal slits. In some Zannichellia, Singh
(1965) noticed 2 distinct groups of xylem ele-
ments, each surrounded by its own phloem, in
connectives of anthers that had 4 chambers. He
interpreted these 2 xylem bundles to indicate
that the stamen was actually composed of 2
stamens fused their entire length. Posluszny and
Tomlinson (1977) indicated there is no develop-
mental evidence to support the interpretation
of Singh, since every stamen, regardless of the
number of microsporangia present, is initiated as
a primordium.
The carpellate flowers are of 1-8, separate,
short-stalked, slightly asymmetrical carpels on a
471
short pedicel. The carpels are surrounded by a
perianth that consists of a closed tubelike struc-
ture in Zannichellia and Pseudalthenia and of 3
separate segments in Althenia and Lepilaena, with
the segments opposite the carpels. Each carpel in
Althenia and Lepilaena is surrounded by a peri-
anth segment, whereas all carpels collectively in
Zannichellia and Pseudalthenia are surrounded
by a tubular perianth. The styluli are elongate
with enlarged, peltate, funnel-shaped, stigmas.
The stigmas have more or less lacerate margins, or
are occasionally feather-shaped (Lepilaena bilo-
cularis). According to Posluszny and Tomlinson
(1977), much controversy exists as to whether the
carpellate organs are flowers or inflorescences,
but they were unable to state with certainty which
the structures are.
EMBRYOLOGY. The type of anther wall formation
is unknown. The tapetum is periplasmodial, mi-
crosporogenesis successive, and the pollen grains
are 3-celled when shed. The carpels are uniovulate
and the ovules bitegmic, anatropous, and pendu-
lous. A parietal cell is cut off from the archesporia!
cell (Dahlgren and Clifford 1982).
The embryo sac is of the Allium type. En-
dosperm formation is Helobial. Embryo develop-
ment is of the Caryophyllad type.
Fruits, achenes, and ripe seeds are without
endosperm or perisperm. The embryo is curved,
spirally coiled, and lacks chlorophyll.
PoLLEN MORPHOLOGY. The pollen grains are dis-
persed within a gelatinous matrix in Lepilaena
(Cox and Knox 1989) and Zannichellia (Haynes
and Holm-Nielsen 1987). Those of Zannichellia
are globose, inaperturate, hydrophilous, and have
a net -shaped exine wall structure (Van Vierssen et
al. 1982). Pollen of Lepilaena is globose, hydro-
phobic, and floats on the surface of the water. It
will not mix with water even following shaking or
stirring (Cox and Knox 1989).
KARYOLOGY. Chromosome numbers of Zanni-
chellia include 2n = 12, 24, 28, 32, 34, and 36 (Van
Vierssen 1982). Numbers for the remaining genera
are unavailable (Les and Philbrick 1993).
PoLLINATION. Pollination has been studied only
in Lepilaena and Zannichellia (Cox 1988). Accord-
ing to Cox and Knox (1989), Lepilaena anthers
from staminate flowers are abscised underwater
and then float to the surface. Once reaching the
surface, the anthers dehisce explosively, scattering
pollen over the water surface. Floating pollen ag-
472 Zannichelliaceae
gregates into "search vehicles", which are numer-
ous pollen grains embedded in a gelatinous ma-
trix. Carpellate flowers terminate in long pedicels,
with their elongate, concave stigmas reaching the
water surface. All 3 carpels are held together by a
membranous bract with the three stigmas func-
tioning as a single unit, creating a very slight
depression in the water surface. Pollination is ac-
complished when a floating "search vehicle" falls
into this depression. Pollination in Zannichellia is
less well understood. Van Vierssen et al. (1982)
indicate the hydrophilous pollen is released un-
derwater and sinks. The peltate stigma is produced
underwater and, presumably, as the pollen sinks, it
contacts a stigma, accomplishing pollination.
FRUIT. The fruits are drupaceous, with a mem-
branaceous exocarp, fleshy mesocarp, and stony
endocarp. The endocarp of Zannichellia often
is coarsely papillose. The seeds are without
endosperm, and the embryo is curved.
DISPERSAL. The fruits are water dispersed, but
the dispersal systems are incompletely known.
Presumably, the fruits can be carried over various
distances in the intestines of waterfowl.
PHYTOCHEMISTRY. Crystals or tannins have not
been observed in the Zannichelliaceae, but fla-
vonoid sulfates are known from Zannichellia. Les
and Haynes (1995) report 0-glycosyl flavones also
in Zannichellia.
AFFINITIES. Most contemporary authors have
segregated Althenia, Lepilaena, Pseudalthenia,
and Zannichellia as a separate family (Zannichel-
liaceae) with uncertain affinitites. The marine gen-
era of Cymodoceaceae and Posidoniaceae have
occasionally been included within a broader con-
cept of Zannichelliaceae, but these families are
highly dissimilar (Tomlinson 1982), and there is no
phylogenetic evidence to warrant such an associa-
tion. Recent molecular studies (Les et al. 1993; Les
and Haynes 1995; D.H. Les, unpubl.) indicate that
Zannichelliaceae are most closely related to
Potamogetonaceae. There studies resolve a robust,
monophyletic clade consisting of Zannichellia and
Lepilaena and Groenlandia and Potamogeton. At
the molecular (rbcL) level, divergence among
Potamogeton subgenus Coleogeton, Groenlandia,
Lepilaena, and Zannichellia is insufficient to re-
solve their interrelationships adequately; however,
Potamogeton subgenus Potamogeton is relatively
more divergent and distinct from these genera
(D.H. Les, unpubl.).
Although indications of the relationship be-
tween Zannichelliaceae and Potamogetonaceae
have been obscured because of extensive reduc-
tion and specialization that characterizes both
vegetative and reproductive structures of these
families, some features, such as the spirally
coiled cotyledons of Groenlandia and genera of
Zannichelliaceae, may be synapomorphic. From a
taxonomic standpoint, it seems as reasonable to
combine the Zannichelliaceae and Potamogetona-
ceae as it is to maintain them as separate. The
Editor has encouraged us to retain the more con-
servative concept and recognize Zannichelliaceae
as a separate family. However, the current phylo-
genetic evidence emphasizes the relationship be-
tween Zannichelliaceae and Potamogetonaceae,
which should be recognized as closely related
sister groups.
DISTRIBUTION AND HABITATS. Zannichellia is
nearly cosmopolitan, Pseudalthenia is endemic to
the Cape Province, S Africa, Althenia is endemic
to the Mediterranean region, and Lepilaena is en-
demic to S Australia, Tasmania, and New Zealand.
All species of Zannichelliaceae occur in aquatic
habitats including freshwater lakes or shallow,
brackish coastal waters. All species grow as sub-
mersed hydrophytes in clones on muddy
substrates. The species of Zannichelliaceae are
important in stabilizing the muddy areas along
the edge of marshes and delta.
KEY TO THE GENERA
1. Distal leaves pseudowhorled, blade accompanied by a
free membranous stipular sheath. Distal branching sympo-
dial, the branch complex associated with a male and a fe-
male flower, the female flower lowest on the branch unit.
Female flower (1-)4-5(-8)-carpellate. Fruit usually with an
abaxial (sometimes also adaxial), warted ridge
1. Zannichellia
- Distal leaves distichous, never pseudowhorled, at most
subopposite, with an attached sheathing base. Distal
branching sympodial or monopodia!, the groups of flowers
cincinnuslike, not fixed in number; lowest flower within
inflorescence unit always male. Female flower never more
than 3-carpellate. Fruit smooth or warted but never ridged
2
2. Leaf blade up to 2 mm wide with a submarginal "vascular
strand" and oblique transverse strands continuous with
midvein. Male flower naked, 8-sporangiate with a pair of
vestigial appendages on the connective. Female flower
always 1-carpellate, with a membranous envelope. Fruit
uniformly warted 2. Pseudalthenia
- Leaf blade less than 2mm wide, with a single midvein. Male
flower with a short 3-lobed perianth immediately below the
2- or 12-sporangiate anther. Female flower (1-)3-carpellate,
each carpel subtended by a separate tepa!. Fruits usually
smooth 3
 Zannichelliaceae
3. Male flower 2-sporangiate; carpels each with a long style
and peltate stigma. Fruit dehiscent. Leaf fibers sometimes
absent. Mediterranean 3. Althenia
- Male flower either 2- or 12-sporangiate; carpels each with a
short style and a funnel-shaped or feathery stigma. Fruit
reportedly indehiscent. Leaf fibers usually well developed.
Australasian 4. Lepilaena
1. Zannichellia L. Fig. 118
Zannichellia L., Sp. Pl. 2: 969 (1753).
Annual or rarely perennial of fresh or brackish
waters. Leaves in pseudowhorls of 3 but also alter-
nate and opposite, entire, stipulate, mostly less
than 1 mm wide, 1- or rarely 3-veined. Inflores-
cence of one staminate and one carpellate flower.
Flowers monoecious, without perianth; staminate
with (2- )4( -8)-sporangiate anther; carpellate
with (1- )4-5( -8) carpels basally surrounded by a
membranous envelope; style less than 1 mm long,
stigma asymmetrically funnel-shaped. Endocarp
often spiny. Perhaps five spp., near cosmopolitan
distribution.
2. Pseudalthenia Nakai
Pseudalthenia Nakai, Ord. Fam. 213 (1943).
Vleisia Tomlinson & Posluszny (1976).
Annual, in seasonally dry watercourses. Leaves
distichous, entire, stipulate, usually more than
1 mm wide, usually 3-veined, to 6.5cm long, apex
pointed, base open, sheathing. Inflorescences
axillary, many-flowered, sympodial, first-order
branch terminating with male flower, followed
by several carpellate and occasionally additional
staminate flowers. Flowers monoecious, staminate
naked, with 8-sporangiate anther; carpellate with
a basal, tubular spathe; carpel 1; style ca. 1 mm
long; stigma obliquely funnel-shaped. Achene
elongated, asymmetrical; pericarp with numerous
warty protuberances; endocarp sclerotic. Only
one sp., Pseudalthenia aschersoniana (Graebner)
Hartog, restricted to S Africa.
3. Althenia Petit
Althenia Petit, Ann. Sci. Observ. 1: 451 (1829).
Annual or perennial, in brackish coastal waters.
Shoots dimorphic, rhizome creeping, sympodially
branched, rooted, scaly, erect shoots leaf-bearing.
Leaves distichous to diffuse-spirally arranged,
entire, stipulate, narrow, less than 1 mm wide,
usually 1-veined; apex pointed; base open, sheath-
473
ing. Inflorescences axillary, sympodial, complex
clusters. Flowers monoecious; staminate termi-
nating main axis of each flower complex, with
perianth of 3 minute scales and 2-sporangiate
anther; carpellate terminating higher order
branches, with perianth of 3 toothed tepals, 3-
carpellate; style long; stigma peltate, symmetri-
cal. Achenes elongated, cylindrical or somewhat
flattened; pericarp smooth, endocarp sclerotic,
with germination suture. Two spp., N Africa and
S Europe.
4. Lepilaena Drumm. ex. Harv.
Lepilaena Drumm. ex. Harv., Hook. J. Bot. 7: 58 (1855).
Annuals or perennials. Rhizomes creeping,
sympodially branched, scaly; erect shoots leaf-
bearing, lateral, from nodes of the rhizome. Leaves
distichous, entire, stipulate, narrow, less than
1 mm wide, usually 1-veined; apex truncate, emar-
ginate or rounded and toothed, base open, sheath-
ing. Inflorescence axillary, sympodial. Flowers
monoecious or dioecious; staminate solitary,
with 4- or 12-sporangiate anther, subtended by 3
minute perianth scales; carpellate with 3 carpels,
subtended by 4 membranous tepals; style long;
stigma expanded, broadly funnel-shaped or feath-
ery. Achenes asymmetric, smooth or tuberculate.
Four spp., Australia and New Zealand.
Selected Bibliography
Cox, P.A. 1988. Hydrophilous pollination. Annu. Rev. Ecol.
Syst. 19: 261-280.
Cox, P.A., Knox, R.B. 1989. Two-dimensional pollination in
hydrophilous plants: convergent evolution in the genera
Halodule, (Cymodoceaceae), Halophila (Hydrocharita-
ceae), Ruppia (Ruppiaceae), and Lepilaena (Zanni-
chelliaceae). Am. J. Bot. 76: 164-175.
Cronquist, A. 1981. See general references.
Dahlgren, R.T.M., Clifford M.T. 1982. See general references.
Haynes, R.R., Holm-Nielsen, L.B. 1985 [1987]. A generic treat-
ment of Alismatidae in the neotropics. Acta Amazonia 15:
153-193.
Haynes, R.R., Holm-Nielsen, L.B. 1987. The Zannichelliaceae
in the Southeastern United States. J. Arnold Arbor. 68: 259-
268.
Les, D.H., Haynes, R.R. 1995. Systematics of subclass
Alismatidae: a synthesis of approaches. In: Rudail, P.J.,
Cribb, P.J., Cutler, D.F., Humphries, C.J. (eds.) Monocotyle-
dons: systematics and evolution. Royal Botanic Gardens,
Kew, pp. 353-377.
Les, D.H., Philbrick, C.T. 1993. Studies of hybridization and
chromosome number variation in aquatic angiosperms:
evolutionary implications. Aquat. Bot. 44: 181-228.
Les D.H., Garvin, D.K., Wimpee, C.F. 1993. Phylogenetic stud-
ies in the monocot subclass Alismatidae: evidence for a
474 Zannichelliaceae/Zingiberaceae
reappraisal of the aquatic order Najadales. Mol. Phylogenet.
Evol. 2: 304-314.
Posluszny, U., Sattler, R. 1976. Floral development of Zanni-
chellia palustris. Can. J. Bot. 54: 6541-6662.
Posluszny, U., Tomlinson, P.B. 1977. Morphology and devel-
opment of floral shoots and organs in certain Zanni-
chelliaceae. Bot. J. Linn. Soc. 75: 21-46.
Singh, V. 1965. Morphological and anatomical studies in
Helobiae. II. Vascular anatomy of the flower of Potamo-
getonaceae. Bot. Gaz. (Crawfordsville) 126: 137-144.
Tomlinson, P.B. 1982. See general references.
Tomlinson, P.B., Posluszny, U. 1976. Generic limits in the
Zannichelliaceae (sensu Dumortier). Taxon 25: 273-279.
Vierssen, W. van 1982. The ecology of communities domi-
nated by Zannichellia-taxa in western Europe I. Character-
ization and autecology of the Zannichel/ia taxa. Aquat. Bot.
12: 103-155.
Vierssen, W. van eta!. 1982. On the pollination mechanism of pseudostem formed by the leaf sheaths. Leaves
some eurysaline Potamogetonaceae. Aquat. Bot. 14: 339- distichous, sometimes appearing tufted, in lower
347.
Zingiberaceae
K. LARSEN, J.M. LocK, H. MAAS,
and P.J.M. MAAS
Zingiberaceae Lind!., Key Bot.: 69 (1835), nom. cons.
Perennial, aromatic herbs; rhizome sympodially
branched, sometimes on stilt roots. Stems short,
poorly developed, mostly lengthened by erect
part of shoot usually without blade; leaf sheath
usually open; ligule present or absent; blade
entire, elliptic, sometimes linear or broadly
elliptic, glabrous or hairy, when young rolled up
from one side to the other; midrib prominent;
lateral veins in a pinnate-parallel arrangement;
petiole present or absent; pulvinus usually absent
(except Zingiber). Inflorescence terminal on the
leafy shoot, or terminal on a short, separate,
sheath-covered, leafless shoot arising directly
from the rhizome, lax or congested, cylindric or
fusiform, sometimes globose, with (1- )few-many-
flowered, sometimes bracteolate cincinni axillary
to the bracts of the inflorescence and then repre-
senting a thyrse, or rarely a raceme or a spike;
bracteole obliquely adaxial to transverse, open
to the base ( = non-tubular), or tubular (but
may split early), or absent. Flowers epigynous,
bisexual, zygomorphic. Calyx tubular, usually
3-lobed or 3-dentate, sometimes split down 1 side.
Corolla tubular at base; lobes three, varying in
size and shape, the median posterior one often
larger than the other ones and cucullate. Only
the median posterior stamen of the inner whorl
functional; the other 2 stamens of the inner whorl
sterile and connate to form the petaloid labellum;
2lateral staminodes of the outer whorl petaloid or
inconspicuous, flanking the stamen or adnate to
the labellum; the median anterior member of
outer whorl always reduced; functional stamen
with long or short filament, sometimes filament
absent, anther dithecal, tetrasporangiate, thecae
opening introrsely by longitudinal slits, occasion-
ally dehiscing by pores, the connective often
apically prolonged in a crest and sometimes
also basally into spurs. Gynoecium 3-carpellate,
ovary in its first stages always 3-locular, when fully
developed either (incompletely) 3-locular with
axile placentation or !-locular with parietal or
 Zingiberaceae 475

basal placentation. Style terminal, very thin, The petiole is of varying length; in Zingiber it is
nearly always placed in a furrow of the filament short, fleshy, collenchymatous, and resembles the
and held between the thecae; stigma ± Wet, papil- pulvinus of the Marantaceae.
late, funnel-shaped, often ciliate, appearing on top The plane of insertion of the distichous leaves
of the anther. Nectaries 2, epigynous. Ovules ± varies from parallel to the length of the rhizome to
many, anatropous, bitegmic, and crassinucellar. transverse to it (Weisse 1932, 1933 ); these differ-
Fruit a dry or sometimes fleshy capsule, ences are constant for the tribes (Fig. 119). When
loculicidally dehiscent from apex to base by 3 the interval between the leaves is short, a distinct
valves, or irregularly dehiscent, or indehiscent, 2-ranked leaf arrangement is sometimes not well
smooth, ridged or verrucose; in some genera fruits marked (Kam 1982).
united in a syncarp. Seeds few to many, ovoid Leaf sheaths are usually open, but in Roscoea
to ellipsoid or subglobose, sometimes angular, and Cautleya closed leaf sheaths have been found
with an operculum next to the radicle, usually (Spearing 1977). The blades vary in size, the
with a prominent, commonly lobed or lacerate largest developing usually in the middle of the
aril; embryo straight, linear, central in the en- shoot. In shape they are elliptic, rarely broadly
dosperm; perisperm abundant, starchy with large, elliptic or linear. Particularly in the Alpinieae the
compound starch grains. Germination hypo- blades are asymmetric. One half of the blade is
geal. completely rolled around the other during devel-
A pantropical family of about 50 genera and opment (Kress 1990).
1300 species with the centre of diversity is S
and SE Asia. VEGETATIVE ANATOMY. (Mainly from Tomlin-
son 1969, where further information is provided.)
VEGETATIVE MoRPHOLOGY. The rhizome is sym- The rhizome usually has suberised surface
podial, varying in size and degree of branching; in layers; periderm is often developed. Rhizomes, as
the genus Curcuma, among others, the rhizome is
repeatedly branched. In some of the smaller
species it is hardly developed, while in most gen-
era it is a thick and fleshy storage organ provided
with distichous, scalelike leaves and roots. Genera
with a small rhizome often develop globose or
fusiform starch-filled tuberous roots, as in Globba,
Siphonochilus, Kaempferia, Roscoea, or the roots
are ending in tubers (Curcuma, Gagnepainia).
In some rainforest taxa (ssp. of Geostachys,
Amomum and Hornstedtia), the rhizome is placed
well above the ground and supported by stout,
unbranched stilt roots. In some species of
Scaphochlamys it ascends obliquely and is sup-
ported by slender stilt roots. Stilt-rooted species
are mainly found in humid evergreen forests with
a thick layer of dead leaves.
Usually, the leafy shoots terminate in an in-
florescence or, less commonly, inflorescence and
leaves appear on separate shoots. According to
Ridley (1899} plants growing in dense jungle as a
rule have the flowers close to the ground, while
those in more open spaces have them on the ends
of leafy stems. The shoots vary considerably in
size and in some rainforest species (Etlingera,
Alpinia) can be up to 8 m tall. The erect vegetative
stems are always unbranched. The true vegetative
stem is short and thin, composed mainly of thin-
walled cells, and the upright vegetative shoots Fig. 119A,B. Zingiberaceae. Plane of distichy of leaf shoots
are largely pseudostems (false stems) formed by in relation to rhizomes. A Parallel (Hedychieae, Globbeae,
intercalation of leaf sheaths. Zingibereae). B Transverse (Alpinieae). (Smith 1985)
476 Zingiberaceae
well as roots, store large quantities of starch. The
grains are nearly always simple, often flattened,
sometimes irregular in outline; the hilum is
usually excentric. Globose or ellipsoid grains are
found in Elettaria.
The roots develop 1 or several layers of thin-
walled exodermis, below which in some genera
(Elettaria, Hornstedtia) a cylinder of thin-walled
fibres is present. The vascular cylinder is polyarch,
and in the tuberous roots of Globba, Kaempferia,
and Roscoea, which have a wide, starch-filled
cortex, the stele is of the same diameter as in non-
tuberous roots. Vessels are mostly restricted to the
roots, rarely also present in the stem; they have
oblique scalariform perforation plates.
The stem is usually rather short and divided into
a wide cortex and central part separated by a
smooth fibrous cylinder. Vascular bundles of the
cortex are placed close together, each with a
massive, fibrous sheath; those of the central
cylinder are more scattered and with less devel-
oped sheaths.
The leaves are always dorsiventral. They have
a thin cuticle; rarely (Globba spp., Kaempferia
rotunda) the epidermis is minutely papillose.
The stomata are mainly present on the abaxial
side of the leaves. Each stoma is surrounded by a
pair of lateral and often also by a pair of terminal
subsidiary cells with even divisions. The guard
cells are at the same level as the epidermis, inner
ledges smaller than the outer ones, or absent. A
hypodermis of 1 layer of large, thin-walled,
colourless cells is usually present on both surfaces,
sometimes containing strands of hypodermal
fibres. Expansion cells, causing the young leaf to
unroll, develop by enlargement of hypodermal
cells, most commonly above larger veins of the
leaf blade. The mesophyl usually includes a 1-3-
layered palisade parenchyma and very loose
sponge parenchyma. One arc of air canals is found
in the midvein from the base to half-way towards
the apex.
Microscopic hairs are common, 1-cellular and
generally unbranched, except for the 2-celled hairs
on the rhizome of Curcuma and the characteristic
furcate to stellate, branched but 1-cellular hairs
of some species of Renealmia, Riedelia and
Rhynchanthus (Maas 1977).
Crystals of calcium oxalate as prismatic, rhom-
boid or tubular bodies are common in the assimi-
lating tissue as well as in the hypodermis of the
leaf blade; raphide sacs are lacking. Crystal sand is
sometimes found (Roscoea) and large solitary
bodies of chrystalline material occur in hypoder-
mal cells in the leaf blade in Globba.
Silica in the form of sand or as small subglobose
bodies is common in the epidermis of the lamina;
sometimes both forms occur in the same epider-
mis cell of the lamina. Silica bodies in unmodified
epidermal cells accompanying the main veins are
almost restricted to the Alpinieae. Silica cells con-
taining large solitary silica bodies are also known
to occur; they resemble the true stegmata found in
related families.
Condensed tannin is common in cells of the
ground parenchyma in all parts; in the leaf blade,
tannin cells are often larger than normal meso-
phyll cells. In Zingiber elongated tannin cells are
arranged in files and resemble articulated latici-
fers. Sometimes tannin is also deposited in inter-
cellular spaces.
Oil cells are characteristic of the family; they
are isodiametric and have suberised walls. They
are abundant in the parenchyma of all parts of
the plant and very common in the abaxial leaf
epidermis.
Epicuticular wax deposits are mostly absent;
rarely irregularly oriented platelets or compound
rodlets are found (Frolich and Barthlott 1988).
The sieve-tube plastids contain cuneate, pro-
teinaceous inclusions.
INFLORESCENCES. The inflorescence is usually a
thyrse, the bracts of which (sometimes called
primary bracts) each subtend a short cincinnus. In
some taxa the cincinni are reduced to a single
flower, so that racemes or spikes result. Unlike
most other monocotyledons, the bracteoles (sec-
ondary bracts) are not positioned adaxially, but
face at right angles to the primary bract, with the
subsequent bracteoles facing alternately in two
directions at right angles to each other, hence the
cincinnate partial inflorescences. In Scapho-
chlamys the bracteole is immediately facing the
primary bract, entire and open to the base, and
enclosing the whole cincinnus with its bracteoles
and flower buds. In Kaempferia the arrangement
is similar, except that only 1 flower is present, and
the 2-keeled bracteole becomes 2-lobed or even
bifid (separated into 2 narrow bracts). Often
the bracteoles are tubular, particularly in the
Alpinieae, while the open form (split to the base or
non-tubular) is typical of the other tribes. In some
species of Alpinia, Geostachys, and Elettaria the
tubular bracteoles are so large that they enclose
the whole cincinnus up to a late stage of its devel-
opment (Holttum 1950).
While leaves and bladeless sheaths on the pe-
duncle are distichously arranged, fertile bracts are
usually spirally placed on the rhachis forming the
 Zingiberaceae
inflorescence. In some genera the inflorescence is
congested and the bracts are closely imbricate.
Flowers can also be subtended by 2-ranked,
distichous bracts especially in much elongated
and often branched inflorescences (Elettariopsis,
Cyphostigma, Elettaria) or in a short, congested
inflorescence as in Distichochlamys, Caulokaem-
pferia, Haplochorema and Boesenbergia.
Occasionally, the bracts are exceptionally large
and in Camptandra and Pyrgophyllum they de-
velop a foliaceous appendage. In Curcuma and
some species of Scaphochlamys, Boesenbergia and
Zingiber the bracts form pouches that hold water;
the partial inflorescences are then constantly em-
bedded in an aqueous mucilage. In Zingiber the
fruits develop and ripen while embedded in the
slimy substance. A similar situation is found in
most species of Plagiostachys, where the bracts
decay and the whole outer part of the inflores-
cence becomes slimy. Other biologically impor-
tant modifications of the inflorescence have been
described by Holttum (1950).
FLORAL MORPHOLOGY. Flower structure is basi-
cally the same throughout the family and de-
scribed above. The morphological nature of the
labellum has long been controversial (see Holttum
1950), but studies by Rao et al. (1954) have left
little doubt that the labellum is formed from
the 2 united antero-lateral staminodes of the
inner whorl, and that the median anterior stamen
of the outer whorl is entirely suppressed. The
homology of the various lateral appendages of the
labellum with the (postero- )lateral staminodes
from the outer whorl rests on more shaky ground
(Burtt 1972), although for the tribe Hedychieae
this is more easily acceptable than for the
Alpinieae.
Many different parts of the flower can be
connate, such as:
- calyx and corolla together forming a beak on
the ovary (Cyphostigma pulchellum, Elettaria
longituba);
- corolla and labellum are connate in Vanover-
bergia, Riedelia, Aframomum, Hornstedtia,
Zingiber, Amomum, Cautleya;
- corolla and staminodes are connate in Globba,
Cautleya;
- filament and labellum are connate in a distinct
tube above insertion of the petals in Geocharis,
Pommereschea, Etlingera, Aulotandra, Stadio-
chilus, Globba;
- filament and lateral staminodes are connate in
Mantisia;
477
staminodes and labellum are connate in
Siphonochilus.
The thecae usually dehisce longitudinally and
introrsely, but in Etlingera and Amomum, thecae
are sometimes dehiscing in the upper half only,
whereas only the lower half of the anther dehisces
in species of Aframomum. In some species of
Elettaria the anthers are dehiscing by pores; the
same was observed for some spp. of Boesenbergia
by the author (K. Larsen).
One of the more important taxonomic charac-
ters is the elaboration of the connective. Generally,
this is extended apically to form the so-called
anther crest. The presence or absence of a crest is
usually a good generic character. The form of the
crest is useful in distinguishing species and varies
from large, flat, coloured and petaloid in Kaemp-
feria to absent or very small in Boesenbergia.
Basal extensions of the connective are found in
Curcuma. Versatile anthers are found in Cautleya,
Roscoea, Camptandra, and Curcuma. The position
of the style in a furrow in the filament and between
the thecae placing the stigma on top of the anther
is unique for the Zingiberaceae and Costaceae. A
character of promising importance is the groove
formed by the inner surface of the corolla tube to
envelope and hold the style. This character has,
among others, been found in Stadiochilus.
The inferior ovary in its early stages of
development seems always to be trilocular. In the
Globbeae it develops into a unilocular ovary with
parietal placentas. In Kaempferia, Caulokaemp-
feria, Siphonochilus and Boesenbergia, all belong-
ing to tribe Hedychieae, there is a tendency
towards the reduction of the septae. The ovules
then become basal on a short columnar placenta
or, in some species of Boesenbergia, only the lower
part of the ovary is trilocular. In one species of
Scaphochlamys the number of ovules is reduced
to one.
The epigynous glands at one time had been
equated with stylodes or even staminodes;
through the work of Rao (1963) it has become
clear, however, that they are mere outgrowths of
the upper surface of the ovary producing nectar
(for further discussion and illustrations see Burtt
1972).
EMBRYOLOGY. The very many studies that have
been carried out in this field have been sum-
marised by Davis (1969) and Johri et al. (1992).
The anther tapetum is secretory as well as
amoeboid; the division of the pollen mother cell
successive, and mature pollen is always binucle-
478 Zingiberaceae

ate. Embryo-sac development is of the Polygonum Mahanty (1970) suggested that 12 is the original
type, as usual in Zingiberales, and endosperm basic number for the family and presumably
development is Helobial (as in Costaceae, but in derived from 11, which may be the basic number
contrast to all other families of the Zingiberales, in for Zingiberales.
which it is Nuclear). Embryogeny is of the Asterad
type. PoLLINATION. Flowers last only 1 day, they are
tubular and contain nectar. Kato (1996) studying
PoLLEN MoRPHOLOGY. The pollen grains are forest understorey species in Sarawak, found birds
spheroidal, subspheroidal, ovoid or rarely ellip- of the genus Arachnothera (Nectariinae, spider-
soid, and often inaperturate, but sometimes hunters) to be the pollinators of Etlingera bre-
sulcate in Zingiber. The exine is thin, hardly resis- vilabis and Hornstedtia tomentosa, both with red-
tant to acetolysis, and does not provide mechani- flowered, basal inflorescences. Small traplining
cal strength for the pollen grain, but the intine is bees of the Halictidae (Nomia and Trinchostoma)
well developed and lamellated, consisting of sev- were observed on Amomum polycarpum and on
eral different layers. The sculpture of the exine is three species of Boesenbergia. Medium traplining
psilate, verrucate, echinate, scabrate, rugulate or bees of the genus Amegilla are the probable polli-
striate. Liang {1988) and Mangaly and Nayar nators of Amomum gyrolophos and Plagiostachys
(1990) found the following pattern: the Globbeae crocydocalyx. Schumann (1904) already suggested
(only Globba studied) have shortly echinate bees to be also active in pollinating species of other
pollen; in both the Alpinieae and Hedychieae the ground-flowering genera like Elettaria, Geostachys
pollen is either long-spined or psilate; within the and Elettariopsis.
tribe Zingibereae (i.e. the genus Zingiber) pollen In Roscoea purpurea Lynch (1882) found the
can be either spheroidal with rugulate ornamenta- anther to be versatile with basal connective out-
tion or ellipsoidal and striate, closely following growths (spurs) which make a visiting insect force
subgeneric limits. Variation thus by and large the anther forward so that pollen is deposited
seems to follow taxonomic borders, although the on its back, a mechanism corresponding to that
tribes, as far as is known, encompass more than found in Salvia. A similar mechanism has
one pollen type. been found in Curcuma (Schumann 1904) and
Camptandra (Holttum 1950).
KARYOLOGY. Cytological data for the family are In Brazil, Hedychium coronarium and most
abundant (see, e.g. Sharma and Bhattacharya species of the Asiatic genus Hedychium are,
1959; Mahanty 1970; Beltran and Kam 1984; and according to Muller (1890) and Kiinckel
the summary given by Chen 1989). The four tribes d'Herculais (1910), pollinated by hawk moths
are to some extent characterised by their karyo- (Sphingidae). However, their long, straight and
logy. The Zingibereae with the single genus stiff proboscis sticks in the narrow, curved corolla
Zingiber has the single basic chromosome number tube, and they are caught by the vacuum created
n = 11 and almost all species are diploids, with the after sucking the nectar. In the process of freeing
exception of cultivars from Japan and China. The themselves, they pollinate the flowers before
Globbeae are more complex with basic numbers n dying. Pollination by butterflies has also been
= 8 and 10. The genus Globba is most diverse, and observed in Hedychium by the same authors.
not only high polyploids but also aneuploids have In Renealmia 2 floral types are described by
been found. The Hedychieae are even more Maas (1977). In species with tubular flowers,
complex, as the basic numbers n = 10, 11, 12, 13, reddish bracts and copious nectar production
14, 17, 21 and 25 have become known. Polyploidy hummingbird pollination has been observed.
has been demonstrated in several genera, such as In another group of species with flowers with a
in Kaempferia where triploids, tetraploids and labellum composed of a tube and a horizontally
pentaploids occur. The Alpinieae are rather uni- spreading limb, usually green bracts, and faintly
form with basic numbers n = 12 and 11, of which developed nectaries, bee pollination is suggested.
the first is by far the commonest number. In this
tribe polyploidy is a common feature, and in FRUIT AND SEED. Fully dehiscent capsules, in
Plagiostachys octoploidy has been found. The two which 3 valves, separate from apex to base and
genera Rhynchanthus and Pommereschea, tradi- spread apart, seem to exist only in a minority of
tionally included in the Alpinieae, are based on n cases. They are known from Hedychium, Zingiber
= 11 and may be wrongly placed; at least the latter (where the dehisced fruit is still enclosed by large
seems to belong to the Hedychieae. bracts), Roscoea and Globba. In Hornstedtia the
 Zingiberaceae
capsules are reported to break irregularly in their
basal parts, while still ± enclosed by the involucra!
bracts (Holttum 1950). Sometimes the capsule
is indehiscent, or fleshy, as in Aframomum (berry)
and Etlingera (fleshy syncarp).
The seeds of Zingiberaceae are black to brown,
enveloped by a thin, sometimes sweet aril which
is lacerate or lobed, and originates from the
funiculus or the base of the ovule. A patch of
sclerenchymatous tissue develops around the mi-
cropyle and functions as the separating layer of an
operculum in most Alpinieae, Globba and Zingi-
bereae; it seems to be absent in Hedychieae. A
hypostase is often developed, but there are excep-
tions. In the mature seed the seed coat is testal, i.e.
it is formed only by the outer integument. The
outer epidermis of the testal layer consists of
narrow, tangentially elongated, fibriform cells,
which sometimes are sclerotic. The inner testal
epidermis is either thin-walled, as in Globba and
Hedychium, or sclerotic and U-shaped in cross-
section, as in Alpinieae and Zingibereae; it regu-
larly contains silica.
The main storage tissue is the perisperm, which
contains large quantities of starch. The grains are
highly compound in Hedychieae and Zingibereae,
but consists of simple concentric minute grains of
less than 2.5 mm diameter in Alpinieae.
The endosperm is in general more scanty than
the perisperm. It generally contains aleurone only.
It is starch-free in all Alpinieae studied.
The embryo is cylindrical and slender to stout
and barrel-shaped, occasionally hooked, capitate,
clavate or pear-shaped, and usually reaching the
extension of the perisperm, rarely shorter.
Cells containing ethereal oils are frequent in the
embryos but have not been found in two genera of
the Alpinieae and in Globba; in the embryo of the
latter genus, tanniniferous idioblasts have been
observed.
Based on the study of seed characters of ten
genera, the tribes can be characterised in the
following way (from data compiled by Johri et al.
1992 and unpubl. observ. by H. Huber).
1. Endotesta lacking U-shaped cells; endosperm containing
starch 2
- Endotesta cells U-shaped. Operculum mostly present 3
2. Operculum present; starch grains of perisperm simple
Globbeae
- Operculum absent; starch grains of perisperm compound
Hedychieae
3. Starch grains of perisperm compound; endosperm and
embryo containing starch Zingibereae
- Starch grains of perisperm simple, concentric; endosperm
and embryo without starch Alpinieae
479
DISPERSAL. Our knowledge of dispersal is as
sparse as is that of pollination. Hedychium has
fleshy capsules, the valves of which roll back on
opening and expose the orange inner wall of the
capsule and the deep red aril of the seeds. This
strongly suggests bird dispersal. In many species
the arillate seeds are shed onto the ground and
dispersal by ants, or mice or squirrels (Ridley
1899) is suggested. The same mode of dispersal is
suggested by Maas (1977) concerning Renealmia.
The inflorescence is partly or almost completely
buried in species of several genera, notably in
Etlingera, Elettariopsis and Elettaria. In some
cases the fruit itself develops below the surface of
the ground.
In Caulokaempferia the author (K. Larsen) has
observed the green, fleshy capsule opening by a
broad lateral slit, through which the small seeds
are dispersed by rain splash and later by running
water. In some Amomum and Etlingera species the
single fleshy fruits adhere to each other to form a
collective fruit (syncarp) the shape of a small pine-
apple. The fruit wall is acid and juicy and seeds
may be spread by birds or small mammals, but
there are no observations on this. In Africa the
indehiscent fleshy fruits of Aframomum are eaten
by primates and elephants (Nishida and Vehara
1983).
VEGETATIVE REPRODUCTION. Most Zingibera-
ceae can be propagated by fragmenting the
rhizome. Several Globba species, maybe most,
produce bulbils either in the axils of the lower
bracts instead of flowers, or further down the
stem, or even on separate, thin shoots. A few
species, such as G. marantina and G. aphanantha,
rarely produce flowers but regularly bulbils. With
increasing ploidy levels, sexual reproduction is
weakened or even lost.
PHYTOCHEMISTRY. Several genera have been in-
vestigated and an updated survey is given by
Hegnauer (1986). Very characteristic for the
family are the oil cells containing ethereal oils.
Phenylpropanoids are found in Zingiber and
Curcuma species and in Aframonum melegueta
(Grains of Paradise) and the related curcumins
(diarylheptanoid) in Curcuma, Alpinia and
Zingiber. Aromatic acids and flavonoids are iso-
lated from several genera, as well as proan-
thocyanins and condensed tannins. Chinones
were found in stems of Aframomum giganteum.
Steroid saponins, common in the Costaceae, have
only been found in Hedychium coronarium.
Diterpenes and triterpenes are probably not rare;
480 Zingiberaceae
they have been isolated from several genera. Lyco-
pene and beta-carotene are present in the red aril
of Hedychium species. Extracts from rhizomes
of Curcuma and Alpinia have shown strong
mycotoxic qualities.
AFFINITIES. The Zingiberaceae are the largest
family within the Zingiberales. They belong to a
group of four families (Costaceae, Zingiberaceae,
Cannaceae and Marantaceae) that are charac-
terised by the possession of only one functional
stamen, while the other stamens are represented
by petaloid staminodes. Among these families
the Costaceae are the closest relative of the
Zingiberaceae. A comprehensive phylogenetic
analysis of the Zingiberales has been undertaken
by Kress (1990), while Clark et al. (1993) have
studied the relationships on the basis of the rbcL
gene.
The arrangement of the 46 genera in 4 tribes,
Alpinieae, Hedychieae, Globbeae and Zingibereae
is not yet final. The place of the genera Rhyn-
chanthus, Pommereschea and Nanochilus in the
Alpinieae is still questionable. Stadiochilus is
placed as incertae sedis.
DISTRIBUTION AND HABITATS. The family is
widely distributed in the tropics with the greatest
diversity and number of genera and species in
the Asian tropics. In the neotropics the only
indigenous genus is Renealmia, and in Africa
four genera are represented, Aframomum,
Aulotandra (mainly in Madagascar), Renealmia
and Siphonochilus. In Asia the family reaches the
subtropical region in Japan at about 33°N, and the
southern limit is in SE Africa (Natal) at about 27°S;
in the Himalayas Roscoea reaches up to an altitude
of 4800m.
The majority of the species are associated with
tropical forests where they play an important role
in the herbaceous layer in places in which light is
not too sparse. The richest diversity is found in
humid tropical lowlands where species-rich
genera such as Alpinia, Amomum and Etlingera
are often dominant, but many endemics, mainly
belonging to genera of the Hedychieae and
Globbeae, are also found in the mountain forests
of the SE Asian monsoon region. Drier forests
such as deciduous dipterocarp forests also
harbour many species mainly of the Hedychieae,
e.g. Kaempferia, Curcuma. Some species of
Hedychium, Curcuma and other genera occur in
temporarily inundated habitats. A few species of
Hedychium are epiphytes in montane forests and
in lowland swamp forests, and some species of
Caulokaempferia grow together with mosses in
shady places on rocks or on boulders in streams.
In Africa species of Siphonochilus occur in
deciduous woodlands and grasslands with a long
dry season. Roscoea is a montane genus found in
the Himalayas from Kashmir eastward to Tibet
and Sichuan in the north and Yunnan in the south.
It is unique among the Zingiberaceae for with-
standing exposure to the sun, to harsh conditions
and to high elevations.
In spite of their preference for moist forests and
humid habitats in the mountains, no waterplants
are found among the Zingiberaceae.
PALAEOBOTANY. Hickey and Peterson (1978) de-
scribed a fossil genus, Zingiberopsis, based on leaf
remains from the Upper Cretaceous/Eocene of N
America. Fruits and seeds of the fossil genus
Spiratospermum, which formerly were ascribed
to the Zingiberaceae, are now referred to the
Musaceae (Rodriguez-de la Rosa and Cavallos-
Ferriz (1994).
EcoNOMIC IMPORTANCE. The rhizomes of
Zingiber officinale (ginger), and of Curcuma
species (turmeric), are very important spices on
the world market. The seeds of Elettaria car-
damomum are equally important as a spice, while
those of some Aframomum species, particularly
A. melegueta, Grains of Paradise or Melegueta
pepper, are of secondary importance as a spice
and drug. Locally, however, a great many species
are used for different purposes: Amomum,
Aframomum, Alpinia and Kaempferia species as
condiment to give relish to food; Boesenbergia
species as spices; Hedychium and Alpinia galanga
for perfumes; Curcuma as a colouring agent. Re-
cently it has been found that fresh rhizomes of
Zingiber officinale contain compounds that re-
duce nausea and vertigo and seem to have a posi-
tive effect on patients with stomach ulcers. Several
species are also used locally as medicine and cos-
metics. Some are also used in tropical gardening,
e.g. Curcuma Hedychium and Globba species,
Alpinia purpurata and A. speciosa, and Etlingera
elatior. The Himalayan genus Roscoea is some-
times cultivated in gardens of the temperate zone.
KEY TO THE GENERA AND TRIBES
1. Style exserted well beyond the anther tip, the elongated
anther crest wrapped round the style (Zingibereae)
24. Zingiber
- Style not exserted beyond the anther; crest, if present,
never wrapped around the style 2
2. Filament usually long-exserted from the flower, arched as
 Zingiberaceae 481

a bow, the style forming the string of the bow; ovary 18. Bracteole tubular; labellum always 2-lobed; anther crest
unilocular with parietal placentation (Globbeae) 25 absent 19
- Filament usually short; iflong, then ovary always trilocular - Bracteole, if present, open to the base; labellum entire or
3 2-lobed to bifid; anther crest variously shaped 21
3. Lateral staminodes petaloid, usually free from labellum, 19. Bracts distichous 15. Distichochlamys
rarely connate to it but then pseudostem very short - Bracts spirally arranged 20
(Siphonochilus and Cautleya); ovary mostly 3-locular, 20. Corolla tube long-exserted from the bracts; bracts some-
rarely incompletely 3-locular with basal placentation; times imbricate, tips never recurved 1. Hedychium
plane of distichy of leaves parallel to rhizome - Corolla tube usually not exserted from the bracts; bracts
(Hedychieae) 4 imbricating with recurved, Curcuma-like tips
- Lateral staminodes absent or represented by small teeth, 2. Hitchenia
rarely linear or petaloid appendages oflower part oflabel- 21. Bracts distichous; inflorescence flowering from apex to
lum, but then pseudostem well developed; ovary always base 22
trilocular; plane of distichy transverse to rhizome - Bracts spirally arranged; inflorescence flowering from base
(Alpinieae) 28 ~~9 M
4. Bracts laterally connate to each other in the lower part, 22. Plants with well-developed leafy pseudostems; corolla tube
forming pouches; anther versatile 6. Curcuma distinctly exceeding the bracts 10. Caulokaempferia
- Bracts not laterally connate to each other; anther versatile - Plants almost acaulescent, leaves radical; corolla tube not
mn~ 5 exceeding the bracts 23
5. Inflorescence surrounded by a bell-shaped involucre 23. Labellum usually concave and entire to emarginate
formed by one single bract enclosing cincinnus 18. Boesenbergia
16. Stahlianthus - Labellum broad and spreading, emarginate to 2-lobed
- Inflorescence not surrounded by a bell-shaped involucre 17. Haplochorema
6 24. Bracteole open to the base and bifid; flowers violet or white
6. Bracts large, concave, their margins basally adnate to the 13. Kaempferia
inflorescence axis, appearing tubular in lower half 7 - Bracteole open to the base and entire; flowers white with
- Bracts free 8 red or yellow marks 12. Scaphochlamys
7. Anther basally spurred, versatile; bracts 1 per inflores- 25. Filament short, not exceeding the petals 23. Hemiorchis
cence, without apical foliaceous appendage - Filament much exceeding the petals, arched as a bow 26
4. Camptandra 26. Labellum 3-lobed, central lobe very short with 2 gland-like
- Anther apically crested, not versatile; bracts 1-3 per inflo- basal swellings; flowers greenish 22. Gagnepainia
rescence, with apical foliaceous appendage - Labellum 2-lobed to entire, flowers differently coloured
11. Pyrgophyllum 27
8. Inflorescence terminal on a separate leafless shoot 9 27. Lateral staminodes elliptic, inserted at about the same
- Inflorescence terminal on the leafy shoot 14 level as the petals; labellum connate to the filament
9. Bracteole absent 10 20. Globba
- Bracteole present 12 - Lateral staminodes very narrow, connate to the filament
10. Lateral staminodes connate to the labellum, their united forming lateral appendages; labellum inserted at about the
basal part enfolding the filament (Africa, Madagascar) same level as the petals 21. Mantisia
14. Siphonochilus 28. Inflorescence terminal on a separate leafless shoot 29
- Filament not enfolded this way 11 - Inflorescence terminal on the leafy shoot or plant acaules-
11. Anther versatile, basally spurred 8. Roscoea cent and inflorescence arising in the centre of a tuft of
- Anther not versatile, not spurred 5. Haniffia radicalleaves 43
12. Flowering shoots appearing before the leaves 29. Calyx caducous, falling with the flower; bracts small and
13. Kaempferia caducous or absent, lateral staminodes absent
- Flowering shoots appearing together with the leafy shoots 29. Riedelia
13 - Calyx persistent; bracts present 30
13. Anther crest conspicuous, reniform 19. Parakaempferia 30. Inflorescence congested, forming a compact head; axis
- Anther crest absent or very small 18. Boesenbergia covered by imbricating sterile or fertile bracts 31
14. Anther versatile with 2long basal spurs held at right angles - Inflorescence lax; axis visible; bracts not imbricating;
to the thecae 15 sterile bracts never present 35
- Anther not versatile, not spurred, but occasionally with 31. Capsule beaked, fleshy, usually red when ripe; anther crest
short, basal appendages 17 usually conspicuous, 3-lobed (Africa and Madagascar)
15. Leaf sheaths tubular; flowers purple to white with broad 39. Aframomum
cucullate dorsal petal 8. Roscoea - Capsule more or less globose, dry or fleshy; anther crest
- Leaf sheaths usually open; flowers yellow 16 variously shaped (Asia, Australia) 32
16. Bracts often brightly red-coloured; ovary 3-locular with 32. Labellum and filament connate into a distinct tube above
axile placentation; lateral staminodes, lateral petals and the insertion of the petals; anther crest absent
labellum basally connate to each other 7. Cautleya 36. Etlingera
- Bracts green; ovary incompletely 3-locular with basal pla- - Labellum and filament not connate; anther usually crested
centation; lateral staminodes, petals and labellum free 33
from each other 9. Paracautleya 33. Leaves 1-5, in radical tufts or on short pseudostems;
17. Bracts minute; bracteole absent; each theca with an apical, anther crest prominent 42. Elettariopsis
long-acuminate crest 3. Siliquamomum - Leaves many, on well-developed pseudostems 34
- Bracts conspicuous; bracteole usually present; anther crest 34. Inflorescence enclosed by an involucrum of several rigid,
when present not as above 18 sterile bracts; corolla tube more than twice as long as the
labellum 38. Hornstedtia
482 Zingiberaceae

- Inflorescence not enclosed by sterile bracts; corolla tube 51. Labellum concave; stigma small, geniculate; calyx short,
about as long as the labellum 37. Amomum campanulate. New Guinea, Australia
35. Inflorescence prostrate, sometimes almost subterranean 31. Pleuranthodium
with only the tips of the flowers visible 36 - Labellum spreading, pendulous, erect or concave; stigma
- Inflorescence erect or decurved 40 variously shaped; calyx tubular 52
36. Flowers resupinate; anther crest about as wide as the 52. Calyx caducous, falling with the flower. New Guinea
labellum. Sri Lanka 43. Cyphostigma 29. Riedelia
- Flowers upright; anther crest narrower than the labellum - Calyx at least partially persistent on fruit 53
37 53. Labellum bifid, lobes linear; bracts conspicuous; each
37. Bracteole open to the base; anther crest petaloid subtending 1 flower; bracteole absent. Philippines
42. Elettariopsis 28. Vanoverberghia
- Bracteole tubular 38 - Plants with a different combination of characters
38. Labellum and filament connate into a tube above the 25. Alpinia
insertion of the petals; bracts each subtending 2 flowers
41. Geocharis
- Labellum and filament not connate into a tube above the
insertion of the petals; bracts each subtending 1-many Tribes and Genera of the Zingiberaceae
flowers 39
39. Calyx tubular, split down l side; labellum entire, obovate. I. Tribe Hedychieae Petersen (1889).
Asia 45. Elettaria
- Calyx tubular, not split; labellum 3-lobed. America, Africa Small or medium-sized herbs with usually well-
26. Renealmia
40. Labellum bifid; filament often with lateral, subapical teeth developed, horizontal rhizome. Plane of distichy
41. Geocharis of leaves parallel to rhizome. Lateral staminodes
- Labellum not so divided; filament without subapical teeth usually well-developed, petaloid, usually free from
41 labellum. Style not exserted beyond anther. Ovary
41. Base of labellum and filament forming a distinct tube
mostly 3-locular, or incompletely 3-locular with
above the petals; anther crest petaloid (Madagascar,
Cameroon) 40. Aulotandra basal placentation.
- Base of labellum and filament not connate; anther crest, if
present, not petaloid 42
42. Leaves in radical tufts; bracteole enclosing a whole 1. Hedychium J. Konig Fig. 120
cincinnus. Asia 44. Geostachys
Hedychium J. Konig in Retz., Obs. Bot.3: 73 (1783).
- Leaves on well developed pseudostems; bracteole not
Brachychilum (R. Br. ex Wall.) Petersen (1893).
enclosing a whole cincinnus. Africa and America
26. Renealmia
43. Neotropical and African plants 26. Renealmia Terrestrial or epiphytic herbs with long, robust
- Asiatic plants 44 rhizome. Inflorescence terminal on the leafy
44. Leaves cordate to subsagittate; labellum basally connate to shoot; bracts broad and densely imbricate, or
the very long, linear (G/obba-like) filament. Myanmar, N
Thailand 35. Pommereschea
narrow and leaving the inflorescence axis visible,
- Leaves and labellum not as above 45 cincinni several-flowered, rarely flowers single;
45. Inflorescence apparently lateral, breaking through the leaf bracteole tubular; flowers white, yellow or red;
sheaths, often mucilaginous; labellum not showy calyx tubular, unequally dentate; corolla tube
27. Plagiostachys usually much longer than calyx and bracts, lobes
- Inflorescence rarely pushed out laterally from the leaf
sheaths; if so, then never mucilaginous and labellum narrow, reflexed; lateral staminodes about as long
showy 46 as petals, but broader and spreading; labellum
46. Labellum reduced, less than 1 em long 47 usually conspicuous and pendulous, emarginate
- Labellum well developed 48 to 2-lobed; filament usually much longer than the
47. Lateral staminodes absent. Sino-Burmeses
anther, linear, thecae elongated, crest absent;
33. Rhynchanthus
- Lateral staminodes petaloid, over 2 em long. Sumatra ovary 3-locular. Capsule trigonous, globose or
32. Nanochilus ellipsoid, dehiscent; seed with red, lacerate aril.
48. Labellum held erect, encircling the exserted filament. 2n = 34, 51, 68. About 50 spp., widely distributed
Myanmar 46. Stadiochilus in S and SE Asia; most diverse in the southern
- Labellum sometimes erect, if so then filament not exserted
49
tropical Himalaya region from N India to S China,
49. Flowers to 12cm long; corolla tube very slender, about but also rich in species in the Malesian region.
twice as long as the calyx. Philippines 34. Leptosolena
- Flowers not exceeding 6 em; corolla tube rarely long
exserted from the calyx 50 2. Hitchenia Wall.
50. Labellum much shorter than the petals, erect; bracteoles
absent. Borneo 30. Burbidgea Hitchenia Wall., Trans. Med. Soc. Calcutta 7: 215 (1835).
- Labellum usually exceeding or about the same length as
the petals, if shorter, then plants with tubular bracteoles Rhizomatous herbs. Inflorescence terminal on the
51 leafy shoot; bracts subcoriaceous, closely imbri-
 Zingiberaceae
Fig. 120A-F. Zingiberaceae-Hedychieae. Hedychium cylin-
dricum. A Inflorescence. B Flower of a two-flowered cincinnus
with primary bract and bracteole. C Bracteole. D Anther with
stigma in ventral, dorsal and lateral view. E Ovary with epigy-
nous glands and base of style. F Stigma. (Burtt and Smith
1972b)
eating with recurved apex (Curcuma-like) but free
from each other, terminal ones sometimes sterile
(coma), few-flowered; bracteole tubular; flowers
purple, yellow, or white; calyx short, tubular, mi-
nutely dentate; corolla tube slender, much longer
than calyx; lateral staminodes from minute to as
long as petals; labellum conspicuous and pendu-
lous, orbicular to spathulate, emarginate to bifid;
filament as long as labellum, thecae separate on a
broad connective, crest absent; ovary 3-locular.
Capsule trigonous, tardily dehiscent; seeds aril-
late. 2n = 42. Three spp., Himalayan region to
Myanmar. In need of careful re-examination.
3. Siliquamomum Baill.
Siliquamomum Bail!., Bull. Mens. Soc. Linn. Paris 2: 1193
(1895}; Loesener in E.-P., Natiirl. Pflanzenfamilien ed. 2,
1Sa: 573 (1930}
Rhizomatous herbs. Leaves few. Inflorescence a
lax raceme, terminal on the leafy shoot; bracts
483
minute, flowers borne singly on the inflorescence
axis; bracteole absent; flower colour unknown;
calyx tubular-campanulate, membranous, 3-
dentate, split down one side; corolla tube as long
as the lobes; lateral staminodes elliptic-obovate;
labellum conspicuous and pendulous, elliptic-
obovate with crenate margin; filament very short,
each theca with a long-acuminate crest; ovary 3-
locular. Capsule narrowly fusiform, more than
10 em long, torulose, dehiscence unknown; seed
with 2-lobed aril. Only one sp., S. tonkinense Baill.,
N Vietnam.
4. Camptandra Ridl.
Camptandra Rid!., J. Straits Branch R. Asiat. Soc. 32: 103
(1899}; Holttum, Gard. Bull. Singapore 13: 78-82
(1950}; R.M. Sm., Notes R. Bot. Gard. Edinb. 31: 205-207
(1972}.
Small herbs with slender stems bearing few leaves.
Leaves asymmetric. Inflorescence terminal on the
leafy shoot, consisting of a single cincinnus in the
axil of 1 large, concave bract, margins of bract
basally adnate to the inflorescence axis, some-
times with apical foliaceous appendage; bracteole
thin, open to the base and entire; flowers white or
lilac; calyx short, funnel-shaped; corolla tube slen-
der, longer than the bracts; lateral staminodes
almost circular, spreading; labellum conspicuous
and pendulous, bifid; filament short and broad,
anther versatile, crest absent, thecae basally
spurred; ovary 3-locular. Capsule ellipsoid, dehis-
cence unknown; seeds with translucent, lacerate
aril. Four spp., S Thailand, W Malaysia and
Sumatra.
5. Haniffia Holttum
Haniffia Holttum, Gard. Bull. Singapore 13: 123 (1950}.
Herbs up to 80 em with rather thin rhizome.
Stems few to many-leaved; leaves with short ligule.
Inflorescence on separate leafless shoots, pe-
duncle very short; flowers single or in few-
flowered cincinni, subtended by thin closely
spaced bracts; bracteole absent; flowers white
with a violet to white labellum; calyx tubular;
corolla tube long; lateral staminodes rather nar-
row, narrowly elliptic; as long as or longer than
petals; labellum obovate, 2-lobed; filament short,
anther with a short rounded crest; ovary incom-
pletely 3-locular. Capsule unknown. Two spp.
(one of which undescribed), Malay Peninsula and
S Thailand.
484 Zingiberaceae

6. Curcuma L.
Curcuma L., Sp. Pl.: 2 (1753), nom. cons.

Low, rhizomatous herbs; roots often with tubers.

Leaf blade often with violet bands on either side of
the midrib. Inflorescence ± cylindric, either ter-
minal on the leafy shoot or on a separate leafless
shoot; bracts large, broad, apex recurved, laterally
connate to each other in the lower part, forming F
pouches around the several-flowered cincinni, ter-
minal ones often sterile and differently coloured
(coma); bracteole open to the base; flowers white,
yellow, red, or violet; calyx short, unequally 3-
dentate and split half way on one side; corolla tube
adnate to the stamina! tube over its entire length;
lateral staminodes elliptic-oblong, inner edges
partly covered by the cucullate dorsal petal; G~
labellum conspicuous and pendulous, obovate;
filament short and broad, anther versatile,
connective sometimes with a short crest, basally
with 2 curved spurs; ovary 3-locular. Capsule
ellipsoid, liberating the seeds into the mucilage
of the bract pouches; seeds with lacerate aril.
2n = 42, 63, 66, 84. Probably about 50 spp. (but
uncertain how many truly wild, numerous
cultivars being described as species), widely
distributed in tropical Australasia from India to c
Queensland.
Fig. 121A-H. Zingiberaceae-Hedychieae. Cautleya spicata. A
7. Cautleya Hook.f. Fig. 121 Habit. B Flower with bract, lateral view. C Flower, front view.
D, E Dissected flower, D showing ovary, epigynous glands,
Cautleya Hook.f., Bot. Mag. ad t. 6991 (1888); Cowley, Kew corolla tube, anther and dorsal petal, E showing labellum with
Bull. 36: 747- 776 (1982). lateral staminodes, also apices of lateral petals visible. F An-
ther and style in lateral and front view. G Ovary, transversal
Small herbs with short rhizome. Leaves with dis- section. H Ovary (opened) and epigynous glands. (Burtt and
Smith 1972b)
tinct, 2-lobed ligule. Inflorescence terminal on the
leafy shoot; bracts often deep red, well spaced,
each subtending 1 flower; bracteole absent; Low herbs with short rhizome; roots tuberous,
flowers yellow; calyx tubular; corolla funnel- fusiform. Leaves with minute ligule, and tubular,
shaped, tube short, lateral petals basaly connate closed sheath. Inflorescence a raceme, terminal on
with labellum; lateral staminodes narrowly obo- the leafy shoot, or on a leafless shoot appearing
vate, erect, connate to base of labellum for about before the leaves are formed; bracts green, each
half of their length; labellum broadly spathulate, subtending 1 flower; bracteole absent; flowers pur-
emarginate or 2-lobed; filament short, anther ver- plish to pale blue or white; calyx tubular, split
satile, crest absent, basally with 2 long spurs at down 1 side; corolla tube slender, as long as or
right angles to the thecae; ovary 3-locular. Capsule longer than calyx, dorsal petal broad, cucullate;
globose, dehiscent; seeds with short, white aril. 2n lateral staminodes petaloid, erect, narrowly
= 34. Two spp., Himalayan region from N India obovate to circular or rhombic; labellum con-
and Nepal to S China and N. Thailand. spicuous and more or less pendulous, obovate,
entire to bifid, shorter than the petals; filament
short, anther versatile, basally with 2long spurs at
8. Roscoea Sm. right angles to the thecae, crest absent; ovary 3-
Roscoea Sm., Exot. Bot. 2: 97 (1806); Spearing, Notes R. Bot.
locular. Capsule ellipsoid to cylindrical, tardily
Gard. Edinb. 35: 217- 220 (1977); Cowley, Kew Bull. 36:747- dehiscent by 3 valves, or by I lateral slit; seeds with
777 (1982), rev. white, lacerate, fleshy aril. 2n = 24, 48. About 17
 Zingiberaceae 485

spp., mountains from the Himalayan region to S Medium-sized herbs with tuberous base. Inflores-
China and N Vietnam; the most "temperate" cence terminal on the leafy shoot; bracts 1-3,
genus of the family. large, concave, with apical foliaceous appendage,
their margins adnate to the inflorescence axis,
enclosing a 1-3-flowered cincinnus; bracteole
9. Paracautleya R.M. Sm.
open to the base or absent; flowers yellow; calyx
Paracautleya R.M. Sm., Notes R. Bot. Gard. Edinb. 35: 367, split down one side, 2-dentate; corolla tube twice
Fig. 1 (1977). as long as calyx; lateral staminodes narrow, about
as long as the petals; labellum conspicuous and
Acaulescent, rhizomatous herbs. Inflorescence ter- pendulous, emarginate or bifid, lobes obtuse; fila-
minal, arising in the centre of a tuft of radical leaves; ment short, broad, anther with a triangular crest;
bracts green, apex recurved, each subtending 1 ovary 3-locular. Capsule unknown. 2n = 42. Only
flower; bracteole absent; flowers yellow; calyx tubu- one sp., P. yunnanensis (Gagnep.) Wu & Chen,
lar; corolla tube short; lateral staminodes petaloid, China (Yunnan and Sichuan). Previously referred
oblong; labellum conspicuous and pendulous, to Kaempferia by Gagnepain, Camptandra by
about as long as the staminodes, obovate, 2-lobed; Schumann, and Caulokaempferia by Larsen and
filament broad, anther versatile, crest minute, ba- R.M. Smith; separated from these on anatomical,
sally spurred; ovary incompletely 3-locular, ovules cytological and morphological grounds.
basal. Capsule subglobose, probably irregularly
dehiscent; seeds with entire aril. Only one sp., P.
bhatii R.M. Sm., known only from the type locality 12. Scaphochlamys Baker
in India, Kanara Province.
Scaphochlamys Baker in Hook.f., Fl. Brit. Ind. 6: 252 (1892);
Holttum, Gard. Bull. Singapore 13: 82 (1950); R.M. Sm.,
Notes R. Bot. Gard. Edinb. 44: 207-210. 1987.
10. Caulokaempferia K. Larsen
Caulokaempferia K. Larsen, Bot. Tidsskr. 60: 166 (1964); Low herbs; rhizome creeping. Leaf blade asym-
K. Larsen & R.M. Sm., Notes R. Bot. Gard. Edinb. 31: 287- metric, often purple on lower side. Inflorescence
295 (1972).
congested or lax, from ovoid to elongate, terminal,
arising in the centre of a tuft of few, radical leaves;
Small, delicate herbs with well developed pseu-
bracts densely imbricate or spaced along the axis,
dostems, terrestrial or epilithic. Inflorescence
usually each subtending several flowers; bracteole
terminal on the leafy shoot, few-flowered, flower- open to the base and entire; flowers white with red
ing from apex to base; bracts distichous, narrowly or yellow marks; calyx tubular, split down 1 side;
elliptic, free or marginally adnate to the rachis, corolla funnel-shaped, tube narrow, much longer
each subtending 1-few flowers; bracteoles usually than the bracts; lateral staminodes broader than
split to the base and entire, or absent; flowers
petals, petaloid, spreading; labellum conspicuous
white or yellow; calyx tubular, not deeply split; and pendulous, spathulate to obovate, 2-lobed or
corolla tube long, narrow, straight or curved; lat- entire; filament broad, short, anther crest 3-lobed
eral staminodes large, petaloid, as long as petals; or entire, thecae with very short basal spurs;
labellum larger than the staminodes, ± orbicular, ovary 1-locular. Capsule subglobose, dehiscence
entire or 2-lobed, spreading to concave; filament unknown; seeds with white, lacerate aril. 2n = 26,
short, anther crest conspicuous, often recurved; 28. About 30 spp., endemic to the Malay Peninsula
ovary 3-locular. Capsule fleshy, irregularly dehis- from S Thailand southwards to Singapore.
cent by 1 broad slit; seeds with short hairs and no
aril. 2n = 20, 24, 42. About ten spp., southern
tropical Himalaya to S China, southwards on the 13. Kaempferia L.
Indochinese Peninsula to S Thailand; mostly in
damp environments, among moss, or on rocks Kaempferia L., Sp. Pl. 1:2 (1753); Kam Yee Kiew, Notes R. Bot.
Gard. Edinb. 38: 1-12 (1980).
in shade along streams.
Low herbs with short, fleshy rhizome; roots
11. Pyrgophyllum (Gagnep.) T.L. Wu & often tuberous. Stem with few (often 2) leaves,
Z.Y. Chen erect or appressed to the soil. Inflorescence short,
terminal, arising in the centre of the radical leaves,
Pyrgophyllum (Gagnep.) T.L. Wu & Z.Y. Chen, Acta Phytotax.
Sin. 27: 126 (1989); K. Larsen & R.M. Sm., Notes R. Bot. rarely on a separate leafless shoot, appearing be-
Gard. Edinb. 31: 287-295 (1972). fore the leaves; bracts each subtending 1 flower;
Kaempferia subg. Pyrgophyllum Gagnep. (1902).
486 Zingiberaceae
bracteole open to the base, usually bifid; flowers
white or violet; calyx tubular, unequally dentate;
corolla tube as long as or mostly much longer than
the calyx; labellum bifid, spreading, together with
the 2 petaloid staminodes often creating a mark-
edly quadrate appearance; filament very short,
crest large, entire or 2-dentate, reflexed; ovary in-
completely 3-locular. Capsule subglobose; seeds
with lacerate aril. 2n = 22, 24, 33, 36, 44, 45, 54.
About 40 spp., widespread in tropical Asia.
14. Siphonochilus J.M. Wood & Franks
Siphonochilus J.M. Wood & Franks in J.M. Wood, Natal Pl. 6
(3): t. 560, 561 (1911) and in Kew Bull. 1911: 274 (1911);
Burtt, Notes R. Bot. Gard. Edinb. 40: 369-373 (1982); Lock,
Fl. Trop. East Afr., Zingiberac.: 14-22 (1985), reg. rev.
Cienkowskia Schweinf. (1867), non Regel & Rach (1858).
Kaempferia subgen. Cienkowskia K. Schum. (1904).
Cienkowskiella Kam (1980).
Herbs from short vertical rhizome; roots often
tuberous. Leaves on very short pseudostems. In-
florescence a terminal, 1-many-flowered raceme
on a separate leafless shoot, often appearing be-
fore the leaves, and often subterranean with only
the tips of the flowers visible; bracts each subtend-
ing a single flower; bracteole absent; flowers white,
yellow, blue or purple; calyx tubular or turbinate,
3-lobed; corolla tube elongated; lateral sta-
minodes petaloid, connate to labellum, their
united basal part enfolding the filament; labellum
conspicuous, about as long as the staminodes, 2-
(3-) lobed; filament short, thecae basal, crest elon-
gated, petaloid; ovary 3-locular. Capsule fleshy,
subglobose, trigonous, dehiscence unknown;
seeds ?arillate. 2n = 26, 28, 42. About 15 spp.,
mostly in the seasonally dry parts of tropical Af-
rica and Madagascar.
15. Distichochlamys M.F. Newman
Distichochlamys M.P. Newman, Edinb. J. Bot. 52: 65 (1995).
Small herbs forming dense tufts of few-leaved
shoots. Inflorescence terminal, arising in the
centre of the radical leaves; bracts distichous, each
subtending a few-flowered cincinnus; bracteole
tubular, with 2 keels; flowers white and yellow;
calyx tubular, 3-lobed; corolla tube slender,
exserting the bracts; lateral staminodes elliptic;
labellum somewhat longer than the staminodes,
spreading, obtriangular, 2-lobed; filament short,
anther crest small, entire; ovary 3-locular, ovules
many. Capsule unknown. 2n = 26. Only one sp., S.
citrea M.F. Newman, Vietnam. Closely related to
Scaphochlamys, from which it is distinguished by
the distichous bracts and tubular bracteoles.
16. Stahlianthus Kuntze
Stahlianthus Kuntze, Rev. Gen. 2: 697 (1891).
Acaulescent herbs from short rhizome; roots fili-
form, terminated by small tubers. Leaves few,
often 2, blade asymmetric. Inflorescence terminal,
arising in the centre of a tuft of radical leaves,
consisting of a single several-flowered cincinnus
surrounded by 1, bell-shaped, 2-lobed bract;
bracteole minute, open to the base and entire;
flowers white or violet with yellow marks; calyx
tubular, 3-dentate; corolla white; lateral sta-
minodes petaloid, oblong, as long as or longer
than corolla; labellum conspicuous and pendu-
lous, spathulate, longer than staminodes, bifid;
anthers sessile, connective with or without a mem-
branous crest; ovary imperfectly 3-locular. Cap-
sule unknown. About six spp., Himalayan region
to S China, Hainan, Vietnam, Cambodia, Laos,
and N Thailand.
17. Haplochorema K. Schum.
Haplochorema K. Schum., Bot. Jahrb. Syst. 27: 331 (1899);
R.M. Sm., Notes R. Bot. Gard. Edinb. 44: 211-214 (1987).
Low plants with creeping rhizome. Leaves few,
sometimes only 1. Inflorescence terminal on the
leafy shoot, few-flowered; bracts distichous,
linear, each subtending a single flower, uppermost
flowers opening first; bracteole boat-shaped or
split to the base; flowers white; calyx membra-
nous, tubular, 3-dentate; corolla funnel-shaped,
tube narrow, not exserted beyond the bract;
labellum conspicuous, broadly deltoid, 2-lobed or
emarginate, spreading, together with the 2 peta-
loid staminodes creating a markedly quadrate
appearance; filament short, broad, anther with
petaloid, reflexed crest; ovary incompletely 3-
locular. Capsule ellipsoid, dehiscence unknown;
seeds with short, lacerate aril. Three or four spp.,
endemic to Borneo. Clearly related to Kaempferia
and Boesenbergia, but generic status in need of
further study.
18. Boesenbergia Kuntze
Boesenbergia Kuntze, Rev. Gen. 2: 685 (1891); Holttum, Gard.
Bull. Singapore 13: 106-117 (1950); Sirirugsa, Nat. Hist.
Bull. Siam Soc. 40: 67-90 (1990), reg. rev.
Gastrochilus Wall. (1829) non D. Don (1825).
Curcumorpha A.S. Rao & D.M. Verma (1974).
 Zingiberaceae 487

Small herbs from short, fleshy rhizome. Leaves
few arranged in tufts. Inflorescence terminal on
the leafy shoot, sometimes terminal on a separate
leafless shoot, uppermost flowers opening first;
bracts distichous (spirally arranged in Cur-
cumorpha), densely imbricating, each subtending
1 flower; bracteole open to the base and entire,
boat-shaped; flowers white, yellow or red; calyx
tubular; corolla tube slender, usually shorter than
the bracts; lateral staminodes oblong, petaloid;
labellum conspicuous and usually concave, entire
to emarginate; lateral staminodes and filament
adnate to base of labellum; anther crest absent
or very small; ovary incompletely 3-locular to 3-
locular. Capsule fleshy, subglobose, dehiscence
unknown; seeds with white, lacerate aril, often
longer than the seed. 2n = 20, 24, 36. About 60
spp., southern tropical Himalaya throughout SE
Asia, most species in the Indochinese monsoon
region and on Borneo.
Small herbs with short, creeping rhizome, roots
tuberous. Inflorescence terminal on the leafy
shoot, lax or congested; bracts green or variously
coloured, each subtending a few- to many-
flowered cincinnus, lower ones often bulbili-
ferous; bracteole open to the base; flowers white,
yellow, pink, orange, or purple; calyx turbinate to
campanulate, 3-lobed to 3-dentate; corolla tube
slender, long; labellum 2-lobed to entire, connate
to the filament above the staminodes; lateral
staminodes petaloid, free from labellum, elliptic,
inserted at about the same level as the petals;
anther with or without lateral appendages, apical
crest absent. Capsule usually globose, smooth,
verrucose or ribbed, dehiscent; seed with lacerate
aril. 2n = 16, 22, 24, 28, 32, 34, 48, 64, 80, 96; 32 and
48 the most frequent numbers. About 100 spp.,
eastern tropical Himalaya to S China and through-
out India and Indo-China to Malesia; most species
in the Indo-Chinese monsoon region.

19. Parakaempferia A.S. Rao & D.M. Verma

Parakaempferia A.S. Rao & D.M. Verma, Bull. Bot. Surv. India
11: 206, Fig. 1-10 ("1969", 1971).

Herbs with stout rhizome. Leafy shoots elongated.

Inflorescence elongate, terminal on a separate
leafless shoot, flowering from apex to base; bracts
membranous, elliptic, coloured, each subtending
1 flower; bracteole open to the base and entire;
flowers white with pink venation; calyx tubular, 3-
dentate; lateral staminodes similar in size to the
petals; labellum conspicuous and pendulous,
obovate, emarginate; filament short, anther with
short, broad connective and conspicuous, reni-
form crest; ovary 3-locular. Capsule subglobose,
dehiscence unknown; seeds unknown. Only one
sp., P. synantha A.S. Rao & D.M. Verma, India
(Assam), known only from type collection.

II. Tribe Globbeae Petersen (1889).

Small herbs with short or poorly developed
rhizome. Plane of distichy of leaves parallel to
rhizome. Lateral staminodes present. Filament
and style long-exserted from the flower, arched as
a bow, the style forming the string of the bow
(except in Hemiorchis). Ovary 1-locular with
parietal placentation. A

20. Globba L. Fig. 122

Fig. 122A-D. Zingiberaceae-Globbeae. Globba muluensis. A
Globba L., Mantissa 2: 170 (1771); Holttum, Gard. Bull. Inflorescence. B Flower. C Anther and upper part of style with
Singapore 13:21 (1950); Lim, Notes R. Bot. Gard. Edinb. 31: stigma, ventral view. D Ovary, transversal section. (Smith
243-285 (1972); R.M. Sm., ibid. 45: 1-19 (1988), reg. rev. 1988)
488 Zingiberaceae

21. Mantisia Sims
Mantisia Sims, Bot. Mag. ad t. 1320 (1810); Burtt & R.M. Sm.,
Notes R. Bot. Gard. Edinb. 28: 287-290 (1968).
Small herbs with short rhizome. Inflorescence
a lax, many-flowered panicle, terminal on the
leafy shoot or on a separate leafless shoot; bracts
rather large, several-flowered; bracteole open to
the base; flowers yellow or orange with violet or
blue; calyx tubular, 3-dentate; corolla tube much
longer than calyx; lateral staminodes connate to
the filament and forming linear or spathulate
appendages on either side of it; labellum 2-lobed
to entire, inserted at the same level as the petals;
anther with 2 or 4 lateral appendages, crest
rounded. Capsule globose, dehiscence unknown.
2n = 20. Four spp., NE India to N Myanmar. The
generic distinction from Globba needs further
study.
cave, entire, ovate, wider than the staminodes;
filament short, not exceeding the petals, anther
recurved, crest absent. Capsule globose or fusi-
form, dehiscence unknown; seeds with short,
white aril. Three spp., central to eastern Himalaya
and Myanmar, very rarely collected. May be mis-
placed in Globbeae.
III. Tribe Zingibereae Petersen (1889).
Rhizome horizontal, robust. Plane of distichy of
leaves parallel to rhizome. Inflorescence con-
gested, terminal on a separate leafless shoot;
bracts imbricate. Lateral staminodes petaloid
and adnate to labellum, thus forming a 3-lobed
structure. Anther with elongated crest wrapped
around the style, exposing the stigma. Ovary
3-locular.

22. Gagnepainia K. Schum. 24. Zingiber Boehm. Fig. 123

Gagnepainia K. Schum. in Engler, Pflanzenreich IV. 46: 129 Zingiber Boehm. in Ludwig, Def. Gen. Pl. ed. 3: 89 ( 1760), nom.
(1904). cons.

Small herbs with short rhizome; roots fleshy, often Herbs with creeping, fleshy rhizome. Inflores-
bearing tubers. Stem short. Inflorescence a cence cylindrical to ovoid, long pedunculate or
raceme, terminal on a separate leafless shoot; sessile; bracts often bright red or yellow, closely
bracts and bracteole absent; flowers single on the imbricate or with the apices free, forming pouches
rachis; flowers greenish; calyx tubular, 3-dentate; filled with mucilage, each enclosing a 1(-2-4)-
corolla tube longer than calyx; lateral staminodes flowered cincinnus, margins patent or inflexed;
petaloid, elliptic to rhombic, inserted at about bracteole open to the base; flower colour white
the same level as the petals; labellum shorter to yellow with red to purple markings; calyx
than the petals, 3-lobed, the central lobe very tubular, split down 1 side; corolla tube about as
short with 2 glandlike basal swellings; anther long as the bracts, lateral lobes connate to each
with very short crest. Capsule ellipsoid, fleshy, other and to the centre of the labellum in the lower
dehiscence unknown; seeds with short, white 1/3; labellum 3-lobed, filament short. Capsule first
aril. Three spp., Thailand to Cambodia and S fleshy, later leathery, subglobose, dehiscent; seeds
Vietnam. with white, lacerate aril. 2n = 22, some cultivars
polyploid. About 100 spp., widespread in tropical
Asia with centres of diversity in Indo-China and
23. Hemiorchis Kurz Borneo.
Hemiorchis Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 42: 108,
t. 8 (1873).
IV. Tribe Alpinieae Meisn., Pl. Vase. Gen. 2: 291
Small, rhizomatous herbs with fleshy roots (1842).
without storage tubers. Inflorescence a rather lax
raceme, terminal on a separate leafless shoot; Often tall herbs. Plane of distichy of leaves
bracts caducous, each subtending 1 flower; transverse to rhizome. Lateral staminodes much
bracteole absent; flowers yellow or red; calyx reduced or absent. Style not exserted beyond
tubular, 3-lobed; corolla tube of about the same anther (except in Pommereschea which may be
length as the calyx; lateral staminodes petaloid, misplaced). Ovary mostly 3-locular, if 1-locular
obovate to suborbicular, arising at same level as then with basal placentation (except in Riedelia
the petals, but somewhat shorter; labellum con- where parietal placentation occurs).
 Zingiberaceae
.· s
© H
Fig. 123A-H. Zingiberaceae-Zingibereae. A-D Zingiber colo-
ratum. A Flowering spike. B Flower with primary bract and
bracteole. C Corolla tube with ovary, epigynous glands, dorsal
petal, style and anther. D Ovary in transversal section. E-H Z.
acuminatum. E Flower. F Anther and dorsal petal. G Labellum
and lateral petals. H Corolla tube in transversal section, s style.
(A-D Burtt and Smith 1972b; E-H Smith 1989)
25. Alpinia Roxb. Fig. 124
Alpinia Roxb. in Asiat. Res. 11: 350 (1810), nom. cons.; R.M.
Sm., Edinb. J. Bot. 47: 1-75 (1990).
Cenolophon Bl. (1827).
Kolowratia C. Pres! (1827).
Eimeria Rid!. (1909), non Elmera Rydb. (1905).
Adelmeria Rid!. (1909).
Languas J. Konig ex Small (1913).
Herbs, often large, from well-developed rhizome.
Leaves many or rarely few on the shoot. Inflores-
cence usually terminal on the leafy shoot, lax or
congested, sometimes surrounded by sterile
bracts, sometimes branched; bracts when present
open to the base, cincinni !-many-flowered;
489
~
G~
Fig. 124A-G. Zingiberaceae-Alpinieae. Alpinia glabra. A
Inflorescence (br bracteole). B Two-flowered cincinnus,
bracteole of first flower fallen, flower showing calyx, dorsal
petal and labellum. C Calyx. D, E Dissected flower, with ovary,
floral tube, stamen and dorsal petal (D) and labellum with
lateral staminodes (E). F Anther. G Ovary, transversal section.
(Smith 1985)
bracteole tubular or open to the base, sometimes
completely enclosing a cincinnus, sometimes
caducous, or absent; flowers varying from small
to large, red, yellowish or white to greenish; calyx
tubular, 3-dentate, often split down one side; co-
rolla tube usually equal in length to the calyx; lat-
eral staminodes small or absent; labellum erect
or concave, sometimes spreading or pendulous,
varying in shape, variously lobed or entire; fila-
ment absent or present, sometimes appendaged,
anther crest present or absent; ovary 3-locular,
rarely 1-locular. Fruit indehiscent, globose or
ellipsoid; seeds arillate. 2n = 36, 48, 50. More than
200 spp., throughout tropical Asia to New Guinea,
Australia, the Solomon Islands, New Hebrides,
New Caledonia, Fiji and Samoa. The largest genus
in the family, which has previously, quite unsatis-
factorily, been split into several segregate genera
(see Holttum 1950). No recent revision exists,
but Smith (1990) gives a framework for a new
infrageneric classification, keys, and a complete
list of synonyms and distribution maps of the
infrageneric groups.
490 Zingiberaceae

26. Renealmia L. f.
Renealmia L. f., Suppl. Pl. Syst. Veg.: 7 (1782), nom. cons.;
Maas, Fl. Neotrop. 18 (1977); Lock in Fl. Trop. East Africa,
Zingiberac.: 10-14 (1985).

Herbs with well-developed rhizomes and pseu-

dostems. Inflorescence usually lax, sometimes
branched, terminal on a separate leafless shoot,
erect or prostrate, or rarely terminal on a leafy
shoot; bracts subtending 1-several-flowered cin-
cinni; bracteoles tubular; flowers white to yellow;
calyx tubular to turbinate, 3-dentate; corolla tube
somewhat longer than the calyx, about as long to
twice as long as the labellum; lateral staminodes
small, toothlike; labellum erect or spreading,
3-lobed, longer than the petals; filament short,
anther without crest; ovary 3-locular. Capsule
globose to ellipsoid, ± fleshy, dehiscent from base
to apex; seeds with often orange aril. 2n = 22 or @ F
II
I'
44. About 75 spp., mostly neotropical (Mexico,
Caribbean, tropical S America), about 20 species
in Africa; in lowland and montane forest, from sea
level up to 2500 m, rarely higher. An informal
subdivision is given by Maas (1977).

27. Plagiostachys Ridl. Fig. 125

Plagiostachys Rid!., J. Straits Branch R. Asiat. Soc. 32: 151
(1899); R.M. Sm., Notes R. Bot. Gard. Edinb. 42: 265-272
(1985).

Herbs with the rhizome on or just below the

ground. Inflorescence congested, terminal on a
leafy shoot but apparently lateral, breaking
through the leaf sheaths, often mucilaginous; Fig. 125A-I. Zingiberaceae-Alpinieae. A-F Plagiostachys
bracts absent or rudimentary, flowers singly on albiflora. A Young inflorescence emerging from leaf sheaths.
the rachis; bracteole usually tubular; flowers B Infructescence. C Calyx. D Flower, dissected, showing dorsal
yellow, white, orange or red; calyx tubular or petal and stamen (left) and labellum (right) . E Anther and
stigma. F Ovary, transversal section. G-1 P. strobilifera. G
funnel-shaped, 3-dentate, split down 1 side; co- Young inflorescence. H Capsule, surmounted by intact calyx. I
rolla tube shorter than calyx; lateral staminodes Flower. (Smith 1985)
small, subulate or swollen; labellum slightly
longer than petals, obovate, margin crenulate; fila-
ment short, broad, anther with a small crest; ovary
3-locular. Capsules globose to ellipsoid, indehis- Tall herbs with robust rhizome. Inflorescence a
cent, more or less united in a syncarp; seeds ?with raceme, terminal on the leafy shoot; bracts con-
aril. 2n = 98. About 20 spp., W Malaysia through- spicuous, each subtending 1 flower; bracteole
out the Malesian region, a single representative in absent; flowers pink to white; calyx funnel-shaped,
S China. The species fall into 2 groups, 1 with 3-dentate; corolla tube shorter than calyx, the
mucilaginous inflorescence, the other without. lateral petals basally connate to each other and to
This division is accompanied by morphological the labellum; lateral staminodes filiform, 1-1.5 em
differences. A genus in need of revision. long; labellum about as long as the petals, bifid,
with linear lobes; filament elongated, enclosing
the style, anther with a broad rounded connective,
28. Vanoverberghia Merr.
crest absent; ovary 3-locular. Capsule ellipsoid,
Vanoverberghia Merr., Philipp. J. Sci., Bot. 7: 76 (1 912). tardily dehiscent, smooth, with an aniselike smell;
 Zingiberaceae
seeds without? aril. Only one sp., V. sepulchrei
Merr., from the Philippines.
29. Riedelia Oliv.
Riedelia Oliv., Hooker's Icon. Pl. 15: 15, t. 1419 (1883), nom.
cons.; Valeton, Bot. Jahrb. Syst. 52:42-100 (1914); R.M. Sm.,
Notes R. Bot. Gard. Edinb. 31: 297-306.
Rhizomatous herbs, terrestrial or sometimes epi-
phytic. Inflorescence usually a lax raceme, ter-
minal on the leafy shoot, or sometimes congested
and terminal on a separate leafless shoot; bracts
small and caducous or absent, cincinni 1-3-
flowered; bracteole absent; flowers yellow to
orange; calyx tubular, split down on 1 side, cadu-
cous, apex entire or 3-dentate; corolla tube shorter
or longer than calyx, dorsal petal cucullate, some-
times rostrate; lateral staminodes absent; labellum
erect, 2-lobed to bifid, joined to the lateral petals
and of similar length; filament short and broad,
anther crest inconspicuous or absent; ovary 1- or
3-locular. Capsule fleshy, globose to ellipsoid,
dehiscent; seeds with basal, sometimes cupular
aril. About 60 spp., E Malesia, New Guinea, the
Bismarck Archipelago and the Moluccas. In need
of revision.
30. Burbidgea Hook.f.
Burbidgea Hook.f., Bot. Mag. ad t. 6403 (1879); R.M. Sm.,
Notes R. Bot. Gard. Edinb. 31: 297-306 (1972).
Terrestrial or sometimes epiphytic herbs with
creeping rhizome. Leaves sometimes somewhat
succulent. Inflorescence a raceme, terminal on the
leafy stem; bracts usually absent, or papery and
caducous, flowers borne singly or in pairs;
bracteole absent; flowers yellow to orange-red or
pinkish; calyx tubular, 2-dentate; corolla tube
longer than the calyx or of the same length,
petals conspicuous, dorsal one cucullate; lateral
staminodes absent; labellum narrow, shorter
than the petals, erect, 2-lobed; anther sessile,
crest elongated, dentate or entire; ovary 3-
locular. Capsule narrowly elongated, irregularly
dehiscent by 1 lateral slit; seeds with lacerate,
chartaceous aril. 2n = 48. Eight spp., Borneo.
31. Pleuranthodium (K. Schum.) R.M. Sm.
Pleuranthodium (K. Schum.) R.M. Sm., Edinb. J. Bot. 48: 63-
68 (1991).
Psychanthus (K. Schum.) Rid!. (1916), non Raf. (1814); R.M.
Sm., Edinb. J. Bot. 47: 77-82 (1990).
491
Large, rhizomatous herbs. Inflorescence a raceme,
terminal on a separate leafless shoot, sometimes
pendulous, sometimes pushed out laterally from
the uppermost leaf sheath; bracts absent or cadu-
cous, flowers borne singly on the inflorescence
axis; bracteole absent or caducous; flower colour
unknown; calyx short, campanulate; corolla lobes
somewhat shorter than the labellum; lateral
staminodes presumably represented by basal
swellings on the labellum; labellum concave,
usually broader than long, the side lobes curving
behind the filament or connate to the base of the
filament; filament boat-shaped or flat, thecae often
divergent, crest absent; ovary 3-locular. Capsule
ellipsoid to subglobose, fleshy, splitting to the base
into 3 equal valves; seeds ?with aril. About 25 spp.,
New Guinea, Bismarck Archipelago and Australia.
32. Nanochilus K. Schum.
Nanochilus K. Schum., Bot. Jahrb. Syst. 27: 341, tab. V: C-D
(1899); R.M. Sm., Notes R. Bot. Gard. Edinb. 38: 13-17
(1980).
Rhizomatous herbs. Inflorescence congested, ter-
minal on the leafy shoot; bracts present, flowers
borne singly on the inflorescence axis; bracteole
tubular; flower colour unknown; calyx tubular,
split down 1 side, 3-dentate; corolla tube narrow,
shorter than calyx; lateral staminodes petaloid,
over 2 em long, shorter than the petals, linear,
fused to the filament; labellum considerably
reduced, linear, less than 1 em long; stamen about
as long as the petals, filament much longer than the
anther, fleshy, margins incurved holding the style,
anther with a very short crest; ovary imperfectly 3-
locular. Capsule somewhat fleshy, subglobose, de-
hiscence unknown; seeds unknown. Only one sp.,
N. palembanicus (Miq.) K. Schum., Sumatra.
33. Rhynchanthus Hook.f.
Rhynchanthus Hook.f., Bot. Mag. ad t. 6861 (1886); R.M. Sm.,
Notes R. Bot. Gard. Edinb. 38: 13-17 (1980).
Low herbs with tuberous rhizome. Inflorescence
terminal on the leafy shoot; bracts large, each sub-
tending 1 sessile flower; bracteole small, tubular;
flowers yellowish to red; calyx tubular, split down
1 side, apically 2-lobed; corolla tube long, lobes
short, erect; lateral staminodes absent; labellum
reduced to a small tooth at base of the filament or
absent; filament long-exserted, petaloid, boat-
shaped, narrowed at apex, anther without connec-
tive crest; ovary 3-locular, ovules with cup-shaped
492 Zingiberaceae
aril. Capsule unknown. 2n 44. Six spp.,
Myanmar to S China.
34. Leptosolena C. Presl
Leptosolena C. Pres!, Reliq. Haenk. 1: 111, t. 18 (1827); Ridley,
Philipp. J. Sci. IV: 180-181 (1909).
Tall herbs. Inflorescence paniculate, terminal on
the leafy shoot, usually with 3-4 bracts at the base
of the inflorescence; bracteole absent; flowers
sessile or pedicellate, to 12 em long, borne singly
on the inflorescence axis, white; calyx tubular,
shortly 3-lobed, split on one side; corolla tube
narrow and about twice as long as the calyx, lobes
linear; lateral staminodes small, linear; labellum
twice as long as the petals, oblong, obtuse, entire,
recurved; filament very short, anther ellipsoid,
crest absent or not; ovary 3-locular. Capsule
unknown. A poorly known genus, endemic to the
Philippines, Luzon, with Leptosolena haenkei
Presl as the only species.
35. Pommereschea Wittm.
Pommereschea Wittm., Gartenfl. 44: 131, fig. 35 (1895); K.
Larsen, Nat. Hist. Bull. Siam Soc. 24: 475-477. Fig. 2. 1973.
Low herbs with short rhizome. Leaves with cor-
date or subsagittate base. Inflorescence a raceme,
terminal on the leafy shoot; each bract subtending
1 subsessile flower; bracteole minute; flowers
yellow; calyx tubular, split down 1 side; corolla
tube longer than calyx; lateral staminodes absent;
labellum about as long as the petals, erect, nar-
rowly spathulate, 2-lobed, basally connate with
filament into a distinct tube above the insertion of
the petals; filament long-exserted, curved, linear,
anther crest absent, thecae basally diverging;
ovary 3-locular. Capsule subglobose, dehiscence
unknown; seeds with short aril. 2n = 22. Two spp.,
N Myanmar and adjacent Thailand.
36. Etlingera Giseke
Etlingera Giseke, Prael. Ord. Nat.: ad 202,209 (1792); Burtt &
R.M. Sm., Notes R. Bot. Gard. Edinb. 43: 235-241 (1986);
R.M. Sm., ibid.: 243-254 (1986); R.M. Sm., ibid.: 440-457,
part. rev.
Geanthus Reinw. (1825), non Raf. (1814).
Achasma Griff. (1851).
Nicolaia Horan. (1862), nom. cons.
Phaeomeria (Rid!.) K. Schum. (1904).
Tall herbs up to 8m with stout rhizome. Leafy
shoots usually with many leaves. Inflorescence
terminal on a separate leafless shoot, subglobose,
very often broader than long, often surrounded by
an involucre of sterile bracts, peduncle long or
very short, then inflorescence subterranean with
only the apical part visible; bracts densely imbri-
cate, subtending 1 flower; bracteole tubular;
flowers red to yellow; calyx tubular, 3-lobed to 3-
dentate, often split down 1 side; corolla tube long,
lobes shorter than the tube; lateral staminodes
absent or rudimentary; labellum conspicuous,
variously lobed to entire, central lobe expanded to
orbicular, labellum and filament connate into a
tube above the insertion of petals; free part of fila-
ment short of absent, anther crest absent; ovary 3-
locular. Fruit indehiscent, globose to obovoid,
longitudinally ribbed, sometimes united in a
fleshy, globose or elongated syncarp; seeds aril-
late. 2n = 48. About 70 spp., SE Asia from the
Himalayas to N Queensland, with main distribu-
tion in Indonesia. Generic limits still unclear, par-
ticularly towards Hornstedtia and Amomum;
however, a tube formed by the fusion of the label-
lum and filament is never found in these genera.
37. Amomum Roxb.
Amomum Roxb., Pl. Coromandel 3: 75 (1820), nom. cons.;
R.M. Sm., Notes R. Bot. Gard. Edinb. 42: 295-314 (1985),
part. rev.
Paramomum S.Q. Tong (1985).
Tall herbs, some stiltrooted, rhizome usually thin.
Leaves usually many. Inflorescence congested, ter-
minal on a separate leafless shoot, peduncle some-
times short, then inflorescence subterranean and
only the apical part visible; bracts closely imbri-
cating, rather large, 1-several-flowered; bracteole
tubular or open to the base; flowers white, yellow,
orange or red; calyx tubular, usually 3-dentate;
corolla tube about as long as calyx or longer,
sometimes lateral petals centrally adnate to each
other and to the labellum in their lower part;
lateral staminodes small or absent; labellum
longer than the petals, spathulate, variously lobed;
stamen shorter than labellum, anther with well-
developed, variously lobed, entire or absent crest,
sometimes basally spurred; ovary trilocular. Cap-
sule dry or fleshy, subglobose to flask-shaped,
smooth ribbed or echinate, dehiscent, sometimes
united in a syncarp; seeds arillate. 2n = 24, 48, 96.
About 150 spp., widely distributed in tropical Asia
to tropical Australia, the majority in the Malesian
region. No satisfactory revision exists and the de-
limitation from genera such as Etlingera and
Elettariopsis is insufficiently understood.
 Zingiberaceae
38. Hornstedtia Retz.
Hornstedtia Retz., Obs. Bot. 6: 18 (1791); R.M. Sm., Notes R.
Bot. Gard. Edinb. 42: 289-295 (1985), part. rev.
Tall herbs; rhizome usually stout. Inflorescence
congested, fusiform, terminal on a leafless
shoot, short- to rarely long-pedunculate, entirely
enclosed by an involucre of several rigid, sterile
bracts; bracts subtending usually only one flower;
bracteole open to the base, sometimes absent;
flowers white, yellow, orange or red; calyx
tubular, split down one side, 2- or 3-dentate;
corolla tube more than twice as long as labellum,
slender, sometimes lateral lobes connate to each
other and to the centre of the labellum in the
lower third; lateral staminodes absent or reduced
to minute teeth at base of labellum; labellum
about the same length as petals, fleshy, concave,
base auriculate; filament sometimes fleshy, or
anther sessile, thecae sometimes villose, some-
times with adnate basal spurs, crest usually
present; ovary 3-locular. Capsules globose or
ellipsoid, irregularly dehiscent, sometimes united
in a syncarp inside the persistent involucre;
seeds with thin, cup-shaped aril. 2n = 48.
About 50 spp., mainly in Malesia and tropical NE
Australia, but also in Thailand, Indochina and
China (Hainan, Tibet).
39. Aframomum K. Schum.
Aframomum K. Schum. in Engler, Pflanzenreich IV. 46: 201
(1904); Lock in Kew Bull. 35: 299-313 (1980) and Fl. Trop.
East Afr., Zingiberac.: 22-37 (1985), reg. rev.
Herbs, with well-developed rhizome. Inflores-
cence usually congested, terminal on a separate
leafless shoot, sometimes subterranean with only
the apical part visible; bracts each subtendig 1
flower; bracteole open to the base; flowers large,
violet, red, or yellow, rarely white; calyx split down
1 side; corolla tube narrow, about as long as the
calyx, sometimes petals, stamen, and staminodes
basally connate into a short tube; lateral sta-
minodes small and toothlike; labellum either peta-
loid, broad and suberect or narrower, thicker and
decurved; filament strap-shaped, anther crest con-
spicuous, usually 3-lobed; ovary 3-locular. Cap-
sule ovoid to globose, beaked, fleshy, ridged or
smooth, indehiscent; seeds with white, translucent
aril. 2n = 48. About 50 spp., tropical Africa,
Mascarenes and Madagascar; one species, A.
melegueta, cultivated for its spicy seeds and
widely cultivated in W Africa and also in S
America.
493
40. Aulotandra Gagnep.
Aulotandra Gagnep., Bull. Soc. Bot. Fr. 48: lxxix (1902);
H. Perr. in Fl. Madagascar, 47. Zingiberaceae: 9-19 (1946).
Low herbs from a short vertical or horizontal rhi-
zome; roots sometimes with tubers. Leaves few.
Inflorescence erect, lax, terminal on a separate
leafless shoot; bracts 1-flowered; bracteole absent;
flowers whitish with yellow, violet or red marks;
calyx shortly tubular, 2-3-dentate, split down 1
side; corolla tube short; lateral staminodes
reduced to 2 basal swellings; labellum petaloid,
variously lobed; base of labellum and filament
forming a distinct tube above the insertion of the
petals; free part of filament short, anther crest
petaloid, variously lobed; ovary 3-locular. Capsule
trigonous or trialate, dehiscent or withering with
age; seeds with undivided aril. About five spp.,
Madagascar, one in Cameroon. A poorly known
genus; more work is needed to establish whether
all species belong to it, and whether the genus is
worthy of recognition at all.
41. Geocharis (K. Schum.) Ridl.
Geocharis (K. Schum.) Ridl., J. Straits Branch Roy. Asiat. Soc.
50: 143 (1908); R.M. Sm., Notes R. Bot. Gard. Edinb. 43:457-
460 (1986)
Alpinia section Geocharis K. Schum. (1904).
Low to tall herbs with robust rhizome. Inflores-
cence lax, terminal on a separate leafless shoot,
erect or prostrate; bracts thin, persistent or not,
spaced, leaving the axis visible, each subtending
2 flowers; bracteole tubular, persistent or not;
flowers yellow, red or white; calyx tubular, long
and slender, 3-lobed; corolla tube slender, as long
as or longer than calyx; lateral staminodes absent;
basal part oflabellum and filament connate into a
tube above the insertion of the petals, free part
of labellum narrow, bifid; filament rather long,
broad, more or less boat-shaped, often with lateral
teeth just below the anther, crest present; ovary 3-
locular. Capsule fleshy, narrowly ellipsoid, ridged
or verrucose, dehiscence unknown; seeds with
thin, fleshy aril. About seven spp., W Malaysia and
Sumatra to Borneo and the Philippines. The genus
has some superficial resemblance with Elettaria
and Hornstedtia, but its relationship and circum-
scription have still to be established.
42. Elettariopsis Baker
Elettariopsis Baker in Hook. f., Fl. Brit. India 6: 251 (1892);
Kam, Notes R. Bot. Gard. Edinb. 40: 139-152 (1982), rev.
494
Herbs with short pseudostems, and slender,
creeping rhizome. Leaves few, usually petiolate
and in radical tufts. Inflorescence terminal on a
separate leafless shoot, congested and erect, or
lax, branched and prostrate, with well-spaced,
distichous bracts, and at least partially subterra-
nean; bracts subtending 1(-2) flowers; bracteole
open to the base; flowers white with red and
yellow; calyx tubular, 3-dentate; corolla tube
slender, longer than the calyx; lateral staminodes
reduced to small basal swellings on the labellum
or absent; labellum broad, erect, variously lobed
or entire, longer than the petals; filament short,
anther crest petaloid, almost quadrangular;
ovary 3-locular. Capsule globose, ridged, dehis-
cence unknown; seeds arillate. 2n = 48. About ten
spp., Indomalesia, mostly in evergreen rainforest;
some insufficiently known. The limits of the genus
and the relationship with Amomum are not yet
clear.
43. Cyphostigma Benth.
Cyphostigma Benth., Hooker's Icon. Pl.: t. 1380 (1882); Burtt &
R.M. Sm., Fl. Ceylon 4: 530, Fig. 5 (1993).
Herbs with creeping rhizome. Stem short with few,
basal, erect leaves. Inflorescence terminal on a
separate leafless shoot, lax, branched, prostrate,
subterranean with only the tips of the flowers
visible; bracts remote, each subtending a single
resupinate flower; bracteole tubular; flowers pink;
calyx tubular, shortly 3-lobed, split down 1 side;
corolla tube about as long as the calyx; lateral
staminodes absent; labellum about as long as the
petals, orbicular, obscurely 3-lobed; filament
short, anther crest petaloid, almost as large as la-
bellum; ovary 3-locular. Capsule globose, ribbed,
dehiscence unknown; seeds arillate? Only one sp.,
C. pulchellum (Thwaites) Benth., endemic to Sri
Lanka.
44. Geostachys (Baker) Ridl.
Geostachys (Baker) Rid!., J. Straits Branch R. Asiat. Soc. 32: 157
(1899); Holttum, Gard. Bull. Singapore 13: 224-236, Fig. 30
(1950).
Alpinia subgen. Geostachys Baker (1892).
Acaulescent herbs, leaves in radical tufts; rhizome
stout, on or above the ground stiltrooted. Inflores-
cence lax, terminal on a separate leafless shoot,
erect or decurved with 1-5 flowers to each bract;
bracts small, inconspicuous; bracteole tubular, in-
flated, enclosing the cincinnus, opening at apex
Zingiberaceae
only; flowers yellow or pink with reddish marks;
calyx tubular, split down 1 side; corolla tube
slightly shorter than calyx; lateral staminodes usu-
ally absent; labellum erect, longer than petals, en-
tire to distinctly 3-lobed; filament strap-shaped, as
long as anther, crest mostly absent, sometimes
entire or 3-lobed, small; ovary 3-locular. Capsule
globose or ellipsoid, sometimes ribbed, dehis-
cence unknown; seeds arillate. About 18 spp., S
Vietnam, S Thailand, W Malaysia to Sumatra, and
Borneo; most species from the Malay Peninsula.
45. Elettaria Maton
Elettaria Maton, Trans. Linn. Soc. Lond. 10: 250 (1811); R.M.
Sm., Notes R. Bot. Gard. Edinb. 43: 461-463 (1986).
Low to robust herbs with creeping rhizome.
Inflorescence elongate, branched, terminal on a
separate leafless shoot, usually prostrate and sub-
terranean with only the tips of the flowers visible;
bracts remote, each subtending a several-many-
flowered cincinnus; bracteole tubular; flowers
white, yellow or orange; calyx tubular, 3-dentate,
split down 1 side; corolla tube more than twice
the length of the lobes; lateral staminodes small,
subulate or absent; labellum about as long as the
petals, obovate; filament short or absent, crest
inconspicuous or absent; ovary 3-locular. Capsule
globose or ellipsoid, indehiscent; seeds arillate.
2n = 48, 50. About seven spp., four of which in
the Malesian region, one, the cardamom, E.
cardamomum, endemic to Sri Lanka, but much
cultivated in India and Guatemala, and
naturalised in E Africa (Tanzania).
Incertae sedis
46. Stadiochilus R.M. Sm.
Stadiochilus R.M. Sm., Notes R. Bot. Gard. Edinb. 38: 15
(1980).
Rhizomatous terrestrial or epiphytic herbs. Leaves
3-7, long acuminate. Inflorescence a congested
raceme, terminal on the leafy stem; bracts large,
each subtending 1 flower; bracteole tubular, split
down 1 side; flowers red-purple; calyx tubular,
split down 1 side; petals linear, as long as the
labellum; lateral staminodes absent; labellum obo-
vate, 2-lobed, erect, the margins overlapping be-
hind the filament, thus resembling a tube beyond
which the filament protrudes; filament elongated,
linear, anther crest minute; ovary 3-locular.
Capsule unknown. Only one sp., S. burmanicus
R.M. Sm., N Myanmar.
 Zingiberaceae
Cornukaempferia J. Mood & K. Larsen, Nat. Hist. Bull. Siam
Soc. 45: 217 (1997).
Monotypic genus from N. Thailand, C.
aurantiflora ]. Mood & K. Larsen. Related to
Kaempferia from which it deviates in the undi-
vided, orange labellum, an anther crest in the form
of a curved, hornlike appendage enclosing the dis-
tal part of the style similar to what is found in
Zingiber.
Selected Bibliography
Beltran, I.C., Kam, Y.K. 1984. Cytotaxonomic studies in the
Zingiberaceae. Notes R. Bot. Gard. Edinb. 41: 541-559.
Burtt, B.L. 1972. General introduction to papers on Zingibera-
ceae. Notes R. Bot. Gard. Edinb. 31: 155-165.
Burtt, B.L., Smith, R.M. 1972a. Tentative keys to the sub-
families, tribes and genera of Zingiberaceae. Notes R. Bot.
Gard. Edinb. 31: 171-176.
Burtt, B.L., Smith, R.M. 1972b. Key species in the taxonomic
history of Zingiberaceae. Notes R. Bot. Gard. Edinb. 31:
177-227.
Chen, Z-Y. 1989. Evolutionary patterns in cytology and pollen
structure of Asian Zingiberaceae. In: Holm-Nielsen, L.B.,
Nielsen, I.C., Balslev, H. (eds.) Tropical forests. Botanical
dynamics, speciation and diversity. London: Academic
Press, pp. 185-199.
Clark, W.D., Gaut, B.S., Duvall, M.R., Clegg, M.T. 1993.
Phylogenetic relationships of the Bromeliifiorae-
Commelinifiorae-Zingiberifiorae complex of monocots
based on rbcL sequence comparisons. Ann. Mo. Bot. Gard.
80: 987-998.
Dahlgren, R.M.T. eta!. 1985. See general references.
Davis, G.L. 1969. See general references.
Frolich, D., Barthlott, W. 1988. See general references.
Hegnauer, R. 1963, 1986. See general references.
Hickey, L.H., Peterson, R.K. 1978. Zingiberopsis, a fossil genus
of the ginger family from Late Cretaceous to Early Eocene
sediments of Western Interior North America. Can. J. Bot.
56: 1136-1152.
Holttum, R.E. 1950. The Zingiberaceae of the Malay Peninsula.
Gard. Bull. Singapore 13: 1-249.
Johri, B.M. eta!. 1992. See general references.
Kam Yee Kiew. 1982. The genus Elettariopsis (Zingiberaceae)
in Malaya. Notes R. Bot. Gard. Edinb. 40: 139-152.
Kato, M. 1996. Plant-pollinator interactions in the under-
storey of a lowland mixed dipterocarp forest in Sarawak.
Am. J. Bot. 83: 732-742.
Koechlin, J. 1965. Flore du Cameroun. Zingiberaceae: 19-95.
Kress, W.J. 1990. The phylogeny and classification of the
Zingiberales. Ann. Mo. Bot. Gard. 77: 698-721.
Kiinckel d'Herculais, J. 1910. Rapport des insectes lepi-
dopteres avec les fieurs des zingiberacees et en particulier
avec celles des Hedychium. C.R. Hebd. Acad. Sc. Paris CLI:
1153-1155.
Liang Y.-H. 1988. Pollen morphology of the family Zingibera-
ceae in China - pollen types and their significance in the
taxonomy. Acta Phytotaxon. Sin. 26: 265-281 (in Chinese
with English summary).
Lock, J.M., Hall, }.B., Abbiw, D.K. 1977. The cultivation of
Melegueta pepper (Aframomum melegueta) in Ghana.
Econ. Bot. 31: 321-330.
495
Loesener, Th. 1930. Zingiberaceae. In: Engler, A., Prantl, K.
(eds.) Natiirliche Pfianzenfamilien, 2nd edn., 15a. Leipzig:
W. Engelmann, pp. 541-640.
Lynch, R.I. 1882. On a contrivance for cross-fertilisation in
Roscoea purpurea; with identical reference to the structure
of Salvia Grahami. }. Linn. Soc. 19: 204-206.
Maas, P.J.M. 1977. Flora Neotropica Monogr. 18. Renealmia,
Costoideae (Additions). Bronx, N.Y.: The New York
Botanical Garden.
Mahanty, H.K. 1970. A cytological study of the Zingiberales
with special reference to their taxonomy. Cytologia 35: 13-
49.
Mangaly, J.K., Nayar, }. 1990. Palynology of South Indian
Zingiberaceae. Bot. J. Linn. Soc. 103: 351-366.
Muller, F. 1890. Miscellen. Kreuzung von Hedychium. Abh.
Naturwiss. Ver. Bremen 11: 444.
Nishida, T., Vehara, S. 1983. Natural diet of chimpanzees
(Pantroglodytes schweinfurthii): long-term record from the
Mahale Mountains, Tanzania. Afr. Stud. Monogr. 3: 109-
130.
Pai, R.M. 1965. The floral anatomy of Elettaria cardamomum
Maton. A reinvestigation. New Phytol. 64: 187-204.
Petersen, O.G. 1889. Zingiberaceae. In: Engler, A., Prantl, K.
(eds.) Die natiirlichen Pflanzenfamilien II, 6. Leipzig: W.
Engelmann, pp. 10-30.
Rao, V.S. 1963. The epigynous glands in Zingiberaceae. New
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805.
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496
Zosteraceae
J. Kuo and A.J. McCOMB
Zosteraceae Dumort., Anal. Fam. Pl.: 65, 66 (1829), nom. cons.
Perennial (rarely annual) herbs with creeping,
monopodially branched rhizomes, wholly sub-
merged, sometimes intertidal, marine, rarely es-
tuarine, sometimes extending to brackish water
habitats. Roots unbranched, with root hairs,
growing on sand or mud substrates (Zostera,
Heterozostera) or on rocks (Phyllospadix). Leaves
alternate, distichous, with distinct blade and
sheathing base; leaf sheath ligulate, auriculate;
blade linear, flat to biconvex, leaf apex variable;
stomata absent. Squamules present within each
leaf sheath. Prophylls at the base of each shoot.
Erect shoot annual, possibly sometimes perennial,
and producing inflorescences. Flowers monoe-
cious or dioecious, naked and arranged in 2
rows (in monoecious species of Zostera and
Heterozostera, male and female flowers alter-
nately) on 1 side of a flattened spike (spadix) en-
veloped within a modified leaf sheath (spathe).
Retinacules present or absent. Male flowers of
one stamen, consisting of 2 free, bilocular,
lengthwise dehiscent, deciduous thecae connected
by a ridgelike connective; pollen filiform.
Female flowers of a unilocular ovary with a
single short style and 2 long stigmas; uniovulate.
Pollination hydrophilous. Fruit an achene.
A family consisting of 3 genera and 18 species
from temperate waters in both hemispheres, some
species extending to the subtropics.
VEGETATIVE MORPHOLOGY AND ANATOMY. The
rhizomes are herbaceous and usually cylindrical,
with obscurely monopodia! and irregular branch-
ing, and bear erect, annual (rarely perennial) leafy
shoots (Tomlinson 1974). The rhizome internodes
are very short with a thick cuticle in Phyllospadix
(Cooper and McRoy 1988; Barnabas 1994), but
are rather long in other genera. The rhizome in-
ternodes have a conspicuous stele with 2 or more
cortical vascular bundles and numerous ligni-
fied fibres. The distribution and number of cor-
tical vascular bundles and cortical fibre strands
differ between genera (Jacobs and Williams 1980;
Zosteraceae
Kuo 1983a); cortical fibres are absent and thick-
walled parenchyma cells with reduced intercel-
lular spaces occur in Phyllospadix (Kuo 1983a;
Barnabas 1994). There are small starch grains in
cortical parenchyma cells, the exodermis is not
well defined, and epidermal cells are thin-walled
(Zostera and Heterozostera) or thick-walled
(Phyllospadix).
Roots occur at the nodes in 2 to 4 regular, linear
series with 1-many roots in each series. The roots
are unbranched, have pronounced root hairs, and
sometimes form extensive mats which probably
play an important role in nutrient uptake. Each
root has a central stele surrounded by an endoder-
mis with a distinct Casparian strip and suberised
walls. Xylem parenchyma cells have cell-wall in-
growths (Barnabas and Arnott 1987). The cortex is
covered by a distinct layer of suberised exodermal
cells, which, in those abutting root hair bases, pos-
sess wall ingrowths. Plasmodesmata are present in
both endodermal and exodermal cells of Zostera
(Barnabas 1994).
Leaves are distichous, and arranged alternately
on the erect shoots. The leaf base is narrow, flat-
tened to biconvex, and is closed or open. The
blade is strap-shaped, with a leaf apex which dif-
fers between species, and contains 3 to 11 or more
parallel, longitudinal veins connected periodically
by transverse veins. In addition several longitu-
dinal striations are present between longitudinal
veins, representing diaphragms which separate
longitudinal gas lacunae (Tomlinson 1982; Kuo et
al. 1992). The blade epidermal cells are uniformly
chlorenchymatous, squamous to rectangular and
with smooth margins in surface view. The epider-
mal cells lack tannin. Leaf mesophyll cells are
thin-walled and have large vacuoles and a thin
peripheral cytoplasm containing few chloroplasts;
gas lacunae are well developed, with 3 to 6 large
lacunae between pairs of longitudinal veins. The
lacunae have regular transverse diaphragms. In
each longitudinal leaf vein there are cell-wall in-
growths in sheath and phloem parenchyma cells.
The xylem and phloem are distinctly separated,
a feature which distinguishes them from other
seagrasses. The xylem is represented by single,
wide lacunae without lignified walls. Nacreous-
walled sieve tubes and phloem parenchyma cells
with wall in growths occur in both leaf blade and
sheath. The sieve tubes contain typical monocoty-
ledonous plastids with protein crystalloids. Fibre
strands of thickened but not lignified cells are
abundant in the leaf margins, occur as small
strands associated with longitudinal veins, and are
found in the hypodermis adjacent to gas lacunae
 Zosteraceae 497

and vascular bundles (Kuo et al. 1988, 1990;

Barnabas 1994).
The leaf sheath is tubular and either open or
closed, with distinct auricles and short ligules.
Chloroplasts, wall ingrowths and subcuticular
cavities are absent from both abaxial and adaxial
epidermis of Zostera and only the abaxial epider-
mis of Phyllospadix. The mesophyll resembles that
in the blade, but fibrous strands are more numer-
ous. The vascular bundles are similar to those in
the blade but there are no cell-wall ingrowths in
vascular sheath cells and phloem parenchyma
(Kuo et al. 1988; Kuo and Stewart 1995).

INFLORESCENCE STRUCTURE. The erect shoot is

annual, possibly sometimes perennial, and pro-
duces inflorescences. The inflorescence shoots are
either sympodially branched (Zostera, Hetero-
zostera and Phyllospadix torreyi) or unbranched
(the other four Phyllospadix species) (Fig. 126).
Each branch has 1-few spadices. Developing
monoecious or dioecious flowers are carried on
flattened, spikelike axes (spadices), and enclosed
by a modified leaf sheath (spathe) (Soros-Pottruff
and Posluszny 1994). The margin of the spadix is
incurved, and usually bears retinacules at regular
intervals. Retinacules are present in all members
of the family except in subgenus Zostera of Zostera
(den Hartog 1970).

FLORAL MORPHOLOGY. The plants are either

monoecious (Heterozostera and Zostera) or dioe-
cious (Phyllospadix) (Fig. 126). In monoecious
plants 2 male flowers and 1 female are arranged

alternately in 2 longitudinal rows in the spadix.
In dioecious species, flowers are situated in 2
longitudinal rows in the spadix. Perianth parts
are absent. Male flowers have 1 stamen with 2
bilocular anthers with a ridgelike connective. De-
hiscence is longitudinal. The female flower has an
ellipsoidal or crescent-shaped carpel with 2 long
stigmas. The solitary ovule is pendulous and
atropous. The developmental morphology of re-
productive structures of two Phyllospadix species
has been described in detail (Soros-Pottruff and
Posluszny 1994).
FLORAL ANATOMY. The floral axis in Zostera
marina has 3 longitudinal bundles, 1 median and
1 in each margin. These connect below each pistil
and stamen pair at a complete, oblique commis-
sure. The vascular supply to the carpel is a single
trace from the commissure; this trace remains un-
branched and ends at the base of the ovule. The
vascular supply to the stamen pair is provided
~
.
..
K
Fig. l26A-L. Zosteraceae. A-F Zostera marina. A Flowering
plant. B Spadix with alternating staminate and pistillate
flowers. C Transversal section of spadix with spathe and
staminate and pistillate flower. D Fruit. E Seed. F Branched
stamen. G- L Phy llospadix scouleri. G Habit. H Pistillate spadix
with retinacula and rudimentary staminate flowers. I Pistillate
flower. J Staminate spadix with retinacula. K Male flower. L
Young offruit. (Takhtajan 1982)
directly from the commissure, which is enlarged
below them (Tomlinson 1982).
In Phyllospadix scouleri the vascular trace of the
carpel diverges from the main vascular bundle
without forming a complete commissure across
498 Zosteraceae
the axis. The carpel trace is branched to produce a
dorsal and a ventral trace, ending below the ovule.
The vascular supply to the retinacule (when
present) is by a marginal branch from the stamen
commissure, the branch remaining individual
within the retinacule (Tomlinson 1982).
EMBRYOLOGY. Tomlinson (1982) records that en-
dosperm formation is Helobial, but Dahlgren and
Clifford (1982) comment that Nuclear endosperm
formation has been observed; more work is re-
quired in this area. The first root of Zostera is
initiated endogenously and by its growth breaks
the surrounding tissue, which constitutes a form
of coleorhiza (Taylor 1957).
PoLLEN MoRPHOLOGY. When meiosis in Phyl-
lospadix torreyi is initiated, the pollen mother
cells have already elongated and at metaphase I
the chromosomes lie in a plane across the narrow
axis of the filamentous cell. However, the pole and
spindles rotate through 90° during the anaphase
of the first and second meiotic divisions, and the
cells divide longitudinally to produce filamentous
pollen (Stewart and Liidenberg 1980). Mature pol-
len is filiform, 500 11m long in Zostera and 1500 11m
long in Phyllospadix. The pollen is binucleate
(Zostera) or trinucleate (Phyllospadix), without an
exine (Pettitt 1984). The pollen of the Zosteraceae
is generally thinner than that of the other
seagrasses with filiform pollen (McConchie and
Knox 1989).
In the filiform pollen there is constant cytoplas-
mic streaming, a feature used by de Cock ( 1980) to
indicate pollen viability in Zostera; pollen could
survive 8h.
KARYOLOGY. Zostera species from the northern
hemisphere have chromosome number 2n = 12,
but those from the southern hemisphere have
2n = 24. The Australian Heterozostera has 2n =
ca. 36 (J. Kuo unpubl. 1994). The Japanese
Phyllospadix species have 2n = 20. Male plants
of the Japanese Phyllospadix have a large
heterochromatic chromosome, which is regarded
as a sex-chromosome (Uchiyama 1993). On the
other hand, the Californian P. torreyi pollen
has n = 9 and possibly a heteropycnotic X-
chromosome (Stewart and Liidenberg 1980).
PoLLINATION. In the intertidal Zostera and
Phyllospadix species pollen is released on the
water surface, where it forms a network, and the
stigmas occur on the same water surface, so that
pollination can take place there (de Cock 1980;
Cox 1988). On the other hand, in subtidal species
pollination must take place under water; the
filiform pollen wraps around a stigma during pol-
lination, maximising the surface area of attach-
ment to the stigma (McConchie and Knox 1989).
Pollen germination may be triggered when pollen
contacts the stigma, but numerous papillate
pollen tubes are produced close to each other.
Pollen germination may also occur before pollen
contacts the stigma (de Cock 1978).
The stigma of Zostera is non-papillate and
breaks off after pollination. However, in Z.
marina, which is protogynous, the stigmas remain
in place until the male phase, ensuring self-
fertilization if pollination has not occurred pre-
viously (de Cock 1980). The small pollen tubes
contain callose but only 1 eventually undergoes
further growth and penetrates the stigma. Obser-
vations of an actual fertilisation event are yet to be
reported for any seagrass.
The period of flowering in Heterozostera is
short; the development of flowers is initiated in
August, followed by anthesis in November. In this
case the anthers detach but are not buoyant.
Pollination occurs underwater.
FRUIT AND SEED. The fruits are achenes, either
ovoid to ellipsoid with a scarious pericarp, or
crescent -shaped with a soft, ephemeral exocarp
and a hard, fibrous en do carp. Development of the
seed and seedling of Z. marina has been described
by Taylor (1957) and Yamashita (1973 ). The seeds
lack endosperm. The embryo has a plumule and
an enlarged hypocotyl, 2 adventitious root apices,
and is said to lack a radicle.
DISPERSAL AND REPRODUCTIVE BIOLOGY.
Dispersal is by water. Flowers form during winter
to spring, the flowers opening in spring, and fruit
maturing in late spring to summer. Seedlings may
be found in summer. Flowering in Zostera marina
is probably controlled by water temperature; in
the Atlantic it is later than in the Pacific at the
same latitude (Phillips et al. 1983).
In contrast to other seagrasses, seeds may
enter a dormant phase. However, for Z. marina
higher germination rates were obtained for
seeds collected from reproductive shoots than
for seeds collected from fresh beach debris;
germination was inhibited at 28-32 oc (McMillan
1983).
Germination and establishment is critical for
annual races, but in established clumps or mead-
 Zosteraceae 499

ows of perennial races, most propagation is five in the order Potamogetonales. Dahlgren and
vegetative by growth of rhizomes. Clifford {1982) placed the Zosteraceae and six
The adventitious shoots have horizontal rhi- other families in the order Zosterales.
zomes arising from mature erect fertile shoots. Cox and Humphries {1993) considered that the
These adventitious shoots may develop roots, be- three seagrass families: the Zosteraceae, Posi-
come detached and act as propagules (Cambridge doniaceae and Cymodoceaceae share a common
et al. 1983). ancestor with filiform pollen. On the other hand,
based on the breeding system, it has been
PHYTOCHEMISTRY. Phenolic compounds such suggested that Zosteraceae are closer to the
as sulphated and non-sulphated flavonoids are Potamogentonaceae/Zannichelliaceae than to the
present in Zostera and Phyllospadix but absent Posidonaeceae and Cymodoceaceae (Waycott
from Heterozostera (McMillan et al. 1980; and Les 1996; Les et al. 1997).
McMillan 1983). Despite the discontinuous distri-
bution in Heterozostera, leaf samples from Aus- DISTRIBUTION AND HABITATS. Zostera is widely
tralia and Chile were found to have identical distributed along the coasts of temperate zones of
patterns of seven phenolic compounds but to lack both hemispheres. Phyllospadix is restricted to the
flavonoids (McMillan 1983). northern Pacific Ocean, while Heterozostera
Soluble sugars such as fructose, glucose and occurs in temperate Australia and two localities
sucrose are present in the family. Phyllospadix in northern Chile.
differs from other genera in the family by possess- Members of the family normally grow along
ing an additional trisaccharide (Drew 1983) and shallow, low energy coastlines or in estuaries, and
apiose (Woolard and Jones 1978). may be either subtidal or intertidal. Zostera ex-
tends into brackish water. The plants may occur in
RELATIONSHIPS WITHIN THE FAMILY. DNA se- small clumps, may form large, mono-specific
quence diversity and data from morphology, meadows, or may contribute (with other seagrass
anatomy, karyotype and isoenzyme studies sug- taxa) to meadows of different species. Both
gest significant differences between two subgenera Zostera and Heterozostera are mostly associated
of Zostera, and that the subgenus Zostera is more with muddy or sandy substrates, but Phyllospadix
closely related to Phyllospadix than to the subge- is normally associated with rocky substrates both
nus Zosterella (Uchiyama 1996). Furthermore, re- intertidally and subtidally. In the subartical circle,
cent DNA fingerprinting study indicates that Z. Zostera continues growth under frozen, icy sur-
marina in Central California possesses a high ge- face water.
netic diversity and is not primarily clonal (Alberte Large underwater meadows are visually striking
et al. 1994). The rbcL data of Les et al. {1997) do and support a diverse population of marine flora
not support the recognition of Heterozostera as a and fauna including commercially important
distinct genus. fisheries such as blue crabs. They are highly pro-
ductive and support food webs either through di-
AFFINITIES. The family is a clearly circumscribed rect grazing or through the production of large
group with a highly specialised arrangement of amounts of organic, nutrient-rich detritus.
reproductive organs and anatomical features.
However, the status and the relationship of this PARASITES. The "wasting disease" of eelgrass,
group among other major taxa is less clear. Zostera marina, which destroyed large meadows
Bentham and Hooker {1883) placed the tribe of the N Atlantic Zostera beds, led to a decline in
Zostereae with another seven tribes in the family the population of marine organisms and water-
Najadaceae. Engler {1904) placed this and four fowl in W Europe in 1930-1940. A new outbreak
other tribes in the family Potamogetonaceae. of "wasting disease" has also been reported from
Ascherson and Graebner {1907) and Eckardt N America. The fungus Labyinthula zosterae
{1964) considered the Zostereae to be a subfamily has been considered as being involved in the dis-
in the Potamogetonaceae, along with Posidonio- ease, but there was no direct proof that fungi
ideae and Potamogetoneae ( = Cymodoceae), an were the primary cause of seagrass loss (den
arrangement followed "for the time being" by den Hartog 1996). Lulworthia medusa and Ophilbolus
Hartog (1970). maritimus were also found in Z. marina, but were
Hutchinson {1934) and Takhtajan {1966) re- not associated with the "wasting disease" (den
garded the group as a family Zosteraceae, one of Hartog 1970).
500 Zosteraceae
PALAEOBOTANY. A well-preserved fossil Archeo-
zostera = Archaeozostera containing several
species was described by Koriba and Mil<i (1931,
1958) from the Cretaceous at several localities
in Japan, and has been proposed as a
"protozosteroid", the ancestor of the modern
Zosteraceae (den Hartog 1970; Larkum and den
Hartog 1989). However, re-examination of these
fossil materials led to the conclusion that
Archaeozostera was not associated with the
Zosteraceae and probably was not even a marine
angiosperm (Kuo et al. 1989).
ECONOMIC IMPORTANCE. Members of this family
are economically significant because their large
biomass in nearshore regions is important in food
webs, providing shelter and energy for a large
variety of organisms including commercially-
important fish and waterfowl. Zostera marina has
also been used as packing material.
KEY TO THE GENERA
1. Rhizome thick with condensed internodes; rhizome cortical
cells thick-walled, cortical fibre bundles absent; roots short,
thick; leaf blade thick, fiat to biconvex; normally growing on
rock substrate. Dioecious 3. Phyllospadix
- Rhizome slender, usually with elongated internodes; rhi-
zome cortical cells thin-walled, cortical fibre bundle
present; roots long, thin; leaf blade thin, fiat; normally
growing on sand/mud substrate. Monoecious 2
2. Rhizome internodes with 4-12 cortical vascular bundles,
fibre bundles distributed throughout cortex, commonly as-
sociated with stele; erect shoots long, wiry, branched and
often forming new rhizomes with plantlets; only in older
plants erect shoots sometimes developing inflorescences
2. Heterozostera
- Rhizome internodes with 2 cortical vascular bundles, fibre
fundles restricted to outer cortex; erect shoots herbaceous
not wiry, always developing inflorescences 1. Zostera
1. Zostera L. Fig. 126A-F
Zostera L., Sp. Pl.: 968 (1753), den Hartog, Verk. K. Ned. Akad.
Wet. II, 59 (1): 42-97 (1970), rev.
Rhizome creeping or suberect, herbaceous, mo-
nopodia!, with 2 cortical vascular bundles in each
internode. Vegetative shoot short, bearing 2-6
leaves. Leaves ligulate, auriculate; blade linear,
deciduous, flattened, with 3-11 longitudinal vas-
cular bundles; squamules 2-4. Monoecious, inflo-
rescence shoots simple or sympodially branched;
spathe stalked, resembling vegetative leaves, bear-
ing alternating male and female flowers. Fruits
with a scarious pericarp. At least 12 spp., mainly
in temperate and subtropical waters of the
world; divided into two subgenera: subgen.
Zostera, with leaf sheath closed, inflorescence ter-
minal, retinacules absent, four spp., one of which
Z. marina L. of a wide distribution in theN Atlan-
tic and Pacific; subgen. Zosterella, with leaf sheath
open, inflorescence lateral, retinacules present,
eight spp., mainly in temperate Eurasia, E Africa
and Australasia, some extending into the tropics.
2. Heterozostera den Hartog
Heterozostera den Hartog, Verh. K. Ned. Akad. Wet. II, 59 (1):
114 (1970).
Rhizome creeping, lignified, sympodial to form
erect shoots; rhizome internodes with 4-10
cortical vascular bundles and many fibre bundles
in outer and inner cortex. Vegetative shoots erect,
unbranched; leaves with open sheath, ligulate,
auriculate; blade linear, caducous, with 3 longitu-
dinal vascular bundles; squamules 2. Fertile erect
shoots simple or sympodially branched, forming
cymose inflorescences. Monoecious, male and
female flowers alternating within each spathe.
Retinacules present. Fruits with scarious pericarp.
Only one sp., H. tasmanica (Martens ex Aschers.)
Hartog, S Australia and N Chile.
3. Phyllospadix Hook.
Phyllospadix Hook., Fl. Bor. Am. 2: 171 (1838); den Hartog,
Verh. K. Ned. Akad. Wet. II, 59 (1): 97-114 (1970), rev.
Rhizome creeping, herbaceous, monopodia!; rhi-
zome internodes short, thick, with 2 cortical vas-
cular bundles. Leaves 1 per node; sheath open,
ligulate, auriculate; blade linear, flattened to bi-
convex, with 3-7 vascular bundles. Inflorescence
shoots sympodially branched, spathe stalked, re-
sembling vegetative leaves. Dioecious; male or fe-
male flowers in a zig-zag arrangement on the
spadix. Male spathe with retinacula inserted
alongside flowers. Female spathe with flowers al-
ternating with retinacula. Fruit crescent-shaped.
Five spp., temperate northern Pacific Ocean.
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Index to Scientific Names
References to main entries in bold-faced print, to illustrations in italics.

Abildgaardia 167 Amauriella 56 A. madagascariensis 22

Abildgaardieae 151, 166 Ambrosina 66 A. ulvaceus 24
Abolboda 468 Ambrosineae 66 A. undulatus 22
A. ciliata 462 Amischophacelus 120 Aponogetonaceae 1, 14, 21,
Abromeitiella 89 Amischotolype 111, 112, 114, 450
Acanthophoenix 374 120 Appertiella 245
Acanthostachys 75, 81, 95 Ammandra 387 Araceae 1, 26
Achasma 492 A. decasperma 318 Araeococcus 78, 93
Achlyphila 467 Amomum 480, 492 Arales 1
Acion 439 Amorphophallus 35, 64 Aratitiyopea 468
Acoelorraphe 331 Amphibolis 136, 137, 138 Archontophoenicinae 359
A. wrightii 308 Amphiphyllum 421 Archontophoenix 360
Acoraceae 1, 7 Amphiscirpus 164 Areae 66
Acorus 10, 30, 32, 37, 38 Amydrium 47 Areca 367
A. calamus 8 Anadendreae 46 A. abdulrahmii 318
Acrocomia 383 Anadendrum 46 A. caliso 318
Actinokentia 361 Ananas 95 Arecaceae 306
Actinoschoenus 174 A. comosus 81, 83 Areceae 354
Actinoscirpus 165 A. lucidus 83 Arecinae 365
Actiorhytis 372 Anaphyllopsis 49 Arecoideae 351
Actoplanes 289 Anaphyllum 49 Arenga 352
Adelmeria 489 Anarthria 20 Aridarum 57
Aechmea 95, 75, 78, 79, 82 A. prolifera 20 Ariopsis 70
Aetheolirion 119 A. scabra 19 Arisaema 33, 35, 69
Aframomum 480, 493 Anarthriaceae 4, 19, 196, 431 A. amurense 69
A. melegueta 480 Anchomanes 62 A. griffithii 69
Afrocalathea 290 Ancistrophyllinae 339 A. thunbergii 69
Afrorhaphidophora 47 Androlepis 75, 78, 96 Arisaemateae 69
Afrotrilepis 183 Andromycia 53 Arisareae 66
Aglaodorum 62 Androtrichum 169 Arisarum 33, 66
Aglaonema 62 Aneilema 110-112, 114, 126 Aristolochiaceae 9
Aglaonemateae 62 A. sebitense 112 Aroideae 51
Aiphanes 319, 383 Anepsias 48 Arophyteae 65
Alcantarea 82, 91 Anigozanthus 221 Arophyton 65
Alexgeorgea 430, 440 A. flavidus 213 Aropsis 55
Aliniella 171 A. uliginosus 213 Arthrostylis 173
Alinula 142, 171 A. humilis 213 Arum 33,67
Alisma 16 A. anglesii 213 A. korolkowii 67
Alismataceae 1, 11, 24, 101, 273 Anosporum 170 Ascolepis 171
Alismatales 1 Anthericopsis 112, 125 Ascopholis 171
Allagoptera 380 Anthochortus 437 Askidiosperma 435
Allardtia 92 Anthracoidea 153 Astelia 224
Alloschemone 48 Anthurieae 45 Asterochaete 177
Alloschmidia 372 Anthurium 34, 46 Asterogyne 385
Alocasia 71 Antongilia 357 A. martiana 310
Alpinia 480, 489 Anubiadeae 56 Asterostigma 53
A. sect. Geocharis 493 Anubias 56 A. luschnathianum 54
A. subgen. Geostachys 494 Apalanthe 244 Astonia 17
A. galanga 480 Aphandra 387 Astrocaryum 384
A. glabra 489 Aphelia 108 Ataenidia 287
A. purpurata 480 Aphyllarum 60 Attalea 381
A. speciosa 480 Apodasmia 443 A. exigua 382
Alpinieae 488 Aponogeton 25 A. maripa 382
Alsmithia 371 A. distachyos 22 A. speciosa 382
Althenia 473 A. elongatus 23 Attaleinae 381
504 Index to Scientific Names

Aulotandra 493 Calameae 339 Centrolepidaceae 4, 106, 233

Ayensua 87 Calaminae 340 Centrolepis 108
Calamoideae 337 C. aristata 107
Bactridinae 382 Calamus 308, 314, 342 Cephalocarpus 182
Bactris 383 C. subgen. Eremospatha 339 C. dracaenula 175
B.concinna 318 C. subgen. Laccosperma 339 Cephalostemon 423
Baeothryon 164, 166 C. subgen. Oncocalamus 345 Ceratocaryum 436
Balaka 364 Calathea 288 Ceratolobus 343
Baldellia 16 Caldesia 17 C. pseudoconcolor 310
Baloskion 439 C. parnassifolia 14 Cercestis 63
Barberetta 219 Calectasia 191, 193 Ceroxyleae 349
Barcella 381 C. cyanea 192 Ceroxyloideae 346
Basselinia 373 Calectasiaceae 190 Ceroxylon 349
Baumea 180 Calla 30, 35, 51 C. alpinum 348
Baxteria 191, 194 Callisia 112, 123 Chaetanthus 442
Baxteriaceae 190 Calloideae 51 Chamaedorea 350
Beccariophoenicinae 378 Callopsideae 64 C. pinnatifrons 308
Beccariophoenix 379 Callopsis 64 C. schiedeana 310
Belosynapsis 120 Calopsis 435 Chamaegyne 166
Bentinckia 370 Calorophus 437 Chamaerops 328
Biarum 27, 68 Calospatha 342 C. humilis 327
Billbergia 79, 82, 94 Calyptrocalyx 362 Chambeyronia 360
B. nutans 94 Calyptrocarya 185 Chelyocarpus 326
Bisboeckelera 184 Calyptrogyne 385 C. ulei 310, 327
Bisboeckelereae 152, 184 C. subgen. Calyptronoma 385 Chevaliera 95
Bismarckia 337 C. subgen. Pholidostachys 384 Chillania 166
Blancoa 221 Calyptronoma 385 Chlorospatha 60
Blastocaulon 206 Campecarpus 372 Chondropetalum 435
Blysmopsis 172 Campelia 123 Chordifex 440
Blysmus 173 Camptandra 483 Chorizandra 161
Blyxa 244 Canistropsis 97 Chrysalidocarpus 357
B. aubertii 238 Canistrum 79, 96 Chrysitricheae 151, 161
Boesenbergia 480, 486 C. aurantium 79 Chrysitrix 147, 162
Bognera 52 Canna 105 Chuniophoenix 332
Bolboschoenus 165 C. indica 104 Cienkowskia 486
Borasseae 334 Cannaceae 2, 103, 229, 285, 480 Cienkowskiella 486
Borassodendron 335 Cannomois 436 Cintractia 153
Borassus 335 Capitularina 160 Cladium 142, 153, 180
Brachychilum 482 Carex 142, 147, 187 Clinosperma 370
Brahea 331 C. bequaertii 142 Clinostigma 370
Brassiophoenix 365 C. dioica 187 C. sect. Lepidorrhachis 369
Brewcaria 82, 89 C. physodes 187 Coccothrinax 328
Brocchinia 75, 86 C. vulpina 187 Cochliostema 110-112, 114,
Brocchinieae 86 Cariceae 152, 186 121
Bromelia 75, 82, 83, 94 Caricoideae 152, 186 Cocoeae 377
Bromeliaceae 3, 74, 393, 467 Carlephyton 65 Cocos 380
Bromeliales 3 Carpentaria 363 C. nucifera 320
Bromelioideae 92 Carpha 177 C. subgen. Butia 379
Brongniartikentia 369 Carpoxylon 371 Coleocarya 438
Bucephalandra 57 Carptotepala 205 Coleochloa 183
Buforrestia 111, 124 Cartonema 110, 116, 118 Coleogeton 413
Bulbostylis 167 Cartonemataceae 115 Coleotrype 114, 120
Burbidgea 491 Cartonemateae 118 Coleotrypinae 120
Burnatia 18 Cartonematoideae 118 Colletogyne 65
Burretiokentia 373 Caryota 352 Colocasia 71
Butia 379 C. mitis 310 C. esculenta 61
B. capitata 310 Caryoteae 351 Colocasieae 70
Butiinae 379 Catacolea 439 Colpothrinax 331
Butomaceae 1, 14, 24, 100, 273 Catoblastus 354 C. wrightii 308
Butomopsis 101, 274 Catopsis 79, 81, 90 Comanthera 205
Butomus 102 C. berteroana 90 Cominsia 292
B. umbellatus 101 Caulokaempferia 485 Commelina 110-112, 114, 126
Caustis 153, 180 Commelinaceae 82, 109, 218, 295,
Caladieae 59 Cautleya 484 419
Caladiopsis 60 C. spicata 484 Commelinales 3
Caladium 60 Cenolophon 489 Commelineae 124
 Index to Scientific Names 505

Commelinoideae 118 Cyrtostachys 361 E. subgen. Barcella 381

Commelinopsis 126 Cytogonidium 441 E. guineensis 382
Connellia 88 Elasis 122
Conostylidoideae 220 Daemonorops 342 Elegia 434
Conostylis 221 D. sparsiflora 318 Eleiodoxa 342
C. aurea 213 Damasonium 16 Eleocharideae 151, 166
Copernicia 331 D. polyspermum 12 Eleocharis 145, 147, 166
C. rigida 31 0 Dapsilanthus 443 Eleogiton 168
Corypha 314, 332 Dasypogon 191, 193 Elettaria 494
C. umbraculifera 308 D. bromeliifolius 192 E. cardamomum 480
Corypheae 224 Dasypogonaceae 2, 190 Elettariopsis 493
Coryphinae 332 Dasypogonales 2 Eimeria 489
Coryphoideae 223 Deckenia 374 Elodea 245
Costaceae 2, 128, 229, 480 Deinacanthon 75, 95 E. canadensis 236
Costularia 178 Dendropogon 92 Elyna 186
Costus 131 Desmocladus 438 Eminium 66
C. cuspidatus 129 Desmoncus 383 E. alberti 67
C. lucanusianus 129 Desmoschoenus 153, 168 Empodisma 438
C. megalobractea 129 Deuterocohnia 75, 89 Encholirium 75, 89
Cottendorfia 82, 87 Diandriella 56 Enhalus 247
Courtoisina 142, 170 Dichorisandra 112, 114, 121 E. acoroides 237
Craspedolepis 435 Dichorisandrinae 121 Ensete 300
Crosslandia 167 Dichromena 174 E. ventricosum 297
Cryptangieae 152, 181 Dictyocaryum 353 Epidryos 422
Cryptangium 181 Dictyosperma 371 Epipremnopsis 47
Cryptanthus 75, 79, 93 Dictyospermum 112, 125 Epipremnum 47
Cryptocoryne 33, 34, 58 Didymiandrum 182 Epischoenus 178
Crysophila 326 Dieffenbachia 52 Eremospatha 339
Ctenanthe 291 Dieffenbachieae 52 Eriocaulaceae 3, 116, 197, 419, 467
Ctenophrynium 291 Dielsia 440 Eriocaulon 202, 203, 204
Culcasia 63 Dilatris 219 E. longipetalum 198
Culcasieae 62 D. pilansii 213 E. setaceum 198
Curculigo seychellensis 309 Dimerocostus 131 E. sieboldianum 198
Curcuma 480, 484 Diplacrum 147, 184 E. ussuriense 198
Curcumorpha 486 Diplasia 161 E. woodii 198
Cuthbertia 123 Disteganthus 97 Eriophorella 164
Cyanotinae 120 Distichia 258 Eriophorum 163
Cyanotis 111, 112, 120 D. muscoides 253 Erioscirpus 163
Cyathochaeta 142, 147, 176 Distichochlamys 486 Eriospora 183
Cyathocoma 178 Donax 289 Etlingera 492
Cyclanthaceae 309, 321 Dovea 435 E. elatior 480
Cyclospathe 348 Dracontioides 49 Eugeissona 340
Cyclospatheae 348 Dracontium 49 Eugeissoninae 339
Cymbispatha 123 D. sect. Urospathopsis 49 Eurychorda 440
Cymodocea 137, 138 Dracunculus 67 Eurystemon 402
C. nodosa 133 Drymophloeus 363 E. mexicanum 397
Cymodoceaceae 2, 133, 406, 447, Duckea 423 Euterpe 358
499 Dulichieae 152, 173 Euterpeinae 358
Cymophyllus 188 Dulichium 173 Evandra 148, 180
Cyperaceae 3, 141, 257, 456 Dyckia 75, 89 Everardia 182
Cypereae 151, 152, 168 Dypsidinae 357 Exocarya 160
Cyperoideae 151, 162 Dypsis 357 Exochogyne 148, 185
Cyperus 145, 146, 170 D. lutescens 308
C. bequaertii 142 Fascicularia 93
C. boreochrysocephalus 142 Ecdeiocolea 196 Felipponia 53
C. halpan 142 Ecdeiocoleaceae 4, 20, 195, 431 Felipponiella 53
C. papyrus 154 Echidnium 49 Fernseea 93
C. pedunculatus 142 Echinodorus 16 Ficinia 146, 168
C. purpureo-glandulosus 142 Edmundoa 96 Filarum 59
C. usitatus 142 Egeria 245 Fimbristylis 166
Cyphokentia 370 E. densa 136 Fintelmannia 182
Cyphophoenix 372 Egleria 166 Flagellaria 210, 224, 250
Cyphosperma 373 Eichhornia 401 F. guineensis 208
Cyphostigma 494 E. crassipes 396 F. indica 208, 209
Cyrtosperma 50 Elaeidinae 381 Flagellariaceae 4, 208, 431
Cyrtostachydinae 361 Elaeis 382 Flagellarisaema 69
506 Index to Scientific Names

Floscopa 111, 124 H. ova/is 237 H. verschaffeltii 348

Fosterella 75, 88 Hanguana 209, 225, 250 Hyophorbeae 350
F. penduliflora 77 H. malayana 223 Hyospathe 359
Fuirena 146, 164 Hanguanaceae 3, 210, 223 Hyphaene 336
F. leptostachya 142 Haniffia 483 H. thebaica 308
Fuireneae 151, 164 Hapaline 61 Hyphaeninae 335
Furtadoa 55 Haplochorema 486 Hyplophytum 95
Harperia 438 Hypodiscus 437
Gagnepainia 488 Haumania 292 H. procurrens 426
Gahnia 142, 179 Hechtia 75, 82, 90 Hypolaena 437, 442
Gaimardia 108 Hedychium 480, 482 Hypolytreae 151, 159
Gamogyne 57 H. cylindricum 483 Hypolytrum 160
Gastrochilus 486 Hedyscepe 360 Hypoxidaceae 218
Gastrococos 383 Heleocharis 166 Hypselodelphys 289
Gaussia 350 Helicodiceros 68
Geanthus 492 Heliconia 229, 454 Iguanura 369
Gearum 54 H. solomonensis 227 Iguanurinae 367
Geocharis 493 H. spathocircinata 228 Iriartea 319, 353
Geogenanthus 112, 121 Heliconiaceae 2, 226 Iriarteeae 352
Geonoma 385 Hellmuthia 162 Iriartella 353
G. macrostachys 318 Helmholtzia 393 I. setigera 318
Geonomeae 384 Helonema 166 Ischnosiphon 288
Georgeantha 196 Hemicarpha 172 Ischyrolepis 434
G. hexandra 195, 196 Hemiorchis 488 Isolepis 168
Geostachys 494 Hemithrinax 327 I. jluitans 142, 163
Gibasis 123 Heteranthera 402 Itaya 327
Gibasoides 122 H. zosterifolia 397
Glaziova 380 Heteroaridarum 58 ]aimenostia 68
Globba 487 Heteroarisaema 69 ]asarum 60
G. muluensis 487 Heterolobium 51 ]essenia 359
Globbeae 487 Heteropsideae 46 ]ohannesteijsmannia 330
Glomeropitcairnia 81, 91 Heteropsis 46 f. altifrons 310
Gonatanthus 70 Heterospathe 369 ]oinvillea 210, 224, 250
Gonatopus 51 Heterozostera 500 f. ascendens 249
G. angustus 31 Hitchenia 482 f. plicata 249
Goniocladus 373 Helicodea 94 Joinvilleaceae 4, 20, 196, 249,
Gorgonidium 53 Hohenbergia 82, 95 431
Gramineae 4 H. stellata 80 ]uania 349
Greigia 93 Hohenbergiopsis 78, 96 ]ubaea 379
G. berteroi 82 Holochlamys 46 ]ubaeopsis 379
Groenlandia 414 Holoschoenus 169 Juncaceae 3, 152, 252, 257, 456
G. densa 409 Homalomena 32, 35, 56 Juncaginaceae 1, 260, 262, 450
Gronophyllum 318, 366 Homalomeneae 55 Juncales 3
Guacamaya 422 Hopkinsia 429, 431, 434 ]uncella 233
Guihaia 329 Hornstedtia 493 ]uncellus 170
Guilielma 383 Hottarum 57 ]uncus 153, 259
Gulubia 366 Howea 362 J. balticus 253
Guringalia 439 Hybophrynium 289 f. triglumis 253
Guzmania 79, 91 Hydatella 108, 233
G. acrifolia 80 H. dioica 232 Kaempferia 480, 485
Gymnoschoenus 176 Hydatellaceae 4, 108, 231 K. subgen. Cienkowskia 486
Gymnostachydoideae 43 Hydriastele 366 K. subgen. Pyrgophyllum 485
Gymnostachys 27, 35, 37, 44 Hydrilla 245 Karinia 169
G. anceps 31 H. verticillata 236 Kentiopsis 361
Hydrocharis 244 Kerriodoxa 332
Hadrodemas 123 H. morsus-ranae 238 Kingia 191, 193
Haemodoraceae 3, 116, 212, 393, Hydrocharitaceae 1, 234, 305 K. australis 191
400 Hydrocleys 14, 101, 274 Kingiaceae 190
Haemodoroideae 219 H. martii 273 Kobresia 142, 153, 186
Haemodorum 219 H. nymphoides 272 Koernickanthe 290
H. spicatum 213 Hydrophilus 436 Kolowratia 489
Hagenbachia 218 Hydrothrix 402 Korthalsia 340
Halmoorea 355 H. gardneri 397 Koyamaea 185
Halodule 138 Hylaeanthe 291 Kulinia 439
Halopegia 292 Hymenochaeta 165 Kunhardtia 422
Halophila 247 Hyophorbe 350 Kyllinga 148, 172
 Index to Scientific Names 507

K. sphaerocephala 142 Lophiola 218 Medemia 337

Kyllingiella 169 Lophoschoenus 178 Meeboldina 442
Lophospatha 342 Megalotheca 441
Laccospadix 362 Lowiaceae 2, 229, 275, 299 Megaphrynium 289
Laccosperma 339 Loxocarya 441 Melanostachya 441
Lachnanthes 219 Loxococcus 365 Mesanthemum 203, 205
Lachnocaulon 205 Luronium 16 M. rutenbergianum 199
Lagarosiphon 246 L. natans 12 Mesomelaena 176
L. muscoides 236 Luzula 259 Metroxylinae 340
Lagenandra 33, 34, 58 L. pilosa 256 Metroxylon 314, 340
Lagenocarpus 181 L. spicata 253 M. amicarum 309, 341
Lamprocaulos 434 Lyginia 429, 431, 434 M. sagu 341
Lamprococcus 95 L. barbata 426 M. salomonense 309, 318, 341
Lanaria 218 L. imberbis 426 M. vitiense 341
Languas 489 Lymania 97 M. warburgii 341
Lasia 50 L. corallina 80 Mezbromelia 91
Lasimorpha 50 Lysichiton 35, 44 Microcasia 57
Lasioideae 49 L. camtschatcensis 44 M. sect. Truncatae 57
Latania 335 Lytocaryum 380 Microculcas 51
Lataniinae 335 Microdracoides 183
Lavoixia 372 Machaerina 147, 180 M. squamosus 183
Lazarum 35, 68 Machaerocarpus 16 Micropapyrus 174
Leiothrix 205 Mackeea 361 Moldenkeanthus 206
L. nubigena 199 Macrochaetium 178 Monochoria 401
Lemna 269 Macrochordion 95 M. vagina/is 396
L. perpusilla 265 Maidenia 246 Monocostus 131
L. trisulca 265 M. rubra 237 Monophrynium 287
Lemnaceae 1, 38, 264 Malortieinae 356 Monophyllanthe 290
Lemnoideae 269 Mangonia 53 Monotagma 289
Lemurophoenicinae 357 Manicaria 356 M. laxum 280
Lemurophoenix 357 Manicariinae 356 M. secundum 280
Leopoldinia 356 Mantisia 488 Monotrema 423
Leopoldiniinae 356 Mapania 145, 160 Monotremeae 423
Lepidobolus 439 Mapanioideae 151, 159 Monstera 48
Lepidocaryeae 345 Mapaniopsis 160 M. obliqua 48
Lepidocaryum 345 Marahuacaea 422 Monstereae 47
Lepidorrhachis 369 Maranta 290 Monsteroideae 46
Lepidosperma 142, 181 M. subgen. Saranthe 291 Mimtrichardia 63
Lepilaena 473 M. amazonica 280 M. arborescens 63
Lepironia 153, 161 M. arundinacea 280 Montrichardieae 63
Leptocarpus 435, 442 Marantaceae 2, 105, 229, 278, 480 Moratia 370
L. elegans 427 Marantochloa 290 Morelotia 179
L. tenax 430 Mariscopis 172 Morenia 350
Leptorhoeo 123 Marisculus 171 Murdannia 110, 112, 124
Leptosolena 492 Mariscus 170 M. cryptantha 111
Lepyrodia 437 Marojejya 377 M. simplex 111
Licuala 330 Marsippospermum 258 Musa 300
Lilaea 263 M. grandiflorum 253 M. acuminata 297
Lilaeaceae 262 Maschalocephalus 424 M. maclayi 297
Lilloa 54 Masoala 376 Musaceae 2, 229, 277, 296, 454
Limnobium 244 Masoalinae 376 M. subfam. Heliconioideae 226
L. laevigatum 238 Mastersiella 437 M. tribe Heliconieae 226
Limnocharis 101, 274 Matudanthus 122 Myrialepis 343
L. flava 272 Maundia 263 Myrosma 291
Limnocharitaceae 1, 14, 101, 271 Mauritia 345
Limnophyton 17 M. flexuosa 310 Najadaceae 2, 242, 301
Lindmania 87 Mauritiella 345 Najas 305
Linospadicinae 361 Maxburretia 329 N. podostemon 302
Linospadix 362 Maximiliana 381 Nannorrhops 332
Lipocarpha 147, 172 Mayaca 295 N. ritchiana 327
L. nana M. baumii 295 Nanochilus 491
Livistona 329 M. fluviatilis 295 Navia 81, 82, 86
Livistoninae 329 M. longipes 295 N. acaulis 87
Lodoicea 335 M. sellowiana 295 N. fontoides 77
L. maldivica 336 M. vandellii 295 Nechamandra 246
Lomandra 191 Mayacaceae 3, 82, 116, 294, 419 N. alternifolia 236
508 Index to Scientific Names

Neesenbeckia 148, 178 Parajubaea 380 Pitcairnieae 86

Nelmesia 168 Parakaempferia 487 Pitcairnioideae 86
Nematanthus 437 Paramapania 148, 161 Plagiostachys 490
Nemum 167 Paramomum 492 P. albiflora 490
Nenga 366 Patosia 258 P. strobilifera 490
Neodypsis 357 Pauella 68 Platycaulos 435
Neoglaziovia 75, 94 Pauridia 218 Platychorda 440
N. variegata 83 Pedicellarum 45 Plectocomia 344
Neonicholsonia 358 Pelagodoxa 369 Plectocomiinae 343
Neophloga 357 Peltandra 35, 65 Plectocomiopsis 344
Neoregelia 75, 82, 97 P. virginica 30 Pleiostachya 288
Neoveitchia 368 Peltandreae 65 P. pruinosa 280
Nephrosperma 376 Pepinia 81, 88 Plesmonium 64
Nephthytideae 61 Phacelophrynium 288 Pleuranthodium 491
Nephthytis 34, 61 Phaeomeria 492 Pleuriarum 69
Nevillea 436 Phaethorninae 228 Pleurostachys 175
Nicolaia 492 Phelepsiella 421 Poaceae 20, 152, 196, 431
Nidularium 97 Phenakospermum 454 Poales 4
N. innocentii 77 Philodendreae 55 Podaechmea 93, 95
N. scheremetiewii 94 Philodendron 33, 55 Podococceae 354
Nipa 346 Philodice 205 Podococcus 354
Normanbya 364 P. hoffmannseggii 199 P. barteri 348
Nypa 312, 346 Philonotion 56 Podolasia 50
N. fruticans 310, 318, 320, 347 Philydraceae 3, ll6, 218, 389, 400 Pogonotium 343
Nypoideae 346 Philydrella 394 Pollia 112, 125
Philydrum 394 Polyandrococos 381
Ochagavia 93 P. lanuginosum 390 Polyspatha 111, 112, 114, 125
0. elegans 82 Phlebocarya 220 Pommereschea 492
Oenocarpus 359 Phloga 357 Pontederia 401
Oncocalaminae 345 Phoeniceae 33 P. cordata 396
Oncocalamus 345 Phoenicophorium 375 Pontederiaceae 3, ll6, 218, 393,
Oncosperma 375 Phoenix 333 395
Oncospermatinae 374 P. canariensis 334 Porandra 112, 114, 120
Onychosepalum 438 P. dactylifera 319, 334 Porphyrospatha 60
Orania 349, 355 P. reclinata 334 Portea 96
Oraniinae 355 P. roebelenii 310 Posidonia 407
Oraniopsis 349 P. sylvestris 334 P. australis 404
Orbignya 381 Pholidocarpus 330 Posidoniaceae 2, 136, 404, 406,
Orchidantha 277 Pholidostachys 384 447, 499
0. borneensis 276 Phrynium 287 Potamogeton 413
0. longijlora 276 Phyllocomos 437 P. alpinus 409
0. maxillarioides 276 Phylloscirpus 163 P. gramineus 409
Orectanthe 468 P. acaulis 163 P. lucens 409
0. sceptrum 467 Phyllospadix 500 P. natans 409
Oreobolopsis 163 P. scouleri 497 P. perfoliatus 409
Oreobolus 177 Phymatarum 57 Potamogetonaceae 2, 408, 447, 499
0. obtusangulus 175 Physocaulis 300 Potarophytum 423
Oreograstis 177 Physokentia 373 Potheae 45
Orontioideae 44 Phytelephantoideae 385 Pothoideae 45
Orontium 34, 44 Phytelephas 386 Pothoidium 45
Ortgiesia 95 P. macrocarpa 386 Pathos 35, 45
Orthophytum 79, 92 Pigafetta 344 Pothuava 95
Orthothylax 393 Pigafettinae 344 Prestoea 358
Ottelia 243 Pinanga 367 Principes 2
0. alismoides 238 P. maculata 318 Principina 160
Oxycaryum 169 Pinellia 32, 69 Prionium 258
Oxychloe 259 Piperaceae 9 P. serratum 253
Piptospatha 57 Prionophyllum 89
Paepalanthus 202, 206 Pistia 27, 30, 35, 38, 71, 268 Prionosepalum 442
Palandra 386 P. stratiotes 31 Pritchardia 330
Palisota llO, ll2, 119 Pistieae 71 Pritchardiopsis 330
Palisotinae 119 Pitcairnia 75, 79, 88 Protarum 70
Palmae 2, 306 P. andreana 79 Pseudaechmea 96
Pandanaceae 321, 459 P. feliciana 82 Pseudalthenia 473
Panzhuyia 71 P. grafii 80 Pseudananas 75, 95
Paracautleya 485 P. mirabilis 88 Pseudodracontium 65
 Index to Scientific Names 509

Pseudoeverardia 182 Rhektophyllum 63 Schizocasia 71

Pseudohomalomena 64 Rhodocoma 436 Schoendoum 442
Pseudohydrosme 62 Rhodospatha 48 Schoeneae 152
Pseudoparis 125 Rhoeo 123 Schoeneae 173
Pseudophoenix 348 Rhopalephora 111, 112, 126 Schoenocephalieae 422
P. sargentii 348 Rhopaloblaste 371 Schoenocephalium 422
Pseudoschoenus 165 Rhopalostylis 361 Schoenoides 177
Pseudowolffia 269 Rhynchanthus 491 Schoenoplectus 147, 153, 154, 165
Psilocarya 174 Rhynchocladium 180 S. lateriflorus 163
Psychanthus 491 Rhynchospora 145, 148, 174 Schoenoxiphium 153, 186
Pteroscleria 184 R. corymbosa 146 S. sparteum 142
Ptilanthelium 175, 176 R. latifolia 175 Schoenus 142, 175
Ptilothrix 176 Riedelia 491 Scholleropsis 402
Ptychococcus 365 Rikliella 172 Schumannianthus 289
Ptychosperma 364 Ringentiarum 69 Schunda-Pana 352
Ptychospermatinae 363 Rondonanthus 202, 203, 204 Scindapsus 47
Puccinia 153 Ronnbergia 97 Scirpeae 151, 162
Puya 75, 89 Roscheria 375 Scirpodendron 147, 159
P. berteroiana 88 Roscoea 480, 484 S. ghaeri 159
P. spathaceae 80 Rostkovia 258 Scirpoides 169
Puyeae 89 Roystonea 359 Scirpus 162
Pycnospatha 49 R. regia 308 Scleria 148, 184
Pycreus 148, 172 Roystoneinae 359 S. distans 146
Pyrgophyllum 485 Ruppia 448 S. hildebrandtii 146
Pyrrorhiza 219 R. maritima 446 S. nyasensis 146
Ruppiaceae 2, 445 Sclerieae 152, 184
Queenslandiella 142, 148, 172 Sclerioideae 152, 152, 181
Quesnelia 96 Saba! 333 Sclerosperma 376
S. etonia 308 S. mannii 318
Racinaea 92 S. palmetto 310, 327 Sclerospermatinae 376
R. insularis 82 Sabalinae 333 Separotheca 123
Ranalisma 16 Sagittaria 17 Serenoa 331
R. rostratum 12 S. lappula 13 S. repens 308, 310
Rapatea 422 S. sagittifolia 13 Siderasis 121
R. celiae 418 S. teres 13 Siliquamomum 483
R. membranacea 417 Sagus 344 Siphokentia 366
Rapateaceae 3, 82, 116, 193, 415, S. sect. Pigafetta 344 Siphonochilus 486
467, 422 Salacca 342 Socratea 319, 353
Rapateoideae 422 S. sect. Eleiodoxa 342 S. exorrhiza 348
Raphia 344 S. wallichiana 308 S. salazarii 310
Raphidophora 47 Sanblasia 288 Sodiroa 91
Raphiinae 344 Saranthe 291 Sommieria 370
Rautanenia 18 Sarcophrynium 290 Sorostachys 170
Ravenala 454 Saropsis 440 Sparganiaceae 457
R. subgen. Phenakospermum 454 Satakentia 371 Sparganium 460
R. subgen. Urania 454 Satranala 337 S. emersum 458
R. madagascariensis 452 Sauromatum 35 S. erectum 458
Ravenea 349 Sauromatum 68 Spathantheum 54
R. musicalis 320 S. venosum 31 Spathanthus 423
Reedia 179 Sauvallea 123 Spathicarpa 55
Reinhardtia 357 Saxofridericia 421 Spathicarpeae 53
Remirea 170 S. grandis 417 Spathiphylleae 46
R. maritima 142 S. petiolata 417 Spathiphyllum 35, 46
Remusatia 35, 70 S. spongiosa 417 S. cochlearispathum 31
Renealmia 490 Saxofridericieae 421 Spatholirion 119
Restio 435, 439-441 Saxofridericioideae 421 Sphaerocyperus 171
R. bolusii 430 Scaphispatha 59 Spirodela 38, 269
Restionaceae 4, 20, 108, 152, 196, Scaphochlamys 485 S. polyrhiza 265
203, 250, 425 Scheelea 381 Sporadanthus 437
Retispatha 343 Scheuchzeria 450 Staberoha 434
Reussia 401 S. palustris 449 Stachyphrynium 288
R. rotundifolia 396 Scheuchzeriaceae 1, 262, 449 Stadiochilus 494
Rhaphidophora 35, 47 Schieckia 220 Stahlianthus 486
Rhapidophyllum 328 Schippia 327 Stanfieldiella 111, 124
Rhapis 329 Schismatoglottideae 56 Stegolepis 422
R. excelsa 313 Schismatoglottis 31, 35, 56 S. angustata 420
510 Index to Scientific Names

Stenospermation 48 Tillandsia 75, 92 Veillonia 373

S. spruceanum 48 T. subgen. Diaphoranthema 75 Veitchia 364
Stenotalis 442 T. subgen. Diaphoranthema SO Velloziaceae 82, 393
Steudnera 70 T. subgen. Tillandsia 80 Verschaffeltia 375
Steyerbromelia 87 T. bertramii 92 Vesicarex 187
Stickmannia 168 T. castellanii 79 Vignea 187
Stratiotes 242, 244 T. cyanea 79 Vleisia 473
Strelitzia 454 T. flexuosa 77 Voanioala 380
Strelitziaceae 2, 229, 299; 451 T. jucunda 80 Voladeria 177, 259
Streptocalyx 97 T. recurvata 77 Volkiella 142, 172
Streptoliriinae 119 T. usneoides 81, 83 Vonitra 357
Streptolirion 111, 119 T. utriculata 92 Vriesea 75, 91
Stromanthe 291 Tillandsioideae 90 V. psittacina 79
Stuckenia 413 Tinantia 114, 122
Stylochaeton 52 Tonina 206 VVachendorjla 220
Stylochaetoneae 51 T. jluviatilis 199 W. thyrsijlora 213
Sumatroscirpus 173 Torulinium 170 VVallichia 352
Susum 225 Trachycarpus 328 VVashingtonia 331
Syagrus 380 Trachyphrynium 289 W. fllifera 308
Symplocarpus 30, 44 T. subgen. Hypselodelphys 289 VVebsteria 166
S. foetidus 44 T. subgen. Lasiodelphys 292 W. confervoides 163
Synandrospadix 54 Trachystylis 174 VVeldenia 111, 122
S. vermitoxicus 30 Tradescantia 123 VVelfia 385
Synechanthus 350 T. sect. Cymbispatha 111 VVendlandiella 351
S. warscewiczianus 348 Tradescantiinae 123 VVerauhia 75, 91
Syngonanthus 203, 205 Tremulina 441 W. gigantea 79
S. anthemidijlorus 199 Trianoptiles 142, 177 VVettinia 354
S. xeranthemoides 202 Tribonanthes 221 W. quinaria 348
Syngonium 34, 60 T. variabilis 213 VViesneria 17
Syntrinema 174 Tricarpelema 125 VVilldenowia 437
Syringodium 137 Triceratella 110, 118 VVindsorina 423
Triceratelleae 118 VVinifredia 438
Taccarum 53 Trichophorum 162, 164 VVittrockia 96
Tapeinochilos 131 Trichoschoenus 174 VVodyetia 364
Tapheocarpa 112, 126 Tricostularia 181 VVolffia 269
Taraxis 441 Triglochin 263 W. arrhiza 265
Tectiphiala 375 T. calcitrapa 261 W. microscopica 265
Tetraria 148, 177 T. palustre 261 VVolffiella 269
Tetroncium 263 Trilepideae 152, 182 W. denticulata 266
T. magellanicum 261 Trilepis 147, 182 Wolffioideae 269
Thalassia 247 Tripogandra 110-112, 123 VVolffiopsis 269
T. testudinum 237 Trithrinax 326 VVurdackia 204
Thalassocharis 137 Trithuria 108, 233
Thalassodendron 136, 137, 138 T. macranthera 232 Xanthorrhoea 191
T. ciliatum 133 T. submersa 232 Xanthorrhoeaceae 193
Thalia 292 Tylocarya 166 Xanthosoma 60
T. pavonii 282 Typha 460 X. platylobum 61
Thamnochortus 436 T. angustifolia 459 X. sagittifolium 61
Thaumatococcus 292 T. latifolia 459, 460 Xenophya 71
Thaumatophyllum 55 Typhaceae 4, 400, 457 Xiphidium 220
Theriophonum 68 Typhales 3 X. coeruleum 213
T. minutum 30 Typhonium 68 Xyridaceae 3, 116, 203, 295, 419,
Thomsonia 64 Typhonodorum 35, 66 461
Thomsonieae 64 Tyrbastes 441 Xyridales 3
Thoracostachyum 160 Xyris 202, 468
Thrinacinae 326 Ulearum 59 X. involucrata 465
Thrinax 327 Uncinia 153, 186
T. radiata 327 Urania 454 Zamioculcadeae 51
Thurnia 257, 456 Urospatha 50 Zamioculcas 38, 51
T. jenmanii 456 Urospathella 50 Z. zamiifolia 52
T. polycephala 456 Ursulaea 93 Zannichellia 473
T. sphaerocephala 456 Ustilaginales 153 Z. palustris 471
Thurniaceae 3, 152, 455 Zannichelliaceae 2, 412, 470, 499
Thymocarpus 288 Vallisneria 246 Zantedeschia 64
Thyrsantheminae 121 V. spiralis 237 Zantedeschieae 63
Thyrsanthemum 122 Vanoverberghia 490 Zebrina 123
 Index to Scientific Names 511

Zingiber 488 Zingiberales 2 Zostera 500

Z. acuminatum 489 Zingibereae 488 Z. marina 497
Z. coloratum 489 Zombia 328 Zosteraceae 2, 136, 406, 497
Z. officinale 480 Zomicarpa 59 Zosterella 402
Zingiberaceae 2, 131, 229, 285, Zomicarpeae 58 Z. dubia 397
474 Zomicarpella 59 Zyganthera 62