Organization for Flora Neotropica

Pilocarpinae (Rutaceae)
Author(s): Roel C. Kaastra
Source: Flora Neotropica, Vol. 33, Pilocarpinae (Rutaceae) (Dec. 13, 1982), pp. 1-197
Published by: New York Botanical Garden Press on behalf of Organization for Flora
Neotropica
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FLORA NEOTROPIC

MONOGRAPHNUMBER 33
PILOCARPINAE
(Rutaceae)

by
Roel C. Kaastra

~C -\ f CANCER
?\TROPIC Of

FLORAL
NEOTROPICA/

TROPIC OF CAPRICORN

Published for

Organization for Flora Neotropica

by
The New York Botanical Garden

New York

Issued 13 December 1982

 Copyright ? 1982

The New York Botanical Garden

Published by
The New York Botanical Garden
Bronx, New York 10458
International Standard Serial Number 0071-5794

Library of Congress Cataloging in Publication Data

Kaastra, Roel, 1942-
Pilocarpinae(Rutaceae)
(Flora neotropica ; monographno. 33)
Bibliography:p.
Includes index.
1. Rutaceae-Classification. 2. Botany-Latin America-
Classification.I. Title. II. Series.
QK495.R98K32 1982 583'.24 82-14088
ISBN 0-89327-242-6

All material subject to this copyright may be photocopied for the non-commercial purpose of
scientific or educational advancement.
 A MONOGRAPH OF THE PILOCARPINAE (RUTACEAE)

ROEL C. KAASTRA1

TABLE OF CONTENTS
Introduction ......................................................... .. ......... 1
H isto ry ... ... ... ... . . ... .. ........ .. . .. . .... . . ............... . . ... .. .............. ... 2
Morphology .......................................... ................................ 3
Anatom y ..................................................... ....................... 9
Pollen and Chromosomes .............................................................. 16
Phytochemistry and Physiological Action ................................................ 17
Relationships and Phylogeny ........................................................... 20
D istribution ......................................... ...... ........................... 22
Systematic Treatment .............................................. .................. 22
Key to Genera ..................................................................... 24
1. Esenbeckia ......................... ..................... ........................ 24
2. Metrodorea ..................................................................... 116
3. Raulinoa .......................... .............................................. 130
4. Pilocarpus ... ................................................. ................. 132
Acknowledgments ......................................... ......................... 181
Literature Cited .................................... .................................. 182
Numerical List of Taxa ............................................................. 185
List of Exsiccatae .......................... ................................ .......... 186
Index of Local N ames ................................................................ 193
Index of Taxa ........................................................................ 194
Addenda ................................. ........................................... 197

INTRODUCTION
Up to now about 120 specific names have been publishedfor taxa belongingto
the subtribePilocarpinae.In this monograph46 species with 16 infraspecifictaxa
are accepted. The subtribe consists of four genera, one of which is monotypic.
In the other subtribe of the Cusparieae, the Cuspariinae, there are about 18
genera, most of them with few species.
The Pilocarpinaeare distinguisedby globular flower buds. The flowers have
yellowish or violet petals which are nearly regular and nearly free. They are
haplostemonic with free stamens and with heart-shapedanthers. The embryos
have plano-convex, eared cotyledons which are not folded. The plants occur in
the Neotropics with one species extending into the U.S.A. They grow in very
differenthabitats, from savannas and thickets to moist tropicalforests, from sea
level to 2500 m.
The subtribeis of great pharmaceuticalinterest because pilocarpinehas been
isolated from several species of Pilocarpus. Pilocarpineis an alkaloid which is
used mainlyin ophthalmology,as a parasympathetic.Some species of Esenbeckia
have also been used in medical practice. Because of that interest, some species
have become depleted in number,as large amountsof leaves and branchletshave
been sampled and exported, e.g. to the markets of Liverpool and Hamburgin
shipments of up to 10 tons (see Duval, 1905:62; Holmes, 1904)!For this reason
Pilocarpus jaborandi and P. microphyllus are in danger of extinction. Already in
1904,pilocarpinewas chiefly obtainedfrom P. microphyllus(Holmes, 1904).Ac-

1Marktweg 68, 8444 AH Heerenveen, Netherlands.

2 Flora Neotropica

cording to a letter from the world's largest factory of pilocarpine,the sources of

the alkaloid today are P. microphyllusand the related species P. trachylophus
and P. jaborandi. The plants are cultivated in Brazil but a significantamount of
the drug still comes from local dealers collecting from wild populations.
All measurementsreported in the descriptions, except for floral dimensions,
were made on dried herbariummaterial. The flowers were boiled and flower
diameterwas taken with the petals fully extended. Dried flowers are ? 15%small-
er than boiled flowers. The description of plane shapes follows the SADT-chart
(Taxon 11: 145-156, 1962).
The cited dates of effective publication follow Stafleu (1967) or Stafleu and
Cowan (1976), or are derived from data in the publications. In the citation of
synonyms at least the next publicationafter the valid one is given, and also the
first publicationin which it is reduced to synonymy. There are numerousillus-
trations of Pilocarpus in the pharmaceuticalliteratureand in non-botanicalpub-
lications which are not always cited. The illustrationsin Englerand Prantl(1931,
ed. 2) are usually not cited because they are reprintedfrom the first edition.
The articles of the International Code of Botanical Nomenclature (ICBN),
Seattle Code, which are applicableto invalidly publishednames are indicated.
In the citation of types and specimens "x" means sheets, e.g. F2x means-2
sheets seen from F.
Materialwas studied and/or informationwas obtainedfrom the following her-
baria:BAB, BM, BR, BRG, C, CGE, COL, CR, DS, E, F, G, GB, GE, GENT,
GH, GL, GOET, HAL, HBG, IAN, K, KRA, KW, L, LD, LE, LP, LY, M,
MICH, MO, MPU, NY, OXF, P, PHA, PR, R, RB, S, SP, TO, TRIN, U, UC,
US, VEN, VT, W, WAG, WIS, Z, ZT.
HISTORY
The first known record pertainingto the (sub)tribeis in Plumier's Catalogus
(1703), where Euonymuslatifolius, racemosus is mentioned.The drawingof this
plant in Plumier's manuscriptswas copied, recopied, and eventually printedfor
Burman in his Plantarum Americanarum (1757). Burman incorrectly added a
phrase to the plate, which he took from Linnaeus' Species plantarum. However,
it is clear that both the drawing and the phrase "Euonymus latifolius .. ,"
printedelsewhere in his book, are referableto Pilocarpus racemosus Vahl.
Pilocarpus Vahl (1797) is the first validly publishedgenus of the Pilocarpinae,
but Vahl did not place Pilocarpus in a higher category. This was attemptedby
de Candolle (1822), who thought it probably correct to place Pilocarpus in his
new tribe Diosmeae. De Candolle, in the same work, published the tribe Cus-
parieae which was florally very different from the other three tribes which he
recognized, the link being found in the fruits accordingto the author. Nees and
Martius(1823)publishedthe new family Fraxinellaeand attributedPilocarpus to
one of its two "sectios." Furthermorethey published the family Xanthoxyleae
and recognized the family Diosmeae R. Brown minus some genera which they
transferredto the Fraxinellae. In his Prodromus de Candolle (1824) recognized
only the family Rutaceae A. L. de Jussieu excluding sectio I, and placed the
families recognized by Nees and Martius(1823) into synonymy. He divided the
family into two tribes, the Diosmeae (comprisingthe three tribes he had published
in 1822),and the Cusparieae.The position of Pilocarpus, however, remainedas
doubtfulas it was when publishedin 1822.Dumortier(1829)confidentlyreferred
Pilocarpus to the Diosmeae.
In 1825KunthpublishedEsenbeckia and placed it in the Diosmeae R. Brown.
History 3

The genus Metrodorea was published in 1825 by Saint-Hilaire, although its

position was indicated by Jussieu (1825: 415, 484) who placed it, together with
Esenbeckia, Pilocarpus, and other genera in "sectio prima" of the Diosmeae
americanae. This sectio, however, was not yet satisfactorilyknown to the author.
Bartling (1830) place Pilocarpus, Esenbeckia, and Metrodorea in his tribe
Pilocarpeae(as "genus Pilocarpea"), together with the tribe Cusparieaede Can-
dolle and three non-Americantribes constitutingthe family Diosmeae R. Brown.
The partitionof the (American)Diosmeae into two tribes was also accepted by
Spach (1834), Lindley (1836, 1846), Endlicher(1840, 1841), and Walpers(1842).
Martius(1835) did not mention it, and Meisner (1837) definitelyrejected the two
"sectios" of Jussieu.
Hooker (1862)did not recognize the system of Bartling.He distinguishedsev-
eral tribes within the Rutaceae, and placed Esenbeckia (includingMetrodorea)
and Pilocarpus in the Zanthoxyleae. This was followed by Baillon (1873, 1884)
and by Durand(1888). In 1874Engler publishedhis Studien (1874b).He divided
the tribe Cusparieaede Candolleinto two subtribes:"Eucusparieae"and "Pilo-
carpeae." In the latter he placed Esenbeckia and Pilocarpus and restored Me-
trodorea to generic level. The same system was adopted by him in Flora brasi-
liensis (1874a) and in later studies, including those published in Engler and
Prantl's Die natiirlichen Pflanzenfamilien (1896, 1931). In these latter works he
amended the orthographyto Cuspariinaeand Pilocarpinae.Engler was followed
by Dalla-Torreand Harms and by modern compilers.
Recently Cowan (1960) added the new genus Raulinoa to the Pilocarpinae.
The increase in numbersof validly publishedtaxa belongingto the Pilocarpinae
is shown in Fig. 1.

MORPHOLOGY
Habit
The Pilocarpinaeare usually shrubs or small trees, mostly up to 10 or 12 m.
Subshrubsand "polypodial" shrubs with numerous ascending branches are re-
ported from dry regions:Esenbeckiapumila from the cerradosof mid-Brazil,and
E. hartmanii in northwestern Mexico. Frequent burning and grazing may be
responsible for this habit. Taller trees occur in E. berlandieri (to 15 m), E. pen-
taphylla subsp. belizensis (to 25 m), Metrodorea flavida (to 18 or rarely 27 m),
and Pilocarpus racemosus (in Venezuela reportedly to 20 m). The diameter of
the trunk is usually reported to 20(-50) cm. The bark is smooth, with minute
cracks.

Roots
Planchon (1875: 296) studied the roots of an unknown Pilocarpus. They are
pale yellow-orangewith the peridermscaling off in papery pieces. The cortex is
pale yellow with dark "dots" containing a resin-like substance, and has fiber
groups. The wood is satin-whiteand shows numerouscontorted fibers.

Spines
Raulinoa is characterizedby the presence of spines, althoughnot on all branch-
es. These spines are modifiedspurs, borne in the axil of the leaves or their scars,
and are usually opposite. They have nothing to do with the sheaths of Metro-
dorea, as Cowan (1960) suggested (Kaastra, 1978a).
4 Flora Neotropica

'
10 ~m species infraspecific taxa

I-

c2-

1800 1825 1850 1875 1900 1925 1950 1975-

-1799 1824 1849 1874 1899 1924 1949 1974
YEAR OF EFFECTIVE PUBLICATION -

FIG. 1. Dates of effective publication of the taxa recognized in the present study.

Leaves
In Pilocarpus the perules of the terminal leaf buds are remarkably larger than
those in the other genera. In Metrodorea the buds are naked, due to their unique
position within the vaginae.
The phyllotaxy is of major importance. The leaves are opposite in Metrodorea,
Raulinoa, and Esenbeckia subgen. Oppositifolia, while the other subgenera of
Esenbeckia, and Pilocarpus, have essentially scattered leaves which may be al-
ternate, or ternate to whorled towards the tip. In Pilocarpus the latter condition
is often shown by simple-leaved species.
The division of the leaves, if any, is easy to observe. There are, however,
species which show a gradual transition from plurifoliolate leaves through uni-
foliolate to simple ones: Esenbeckia pumila, E. densiflora, and Pilocarpus ra-
cemosus. Species with simple leaves are unknown in Metrodorea, while pinnately
divided leaves are known only from Pilocarpus. Pilocarpus racemosus is espe-
cially remarkable in that some specimens have simple or unifoliolate leaves,
whereas others have l-3-foliolate or 1-3-jugate leaves. However, I have evidence
that simple leaves occur especially towards the tips.
The petiole may be narrowly winged or not. The presence of wings appears to
be a useful key character. Species of Esenbeckia with simple leaves are devoid
of wings. Species of Pilocarpus with simple leaves have the base of the blade
decurrent along the petiole which seems to be winged. Occasionally the wings
can hardly be distinguished from ribs that frequently occur on the petioles. There-
fore some species are inserted more than once in the keys. The petioles of some
Morphology 5

species of Esenbeckia with divided leaves show an adaxial, distal structure.Al-

though this is mostly a densely hairy tubercle, it attains a relatively large size in
some species, reachingbeyond the articulationof the leaflet. In such cases it calls
to mind the hastula of palms.
The feature which distinguishes Metrodorea from all other Rutaceae is the
presence of a remarkablydeveloped sheath on the leaves (see Kaastra, 1978a).
The two sheaths of each pair of leaves stick to each other until the developing
terminalbud, which is enclosed by them, forces them to separate. After sepa-
ration the sheaths are still connected with the branchletsat their base. For prac-
tical reasons in the technical descriptions which follow the length of the petiole
refers to the free part only.
Van Hall (1860) reported on the articulationof the terminal leaflet, and de-
scribed the terminalpetiolule as being much longer than the lateralones. Lemaire
(1852) reported this petiolule as being distinctly shorter. Van Hall imputed this
to a differentconcept of the term "petiolule." He named the stalk of the terminal
leaflet a petiolule, irrespective of its having one articulationat the base or, as in
his case, being doubly articulateby a second joint near the tip. It is probablethat
Lemairehad the same idea about this, but by accident studied only a small plant
with short terminalstalks which were once-articulate.In the present study I found
terminalstalks to 2 cm long in Pilocarpus pennatifolius, which were not articu-
late. The terminalstalks, therefore, seem to vary in length and in the numberof
articulations. I do not agree with van Hall's terminology since in my opinion
every stalk proximalto an articulationshould be described as part of the rachis
(or petiole), while the distal part merits the name petiolule.
The shape of the leaves is occasionally of some importance. The narrowly
obovate, cuneate leaves of Pilocarpusgiganteus are remarkableoften resembling
those of some species of Angostura. Pilocarpus jaborandi and P. trachylophus
can be distinguishedvegetatively from P. pennatifolius by the base of the leaflets.
This charactershould have been used much more in the past to identify leaves
imported as drugs (see ANATOMY). The base of the leaf(lets) in Esenbeckia
leiocarpa and E. runyonii provides a complementarycharacterfor easy identifi-
cation. The general shape of the base is attenuate in these species, but the very
base terminatesabruptlyon the stalk. In species with attenuateleaflet bases the
limit between the base and the petiolule is often arbitrary.The bases of lateral
leaflets are usually unequal. In most species the marginof the leaves is somewhat
recurved. The apex of the leaves and leaflets in Pilocarpus is nearly always
emarginate. The texture of the leaves is mostly subcoriaceous. Specimens of
Esenbeckia grandiflora with truly coriaceous leaves were collected along the
eastern Braziliancoast; the leaves are thicker possibly due to a differenthabitat
(see the descriptionfor more details).
The venation is described accordingto the system of Hickey (1973).Raulinoa
is somewhat different from other genera in the lateral nerves of the brochidod-
romous system since they join at an obtuse angle to make an intramarginalvein.
The numberof lateralnerves is not easy to determine,due to the gradualdecrease
in thickness of the nerves towardsthe apex. The angle of divergenceof the lateral
nerves hardlydiffersbetween the species, varyingfrom45-80?(measuredmidway
on matureleaves).

Inflorescences
The inflorescenceis of primaryinterest for taxonomy. It is paniculatein Esen-
beckia and Metrodorea, althoughboth true panicles and thyrses are found, the
latter in species with secondary pedicels borne in the axil of bractlets as in E.
6 Flora Neotropica

scrotiformis. Some species of Esenbeckia have a leafy panicle, e.g. E. flava, E.

leiocarpa, and E. oligantha. Raulinoa has raceme-like subfasciate inflorescences.
Pilocarpus has racemes only. These racemes have a long-persistentjuvenile stage
which is spicate; for illustration see Troll (1969: fig. 430). In P. pennatifolius
some specimens were observed in which the pedicels branch in the axil of a
bractlet. This may be an atavism, but shifting and fasciation is also possible.
Esenbeckia pilocarpoides and E. warscewiczii have a narrowly paniculate inflo-
rescence which is more or less racemiform.The inflorescences of Esenbeckia
subgen. Lateriflorensand of Raulinoa are much reduced in size, and are borne
axillary in large numberslaterallyalong the branchlets. There are 1-2 (to rarely
4) bractlets on each pedicel. Three or four bractlets sometimes occur in connec-
tion with a partialdoubling of the floral organs (see P. pennatifolius). This may
be an example of fasciation.

Flowers
The flower buds of subtribePilocarpinaeare globose, while those of subtribe
Cuspariinaeare oblong. This characteris connected with the shape of the anthers,
heart-shapedin the Pilocarpinaeand linear in the Cuspariinae.
The flowers are essentially pentamerous,except in Raulinoa and in Pilocarpus
spicatus (mainly in var. lealii), where they are tetramerous.This charactercan
be used provisionally for distinguishingbetween P. spicatus var. spicatus and
var. lealii.
The perianthsegments often continueto grow somewhatafteranthesis;notably
the calyx lobes enlarge. The petals of Metrodorea become thinner in age. All
parts of the flower may be adnate to each other at the base, althoughonly to a
slight degree. This is correlatedwith the continuousgrowth of the floralaxis (see
Gut, 1966).The calyx lobes of Metrodorea, however, are free from the petals.
In some perianthsegments, as occasionally in bracts and bractlets, the midvein
(or the central nerve of a parallel system) is embedded in yellowish glandular
tissue. This suggests that there should be a single, thick nerve only (pseudocosta
or false midrib).In petals a true single nerve is presentonly in Esenbeckia subgen.
Lateriflorens. [The presence of a single (branched)midrib(cladodromousvena-
tion) in petals is a characterwhich can be found otherwise in Balfourodendron
and Helietta, species of which often have been confused with Esenbeckia.
Anotherdistinguishingcharacterof floweringspecimens may be found in the base
of the petals: unguiculatein Helietta, unnarrowedin Esenbeckia. Specimens in
fruit or in postfloral stage are easily identifiedto genus, due to the presence of
wings in Balfourodendron and Helietta.]
The calyx has been variouslydescribedas toothed, lobed, or even as segmented
into free sepals. Frequently in Raulinoa there is hardly any coalescence, while
in other genera the segments are nearly always connate at the base, though to a
lesser extent than in subtribeCuspariinae.The calyx in Metrodoreavaries in the
degree of coalescence from one species to another; consequently it is described
as toothed or lobed.
The corollas are taxonomicallyimportant.The petals are always free from each
other, or nearly so, but in bud they more or less cohere at the tip. This is in
contrast to subtribe Cuspariinae.The aestivation is valvate in Metrodorea and
Esenbeckia subgen. Lateriflorens, and imbricate to valvate in Esenbeckia subgen.
Esenbeckia and subgen. Oppositifolia. Raulinoa has imbricate aestivation. The
petals of Pilocarpus are curious. Aestivation is essentially valvate, but the petals
in bud are always distinctly coherent by their uncinatelyinflexed tips. These tips
point towards the center of the bud and reach among the anthers. They make
Morphology 7

contact by means of a median keel on the upper side of the petals. This keel is
highly developed in P. trachylophuswhere it is provided with lateral wings. In
the petals of Pilocarpus there is an impression on each side of the keel corre-
spondingto the thecae of the two adjacentanthers.These impressionsare usually
permanent,although in a few species they disappearwhen the bud opens. The
apex of the petals is also inflexed in E. cornuta and E. oligantha, whereas other
species with valvate aestivation do not show this phenomenon, or hardly so.
The filamentsprovide a useful characterin Esenbeckia sect. Esenbeckia. Here
they are provided with a basal swelling, which is extremely large in E. alata.
These swellings must not be confused with the appendages of the filaments in
other Rutaceae and in Simaroubaceae.The latter, sometimes called "ligulae,"
are placed adaxially, and they are less swollen; the swellings in Esenbeckia, in
contrast, are abaxially arranged.
The filamentsare usually subulate, at least at their tip, but in some species of
Pilocarpus they are abruptlytruncate. Due to this, the anthers, which are ver-
satile in other species, are not able to switch, but instead recurve. All species of
Pilocarpus with simple leaves have such stamens. In species with divided leaves
the anthers are versatile, except in P. grandiflorus, while those of P. spicatus
are intermediate.
The anthers are dorsifixed, usually near the middle, never basifixed as some-
times suggested. Soon after sheddingthe pollen the anthers often fall off and the
filamentsbecome reflexed, or in Pilocarpus remain inflexed or become spread-
ing. Finally, the filaments in all genera are usually deciduous. Pilocarpus has
a dorsal gland near the anther tip, whereas the other genera have a protruding
connective.
In Pilocarpus the disc is completely adnate to the ovary (Gut, 1966).In other
genera it is partiallyadnate.
The carpels at anthesis are connate to variable degrees. They are completely
connate in Metrodorea and Raulinoa, connate at the base only in Pilocarpus,
and transitionalin Esenbeckia, being connate in some species (e.g. E. leiocarpa,
E. oligantha, and E. scrotiformis)and hardlyconnate or connate at the base only
in others (e.g. E. amazonica and E. feddemae). True apocarpy, as in Zantho-
xylum, does not occur in this subtribe.An interestingstudy concerningthis matter
was published by Gut (1966). The carpels are somewhat immersed into the re-
ceptacle, except in Pilocarpus, where they are often elevated. In the upperdorsal
part the carpels are provided (or become so) with an outgrowth, the apophysis
("Fruchtknotenkapuze"of Gut). The apophyses are always free from each other
and are absent in Pilocarpus. The style is always single, but often it is angledby
the five constituent strings which unite postgenitally.In Pilocarpus the stringsof
the style separatepostflorally,their remainsformingthe mucroat the apex of the
mericarps.The insertion of the style varies from gynobasic to anacrostylous,but
is never acrostylous. This is due to the dorsalgrowthof the carpels, as Gut (1966)
has demonstrated.
Although it is generally assumed that there are two ovules per carpel, it now
appears that species of Pilocarpus with simple leaves have but one ovule per
carpel. The ovules are always collateral, although if both develop, they grow
superposed.
Fruits
The fruits are usually quinquelocular,althoughsome species are tetramerous.
The parts of the fruits can be consideredfree in Pilocarpus ("mericarps"),where-
as they are connate in the other genera. Often in Pilocarpus one to four of the
8 Flora Neotropica

mericarpsdo not develop. This feature, which also occurs in Angostura (subtribe
Cuspariinae),could be an indication of separate paths for the pollen tubes. The
appendagesof the pericarpare often of taxonomic value, as is the general shape
of the fruits. The fruits of Pilocarpus may be confused with those of Angostura.
The fruits of the latter, however, often show a rim at the tip of each valve by
which they were attachedbefore dehiscence. Also, there are frequentlyremnants
of the perianthwhich can give an indicationabout generic identity.
The exocarp always has very prominentnervation. The nerves are subparallel
(reticulatein Esenbeckiaflava and E. hartmanii)and usually well observable on
the inside (but not in E. cowanii and hardly so in E. leiocarpa). The nerves are
often visible externally also, especially on the sides.
The endocarphas a remarkablestructure.In the Pilocarpinaeit is glabrousand
only rarely impressed by the nerves of the exocarp. Following Vahl (1797), the
endocarpis usually called "cartilaginous"and "elastically" dehiscent. The origin
of the exocarp and its anatomy were discussed by Hartl (1958). The innermost
layers are not lignified, however. From sections of Esenbeckia grandiflora it
appearsthat they are sclerified,while the outermostlayers are lignified.This was
shown by stainingwith phloroglucine-hydrochloric acid accordingto von Aufsesz
(1973). The outermost layer (borderingthe exocarp) is three cells thick in E.
grandiflora. The endocarp in E. hartmanii is distinctly thinner than in other Pi-
locarpinae.
The axial part of the endocarpis remarkablythinnerand probablynot lignified.
It is rupturedfrom the rest of the endocarpwhen the fruit dehisces. I join Hartl
(1958) in calling this structurethe "axial part." Wilson (1970) called it the "pla-
cental endocarp" but the use of the term placental is ambiguous in this case,
especially when compared with the idea of Holmes (1875: 583), who was of the
opinionthat the part concernedis a dilatationof the placenta.Anatomicalsections
may elucidate the origin. It is a pity that the area concerned is omitted in fig. 4
of Hartl's treatise.

Seeds
The seeds vary from ?4.5 mmin lengthin Esenbeckiahieronymito 19mmin Pi-
locarpus giganteus. The micropyle is sometimes surroundedby a caruncle of
verruculate, brown-black spots (E. grandiflora, E. macrantha, and E. pilocar-
poides). The seeds are keeled on the abaxial side except in Metrodorea and
Raulinoa. Their adaxial side is provided with a hilum runningdownwardsfrom
the micropyle. This scar is caused for the greater part by the accubation of an
obturatorduringthe development of the seed. The obturatorbecomes detached
before the seed is shedded (see Kaastra, 1978a).Near the base of the hilum the
vascularbundle merges into the chalazalarea. This area (the place on the outside
of the testa, above the true chalaza) is distinctly observablein several species of
Esenbeckia and Metrodorea, due to its elevation, its color, and its epidermal
structure, which is finer than in the rest of the testa. Wilson (1970) and Corner
(1976: 22) describe the chalazal area as "chalaza." This is incorrect, as the cha-
laza proper is not observable on the outside. The testa is coriaceous, up to 0.5
mm thick in E. berlandieri subsp. litoralis and in E. macrantha, but thinly co-
riaceous in Pilocarpus. The cells of the epidermishave their outer tangentialwall
dented or arched. They are arrangedin often characteristicreticulate or colli-
culate patterns. In Esenbeckia the interspaces are often linear and parallel in
patches.
On the inside of the chalazalarea there is the hypostase, a brown and relatively
Morphology 9

thick area of some millimeterslong. It is at this place that both integumentsare

inserted.
The embryosin all species are enclosed in a transparentmembranousenvelope.
This bag is attached to the testa by the hypostase but is otherwise free from the
seed. Its wall is composed of several layers of cells. This structure has been
called "endopleura"by Holmes (1875: 583) and "nucellar cuticle and remains"
by Corner (1976: fig. 491). The term "cuticular nucellus remains," however, is
better because there is no extensive cuticle. In the species of Esenbeckia subgen.
Oppositifolia this wall is further characterizedby the presence of granularen-
dosperm remains.
Most species of Pilocarpinaehave only one embryo per seed, but I have ob-
served up to eight in Esenbeckia grandiflora, E. oligantha, E. pilocarpoides,
Pilocarpus pauciflorus, and P. riedelianus. I do not know whether or not this is
due to nucellarpolyembryony, which frequentlyoccurs in the Rutaceae (see for
citations Desai, 1962, and Mauritzon, 1860).I observed pleiocotyly (Napp-Zinn,
1974:891) in E. leiocarpa and in P. pauciflorus. The embryos are all of the same
type having plano-convex, thick, eared-cotyledons,which are usually somewhat
unequal. The cotyledons lack starch, but have fat and aleuron, and secretory
cavities (Geiger, 1897:380). The cavities are often easily observable. The radicle
points towards the ears and is usually observable between or beyond them. In
E. hieronymithe radicle was seen to be reflexed outside the ears in the direction
of the chalaza. In all species of Pilocarpus with simple leaves the plumules and
radicles are strigillose, while the cotyledons are dirty greenish in P. giganteus
and P. pauciflorus. Hairy plumuleswere also observed by Corner(1976: 235) in
Glycosmis. Glandularhairs on the plumules were reportedfrom species of Me-
liaceae (Harms, 1940:6), otherwise hairy embryos seem to be rare accordingto
Napp-Zinn(1974: 889).

ANATOMY
Secretory Cavities and Related Structures
Secretory cavities are present in all species of Rutaceae sensu Engler. It is
often by these cavities alone that certain species can be recognized as belonging
to the Rutaceae and not, for instance, to the allied Simaroubaceae.The cavities
are usually referredto as (oil) glands or pellucid dots (also in this study), because
in foliar organsthey often appearon the outside as transparentpoints. There are,
however, several species in this family where, especially in the perianth, only
few glands are visible. Sometimes they are hardlyobservable, but in microscopic
sections they always appear to be present, e.g. in Esenbeckia cornuta, E. war-
scewiczii, and E. almawillia. Secretory cavities occur in most parts of the plant,
including the exocarp, but not in the endocarp and, according to van Tieghem
(1885: 58), not in the roots or in xylem and phloem. The cavities in the disc of
Pilocarpus species frequently unite to form a continuous ring. A survey of the
ideas and facts about the schizo-lysigenousdevelopmentof the cavities was given
by Schulze (1902: 59 ff.), but see also Gilg and Schiirhoff(1930), who reported
a lysigenous development (in Citrus and Ruta). Haberlandt(1898: 1238)has pro-
vided an outstandingdescription of the subepidermal,sunken secretory cavities
in Pilocarpuspennatifolius. These are covered by 3-7 (usually4) cover cells, and
easily release their contents when the leaflets are bent, but do not do so spon-
taneously. I have also observed these "sunken cavities" in P. demerarae, P.
grandiflorus, and in Metrodorea, but I did not look for them in other taxa. Ac-
10 Flora Neotropica

cording to Dede (1962) the situation of the secretory cavities is sometimes cor-
related with the venation pattern. I have some evidence that the situationof the
cavities depends also on the age of the leaves. This may account for the differ-
ences between type V of Dede (E. berlandierisubsp. acapulcensis, etc.) and type
VII (P. racemosus ("P. longipes"), Metrodoreaflavida).
The contents of the Rutaceous cavities are yellowish essential oils (see Heg-
nauer, 1973: 177ff.).
Solereder (1899: 205) observed secretory cells in branchletsof Pilocarpus spi-
catus ("P. subcoriaceus"). They are present in the parenchyma of cortex,
phloem, rays, and (sometimes) pith. The cells in the phloem parenchymaare
extended in an axillary directionwith several cells above each other.

Trichomes
Although hair-cover appears to be of practical value in identification,its im-
portance should not be overemphasized. Taxa have previously been described
which differsolely in the density of theirindument.However, the more specimens
that are examined, the more intermediateplants appearto exist; therefore, such
taxa should not be maintainedas distinct, e.g. Esenbeckiapumila vs. E. leuco-
phylla, Pilocarpus parviflorus vs. P. spicatus, and the hairy and smooth "vari-
eties" of Holmes (1875) in Pilocarpus jaborandi.
There are two types of trichomes,viz. secretory hairs and those of the clothing
type. Clothing hairs are simple and usually unicellular.The hairs on the leaves
of Esenbeckia flava are densely pustulate externally. I observed multicellular
hairs on the inner side of the sheaths in Metrodorea. The multicellularhairs
mentionedby Duval (1905: 119)in Pilocarpuspennatifolius are in fact unicellular
(cf. Solereder, 1908:64). I did not observe stellate hairs, althoughWardleworth
(1893b) reported their scanty presence in P. microphyllus.I agree with Geiger
(1897: 387) in the assumption that Wardleworthprobably described glandular
hairs with resin-likeexudate. The hair-coveron the upperside of the petals varies.
In some species of Esenbeckia there are minute papillae observable when mag-
nified x80; these petals are called "glabrousabove" in the systematicpartof this
paper. In the other species of Esenbeckia the papillaeare observablewith a hand-
lens, or grow out into unicellularhairs reachinga length of 1 mm in E. scrotifor-
mis. The trichomes in the latter species have longitudinalthickenings.Raulinoa
has papillose petals, Metrodorea has hairy ones, and Pilocarpus has them gla-
brous above. Remarkableis the occurrence of hairs on the embryo in some
species of Pilocarpus (see MORPHOLOGY).
Secretory hairs are externalglands with usually globular,multicellular,stalked
heads. Geiger (1897:381) gave a detailed account of the glandularhairs (as of the
clothing hairs). They are reportedto be present in all species of Pilocarpus, but
are very sparse in P. microphyllusandP. spicatus, and they can only be observed
microscopically. I could find them in P. riedelianus only with difficulty;in P.
pennatifolius the glandularhairs are almost completely immersed into the epi-
dermis, in P. jaborandi only partly so. In P. microphyllus, P. trachylophus, and
P. spicatus they are not sunken and therefore are often broken off. Geiger ob-
served clavate glandularhairs in P. trachylophus.The structureof the glandular
hairs can best be studied when they are young, because in age they produce a
resin-like exudate which often deforms their shape and structure. Duval is re-
ported to have seen glandularhairs in Esenbeckiafebrifuga (Solereder, 1908),
but I did not see them there, nor in any other species of Esenbeckia except both
species of subgen. Lateriflorens(see Fig. 2).
Anatomy 11

FIG 2.
FIG. 2. Cross-sectionof leaflet showinga sunken multicellularglandularhair (Ule 9493, K).

Roots
The microscopic structure of the roots of Pilocarpus was studied by Planchon
(1875: 296) and Duval (1905: 22, t. 2, fig. 11). The cortex parenchyma contains
starch and rosette-like crystals intermingled with numerous large secretory cav-
ities. The phloem fibers are single or grouped together, and intermingled with
stone cells. The rays are generally three cells wide and contain starch grains.
Some of the vessels contain yellow-green substances.

Branchlets
Knowledge of these parts is fragmentary. The initiation of the phellem in Pi-
locarpus is subepidermal, not epidermal as stated by Solereder (1899: 204).
The cortex of Esenbeckiafebrifuga was studied more extensively. The phellem
is 6-10 layers thick. Below it there is a phelloderm of 6-8 layers of colorless,
thin-walled cells. In the cortex there are tangentially arranged sclereids, inter-
rupted by rays of 1-2(-3) cells wide and 10-20 cells high. The parenchyma con-
tains starch, calcium oxalate, and cells with tannin-colored contents. There is a
specific color reaction for the phelloderm. The cells contain "evodine," renamed
esenbeckine, which gives a blue-green color when treated with K2Cr2O7and con-
centrated H2SO4 (Gamper, 1900).
The cortex of "Metrodorea pubescens Saint-Hilaire & Tulasne" (=M. stipu-
laris?) is reported to have numerous branched sclereids (Solereder, 1899: 205).
The cortex of Pilocarpus shows many cells with resin-like contents (Fliickiger
& Hanbury, 1878: 254), oxalate, and starch (Geiger, 1897: 379; Stiles, 1877).
While there are usually isolated bundles of phloem fibers in the rutaceous peri-
cycle, some species of Pilocarpus are different. Solereder (1899: 204) observed
in the pericycle of P. spicatus a mixed continuous sclerenchyma ring consisting
of primary phloem fibers intermingled with groups of sclereids. Rocher (1898)
observed this in P. racemosus and in the same species found medullar crystals.
Some Pilocarpinae can be distinguished by their branch anatomy. Pilocarpus
12 Flora Neotropica

jaborandi has characteristicradiallyelongated sclereids in its pericycle (see Gei-

ger, 1897); in P. pennatifolius the sclereids are tangentially stretched (Duval,
1905: 34). Pilocarpus spicatus shows abundantsimple calcium oxalate crystals
in the cortex (Geiger, 1897).Pilocarpus trachylophushas a red medulla(Geiger).
Good drawings of P. jaborandi can be found in Fliickiger and Hanbury (1878:
fig. 76), Geiger (1897:figs. 11-12), and Duval (1905:t. 2, fig. 12). Descriptionsof
the branchletsof P. jaborandi were given by Poehl (1880: 140), and Stiles (1877).

Secondary Wood
The first good general descriptionswere given by Record in Record and Hess
(1940), and are of easy access so that they do not need to be quoted here. They
include the genera Esenbeckia (based on seven species and, in addition, on He-
lietta plaeana = "E. atata"), Metrodorea (based on M. flavida and M. nigra),
and Pilocarpus (based on P. racemosus and P. pennatifolius). Heimsch (1942)
added that the rays are heterogeneous in Esenbeckia and Metrodorea (type IIB
of Kribs prevails), or homogeneous in Metrodorea, of type I or II of Kribs'
system of 1935. The rays in Esenbeckia (based on two species) should be up to
six or seven cells wide, whilst other authorsmentionup to three or four cells (see
also below). There are oxalate crystals in the medulla and Geiger (1897: 379)
observed composed starch grains in medullaand rays.
Milanez (1943) published a description with plate of E. leiocarpa which does
not essentially differ from the general description by Record and Hess (1940).
Furthermorehe gave a key to the genera. Other descriptions were publishedby
Kribs (1968: 142, fig. 302) based on E. febrifuga, E. leiocarpa, and Helietta
plaeana ("E. atata"), by Mainieriand Pereira (1965: 153 with plate) of E. leio-
carpa, and by Pereira(1933: 150 with plate) also of E. leiocarpa.
An unpublishedstudy of the wood of two species by Koek-Noormanand A.
M. W. Mennega(Utrecht) is given here to report recent progress.

Esenbeckia pilocarpoides Kunth, Fig. 3A-C.

Material: Cowan & Lindeman 39194, Surinam, Nassau Geb.; stem 3 cm in
diam.
General characters: wood light yellowish-brown, texture very fine, straight-
grained, hard, and heavy. Growth rings faintly visible with the naked eye. Mi-
croscopic characters:growth rings probablyindicatedby + regularand straight
marginalparenchymabands. Vessels (?80%) in short radial rows of 2-3, very
numerous, ?90(80-100) per sq. mm, evenly distributed;diam. very small, 25-40
,tm; perforationssimple, intervascularpits very fine, 3 /um, the borders vague,
the apertures included or coalescent; vessel element length 350(275-420) ,um;
often a granularyellow substance plugging the perforationplates. Fibers non-
septate, very irregularlyarrangedas seen in transverse section, angular, thick-
walled, diam. ?12 /am, lumen 3 /am; pits small, slit-like, restricted to the radial
walls; length 1200(875-1400)/,m. Ratio of fiber/vessel element length 3.43. Rays
uni- and biseriate, the uniseriate consisting of square and erect cells, 2-8 cells
high, the biseriate consisting of a middlepart of procumbentcells 6-11 cells high,
uniseriate extensions of square and short erect cells usually 2-3 cells, though
sometimes many more cells high; width of the biseriate rays 20 ,/m, height 200-
400 ,/m; pits to the vessels as the intervascularpits, crowded; rhombic crystals
in the procumbentand square cells occasionally present; number 9.5(7-11) per
mm. Parenchymaapotrachealin (probablymarginal)bands, the bands 2-5 cells
wide, nearly straight, occasionally anastomosing, 3-4 per mm, diffuse strands
Anatomy 13

scarce; paratrachealparenchymausually present as a few cells borderinga vessel,

but seldom forming a complete ring; strands of 2-4 cells, also fusiform cells
present; crystals rare; average length 340 um.

Pilocarpus demerarae Sandwith, Fig. 3D-E.

Material:Fanshawe 2065 (FD 4801), Guyana, Essequibo; a plank of a stem at
least 4.5 cm in diam.
Generalcharacters:wood light yellowish-brown,texture fine, straight-grained,
very hard, heavy, vol. weight 1.1. A faint yellowish stripingon tangentialsur-
face is due to (probably marginal)parenchymabands. Microscopic characters:
growth rings present, indicated by a narrow straight marginalband of paren-
chyma, the spacing of the bands irregular,at least in part of the circumference
of the stem, in some of the rings the vessels in the first formed wood more
numerous,in the majorityof rings the distributionof the vessels regular.Vessels
for the greater part (75-80%)in radialrows of 2-4(-6) or in clusters, numerous,
40(32-56) per sq. mm and very numerous, -+100in the closely spaced narrow
rings; diam. small, 40-60 ,m to very small, 20 ,um; perforationssimple, inter-
vascular pits very fine, 3 ,um, aperturesincluded;vessel element length 456(360-
580),um;a yellowish granularsubstanceoften present near the perforationplates.
Fibers non-septate, in rather irregulararrangementas seen on a transverse sur-
face, angularto rounded, diam. ?16 ,um, the walls 4 Aimand the lumen 8 ,um;
pits slit-like, restricted to the radial walls; length 1200(900-1400),m. Ratio of
fiber/vesselelement length 2.6. Rays uni- and 2-4-seriate, the uniseriateconsisting
of square and erect cells, 2-8 cells to 200 tim high, the multiseriateconsisting of
a middle part of procumbentcells, 2-32 (mostly 18-25) cells high, the uniseriate
extensions of square and erect cells usually 1-4 (sometimes more) rows high,
rarely vertically fused rays present; width of 2-4-seriate rays to 50 Am, height
mostly 350-650 (to 900) mm;pits to vessels as the intervascularpits, crowded;
no crystals noticed; number8 per mm. Parenchymapredominantlyapotracheal
as (probably)marginalbands, the bands straight, uninterrupted,2-4 cells wide,
mostly rather widely spaced, 1-2 per mm, elsewhere close together; solitary
strands or small patches of diffuse parenchyma virtually absent; paratracheal
parenchymapresent as complete or partialnarrowrings aroundpart of the ves-
sels; strands of 2-4 cells, on average 458 ,im long.
Petiole
The petiole was studied only in Pilocarpus. The petiole of P. pennatifolius was
studiedby Plitt (1886:39), Geiger(1897:388, fig. 1), and Schulze (1902:82). There
is a closed vascularcylinderat the distal end, which is triangularin cross-section.
The vascularbundle consists of a xylem-phloemring which is interruptedby rays
1-2 cells wide. A ring of sclerenchyma groups encircles the vascular bundle.
Some data about the petioles of P. jaborandi, P. microphyllus, P. trachylophus,
and P. spicatus can be found in Geiger (1897). The latter species may be char-
acterized by the presence of simple calcium oxalate crystals in cortex, phloem,
and medulla, as were also observed in P. racemosus by Rocher (1898: 186).
The abscission zone in the petiole above the insertion of the leaf is curious in
Pilocarpusjaborandi (Geiger, 1897:384). Three vascular strands enter the leaf,
each surroundedby groups of collenchyma cells. Distal to the place where the
strands unite into one bundle the collenchyma is replaced by a closed fiber ring
which can easily be broken. However, I did not actually see the resultingbreak-
ing-off in any herbariumspecimen.
14 Flora Neotropica

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Anatomy 15

Leaf Blade
When I sectioned Esenbeckiafebrifuga leaves to look for glandularhairs, large
oblong crystals appearedto be present in the pericycle. In the leaves of E. pumila
I observed transversecolumns of 3-5 cells thickness, consisting of sclerenchyma
and extendingfromjust below the upperepidermisdown to the lower epidermis.
All other confirmeddata are reportedfrom Pilocarpus. The results of some au-
thors are doubtful due to misidentifications,e.g. from P. jaborandi where the
plants were identifiedas P. pennatifoliusby Semenow (Geiger, 1897:385), Fliick-
iger and Hanbury(1878: 250), and Karsten (1903).
The leaves are bifacial with a strong cuticle on both sides (Geiger, 1897:377).
The cuticle is striate in P. jaborandi (Poehl, 1880: 137) and in P. racemosus
(Rocher, 1898: 183). The lower epidermis is papillose in P. giganteus (Duval,
1903: 50), P. pennatifolius, and P. trachylophus, but not in P. jaborandi (Geiger,
1897:388, 391). The numberof subsidiarycells is 4-5 accordingto most authors,
but 24 in P. jaborandi (Meyer, 1892), and up to 6 in P. trachylophus(Geiger,
1897:391). Some authorshave observed crystals in the epidermalcells of certain
species. These were supposed to be crystals of the flavonone glycoside hesperi-
dine. Geiger (1897) observed crystal aggregates in the epidermis of P. pennati-
folius and especially in that of P. trachylophus. He doubted the presence of
hesperidinebecause it had not been identifiedwith certainty. Accordingto Heg-
nauer (1973: 196)the crystals are probablya flavone glycoside, viz. diosmine.
A hypodermiswas observed in P. giganteus (2-3 layers, Duval, 1905:57), P.
goudotianus (1 layer, Duval, 1905: 53), P. jaborandi (Poehl, 1880: 138), P. ra-
cemosus ("P. latifolius") (1-2 layers, Duval, 1905: 52; Geiger, 1897: 396), and
P. spicatus (Geiger, 1897: 396). The hypodermis is especially developed at the
margin,accordingto Geiger (1897).
The radialwalls of the palisade cells are not closely joined (Geiger, 1897:377).
This is also shown by Meyer (1892:fig. 442) in P. pennatifolius, by Poehl (1880:
138)in P. jaborandi, and by Rocher (1898: 184)in P. racemosus. Some palisade
cells are septate and contain an oxalate druse in each compartment[see Duval
(1905:t. 4, fig. 11),Geiger (1897:377), and Rocher (1898: 184)].Cells with oxalate
crystals frequentlyobstruct the substomatalchambers(see Geiger, 1897:fig. 21,
and p. 377).
The midnerveis collateral, and encircled by a sclerenchymaring, the structure
of which can be used for identification(see Geiger, 1897:fig. 2).
Originaldescriptionsor drawingsof the leaf anatomyare found in the following:
P. jaborandi (Duval, 1903: 47, t. 2; Fliickiger & Hanbury, 1878: 253; Geiger,
1897: 380; Meyer, 1892: 228; Poehl, 1880: 135);P. microphyllus(Duval, 1903:
101; 1905:fig. 2; Geiger: 394); P. pennatifolius (Duval, 1903:50; 1905:t. 6, fig.
1; Geiger:386);P. racemosus (Duval, 1905:t. 6, fig. 3; Rocher, 1898: 183,fig. 4);
P. spicatus (Duval, 1903:102;1905:fig. 2; Geiger, 1897:395);andP. trachylophus
(Duval, 1903:98, t. 3, fig. 4-8; 1905:t. 6, fig. 5; Geiger (based on Vogl), 1897:
390). Duval (1903: 107)gives a key to the species of Pilocarpus.

FIG. 3. Secondary wood. A-C, Esenbeckia pilocarpoides (Cowan & Lindeman 39194, U). D-E,
Pilocarpus demerarae (Fanshawe 2065, type, U). A, x.s. x90. B, t.s. x90. C, r.s. x225. D, x.s.
x90. E, t.s. x90.
16 Flora Neotropica

Anthers
The tapetum of Pilocarpus pennatifolius is of the secretory type (Honsell,
1954).
Ovules
The ovules of Pilocarpus pennatifolius have been studied by Mauritzon (1935)
and by Honsell (1954). They are crassinucellatewith two integuments. Both in-
teguments form the micropyle, but according to Mauritzon(p. 330 and fig. 4N)
they set free the upper part of the nucellus duringfertilization. The nucellus is
provided with a cap when mature;the cells of the nucellus cap seem to aggregate
into clusters (Mauritzon:fig. 41). There is sometimes more than one embryo-sac
mother cell but only one of them matures(Mauritzon:323, fig. 2A-B).
Fruit Wall
The exocarp was studiedby Geiger(1897:379, 393). It is parenchymatous,with
the outermostlayers sclerified, especially on the adaxial side. The nerves on the
inner side are partly surroundedby fibers, and partly by sclerified parenchyma
cells. Pilocarpus trachylophusdiffersin that its nerves send small side-branchlets
into the tubercles on the outside. A drawingof the fruit wall of P. pennatifolius
was given by Corner(1976: fig. 490).
The endocarp was discussed in MORPHOLOGY.
Seed Coat
The seed coat seems to be of systematic importance. According to Geiger
(1897: 380), Duval (1905: t. 7), and Corner (1976: 235, fig. 491) there are the
following tissues in seeds of Pilocarpus (from the outside inwards). The testa
consists of a distinct dark brown cuticle with several layers; a simple epidermis,
the cells of which are extended tangentiallyin P. spicatus (Geiger);several layers
of loosely connected parenchymawith thickenedpitted walls, the outermostlay-
ers of which are suberized;and a layer of small, colored cells in P. pennatifolius
only, accordingto Geiger (not seen by me). The tegmen consists of one or more
layers of cells with spiral or annularthickenings;a layer of thin-walledlarge cells
(with spiral thickenings in P. jaborandi, according to Geiger); and a layer of
small, crushedcells which connects the tegmen with the nucellus (Corner).Draw-
ings of the seed coat of the following species can be found: P. jaborandi (Duval:
t. 7, fig. 1; Geiger: fig. 15); P. giganteus (Duval: t. 7, fig. 5); P. microphyllus
(Duval: t. 7, fig. 6; Geiger: fig. 30); P. pennatifolius (Corner:fig. 491; Duval: t.
7, fig. 2; Geiger:figs. 7-8); P. racemosus (Duval: t. 7, fig. 4); P. spicatus (Duval:
t. 7, fig. 3; Geiger: fig. 27); P. trachylophus(Duval: t. 7, fig. 7; Geiger:fig. 22).
I have made sections of Esenbeckia grandiflora seeds, and the structure is
generally similar to that in Pilocarpus. The cells of the outer epidermis show a
fine striation in their papillose cuticle. There are 9-11 layers of thick-walled,
suberized cells below them, while the inner 3 layers consist of very thick-walled
parenchyma,the innermostlayer of which is not suberized. The tegmen consists
of tracheidswith annularor spiralwall thickenings,with a parenchymatous,thin-
walled layer on the inner side.
POLLEN AND CHROMOSOMES
Pollen
There is no published record concerning the morphology of the pollen. The
grains are large, but I have no statistically reliable data. According to Honsell
(1954) the pollen of Pilocarpus pennatifolius is binucleate.
Pollen and Chromosomes 17

Chromosomes
Some counts of the diploid numberwere made from root apices in Pilocarpus
pennatifolius: 2n=36 (Honsell, 1954); 2n =44 (Kaastra, 1978b); Esenbeckia febri-
fuga: 2n=64 (Kaastra, 1978b).The only chromosomenumbersknown from sub-
tribe Cuspariinaeare accordingto Moore (1973):2n = 36 in Ravenia spectabilis,
and 2n=89-90 in Erythrochitonbrasiliensis. In the Boronieae related chromo-
some numbers are known (Darlington& Wylie, 1961):2n=22 in species of Bo-
ronia; 2n =32 in species of Boronia, Phebalium, and Correa; 2n =64 in species of
Phebalium. With so few counts it is hard to draw conclusions.

PHYTOCHEMISTRYAND PHYSIOLOGICALACTION
General
Although the Rutaceae have been frequently studied phytochemically, espe-
cially duringthe last 25 years (cf. Fish & Waterman,1973;Hegnauer, 1973),the
Pilocarpinae, except P. microphyllus and possibly P. jaborandi (?) (Loewe &
Pook, 1973;Tedeschi et al., 1973, 1974)have largely been ignored. There was an
enormousinterest in Pilocarpus from 1874until ca. 1900where I have seen more
than 55 articles, 30 of which appearedwithin the first four months of 1875. The
introductionof the drugs into Europe caused a stir in the medical and pharma-
ceutical world, due to the physiological action of Pilocarpus, therefore I give a
survey of the therapeuticaluses at the end of this chapter. A summaryof the
isolates from the Pilocarpinaeis given below and is restricted mainly to the al-
kaloids. No data are availablefrom Metrodoreaand Raulinoa.

Esenbeckia
Vitaglio and Comin (1970a)reportedthe isolation of rutaevinfrom the bark of
E. febrifuga. Later (Vitaglio & Comin, 1970b),they reportedtheir isolates, flin-
dersiamine, maculine, and skimmianine (all from E. febrifuga). Oberlin and
Schlagdenhauffen(1878)and Gamper(1900)extracteda compoundfrom the bark
of E. febrifuga which is probablylimonin, but which was called "evodin" by the
former and "esenbeckin" by the latter. In 1829 Buchner had already isolated
:'esenbeckine" from bark of E. febrifuga but it is not clear whether this com-
pound is identical with limonin (cf. Peckolt, 1899:337).
Dreyer et al. (1972)reportedthe isolation of the following alkaloidsfrom stems
and branchesof E. hartmanii:limosin, maculosidine,rutaevin, skimmianine,and
probablyalso limonin diosphenol.
The wood of E. leiocarpa was studied by Freise (1936). The presence of tannic
acid was demonstrated, as well as that of the rare galloyl tannin, of catechin
tannin, and of saponine.

Pilocarpus
The list of data concerningPilocarpus is too large to give more than a fraction
here. Several results of chemical purificationand analysis in the Pilocarpinae
should be criticallylooked at again because of misidentificationsof the extracted
plants. This can be shown for what has been called "jaborandi." Plants of
several families have been treatedunder this name, mainlyPiper, Herpestis, and
Pilocarpus since all are known to have more or less the same physiologicalprop-
erties (for a review see Geiger, 1897).Pilocarpus alone yields materialin some
quantityfor export, and will be discussed here. It came to Europe at the end of
18 Flora Neotropica

1873,broughtby Coutinho(Gubler, 1875),who obtainedit from Pernambucoand

sent it to Gubler in Paris. The latter confirmedthe therapeuticalaction which
Coutinhoused in his Brazilianmedicalpractice. He gave a leaf fragmentto Baillon
for identification. Baillon misidentifiedthis scanty material as belonging to P.
pennatifolius;he (correctly)identifiedmaterialfrom a later importas P. selloanus
(=P. pennatifolius) (Baillon, 1875, 1878; Gubler, 1875). Pilocarpus pennatifolius
was indeed used in Paraguay and sampled for export to Europe (see Baillon,
1878),but the first lot, sent by Coutinho, was reportedto have been collected in
northernBrazil, where P. pennatifolius does not occur. Moreover, Martindale,
Ringer, and others doubted the correctness of the identificationby Baillon on
account of the medical properties, when comparedwith later importsfrom Par-
aguay (Poehl, 1880). Holmes, early in 1875, obtained pieces of the plant from a
parcel sent from Pernambucoto England.He describedand figureda plant which
about twenty years later was validly published as P. jaborandi. Holmes has
argued several times that the identificationby Baillon was false (it should have
been the same plant as Holmes had studied), but due to the influence of Baillon
the idea, that the firstjaborandi importedinto Europe should have been P. pen-
natifolius, persisted. This fiction can be found in several papers and books, also
in most pharmacopoeias.
Fliickiger and Hanbury (1878) were of the opinion that the identificationby
Baillon was right and that by Holmes wrong, but their descriptionwith drawing
obviously refers to P. jaborandi!
Planchon (1875) obtained material which had arrived in Paris in early April
1875. His descriptionimplies that there were fruits and infructescencesamong it
which belong to P. giganteus (P. macrocarpus). The identificationof the fruits
as P. heterophyllus (=P. racemosus) is surely incorrect. Whether the other ma-
terial representedP. pennatifolius or P. jaborandi can not be concluded. How-
ever, because of the admixtureof P. giganteus it is likely that it was P. penna-
tifolius, because these occur in the same area whereas P. jaborandi grows 1700
km to the north.
Bentley and Trimen(1878) were of the opinion thatjaborandiwas affordedby
both P. selloanus and P. pennatifolius. They based this idea however on a spec-
imen from the hortus at Lisboa (Goeze s.n.) which is indeed P. pennatifolius.
The fruit which they described from a specimen of Holmes belongs to P. jabor-
andi.
In 1884Baillon described and figuredajaborandi which is P. pennatifolius. At
this time he conceded that the jaborandiof Holmes originatedfrom a different
species.
The reports of large differences in alkaloid contents (see Dragendorff, 1898:
353; Wehmer, 1929)can be explained by several factors includingimperfectiso-
lation techniques, differences due to the age and time of collecting of the plants,
the misidentificationsmentioned above (see also under P. microphyllus,in this
chapter!), and particularlythe usual mixing of the materialin commerce. Fur-
thermore,the alkaloidpercentagein commercialmaterialdecreases with increas-
ing moisture, and is lower in plants cultivatedin Europe (Duval, 1903: 109;Heg-
nauer, 1973:226).
Chemical compounds are extracted from all aerial parts of the plants, but es-
pecially from the leaves. The main alkaloids isolated from Pilocarpus are all
imidazole alkaloids of the type of pilocarpine.In additionto pilocarpinethe fol-
lowing allied compoundshave been isolated from Pilocarpus: isopilocarpine,pi-
losine, isopilosine, and epiisopilosine. Reports of the presence of the following
compoundsin Pilocarpus is erroneous, as they are artifacts:jaborine,jaboridine,
Phytochemistryand PhysiologicalAction 19

pilocarpidine,and isopilocarpidine.For a survey of the history of chemical iso-

lations from Pilocarpus see Rocher (1898).
Pilocarpus giganteus ("P. macrocarpus") is mentioned by Dragendorff (1898)
and could possibly have small amounts of pilocarpine.
Pilocarpusjaborandi (Pernambucojaborandi),as stated above, has frequently
been confused with P. pennatifolius. The alkaloid content is much higher, how-
ever. Apart from earlier isolations, this species has recently been studied by
Tedeschi et al. (1973), who report the isolation of pilosine, isopilosine, and epi-
isopilosine from its leaves (but see below, under P. microphyllus).This species
has probablyrarely been involved in shipments(cf. Holmes, 1894).
Pilocarpus microphyllus(Maranhamjaborandi)was imported to Liverpool in
1893. It was called Guadeloupejaborandiby Wehmer(1921: 616), a name origi-
nally given to P. racemosus (cf. Holmes, 1904;Rocher, 1898;Duval, 1905:46).
Even fragments of P. microphyllusare easy to identify, so that the source of
isolations is probably reliable. Apart from pilocarpineand isopilocarpine"pilo-
sine" has been isolated by Pyman (see Voigtlander& Rosenberg, 1959:579) and
by Leger and Roques (1912), the latter calling it carpiline.This compoundshould
be renamedisopilosine (Voigtlinder& Rosenberg, 1959:579). In 1959Voigtlander
and Rosenbergreportedthe isolation of 0.06-0.1% of alkaloidfrom 50 kg leaves,
from which they isolated "true" pilosine. The physiological action of pilosine
and isopilosine is very low according to Voigtlinder and Rosenberg. Recently
pure pilosine, isopilosine, and epiisopilosinewere isolated (Loewe & Pook, 1973;
Tedeschi et al., 1973, 1974). The botanical source of the extractions reportedin
the last two papers, however, is obscure, as "P. macrophyllus," P. microphyllus,
and P. jaborandi are alternatelyused in those publications. Tedeschi said (pers.
comm.) that the crude pilosine was extracted from P. microphyllusand the epi-
isopilosine from P. jaborandi. According to Wehmer (1929: 615) and Karsten
(1962: 248) the Folia Jaborandishould refer to this species only, but the com-
mercial material was often mixed with other species. The Pharmacopoeia hel-
vetica (correctly) refers the Folia Jaborandito P. jaborandi, however.
The firstmentionof the export of Pilocarpuspennatifolius(Paraguayjaborandi)
to Europe was by Balansa. He, as well as the Paraguayanbotanist Parodi have
seen the medical utilization of this species in Paraguay(Baillon, 1878). Wehmer
(1929) mentioned some isolations, but there is reason to doubt them, due to the
misidentificationsmentioned earlier. It seems to be (or has been) cultivated in
Sicily, where isolations showed a very low percentage of alkaloids (Hegnauer,
1973).This species is usually referredto as Folia Jaborandi,but incorrectly. The
pharmaceuticalname was based by Holmes on P. jaborandi.
Rocher (1898) isolated 0.6% pilocarpinefrom Pilocarpus racemosus (Guade-
loupe jaborandi). I do not doubt his identificationbecause he had a reference
collection from Geiger and the descriptionand photos also agree. Steineggerand
Hansel (1968)mentionthe isolation of 0.25%of alkaloids, one-thirdof which was
pilocarpine (from "P. heterophyllus").
Pilocarpus spicatus subsp. aracatensis (Aracati jaborandi) was imported into
Englandin 1895(Holmes, 1895a).Bearingin mindthe originof Aracatijaborandi,
and the commercialleaves preserved by Holmes, this drug must come from this
subspecies. The only isolations of alkaloidsknown are those by Pauland Cownley
(1896) which contained0.16%amorphousalkaloids, and by Petit and Polonowski
(see Wehmer, 1929)which contained 0.3% of compounds. Neither isolation was
pilocarpineaccordingto Wehmer(1929). The identity of the plants used by Paul
and Cownley was probably correct as it was based on material provided by
Holmes; I do not know the identity of the plants used by Petit and Polonowski.
20 Flora Neotropica

Pilocarpus trachylophus (Cearhjaborandi) appeared in Europe in February

1894 (Holmes, 1894). Paul and Cownley (1896) isolated only 0.02% crystalline
nitrate. Petit obtained 0.1% crystallinenitrate (Holmes, 1895b).

Physiological Action and Medical Utilization

Esenbeckiafebrifuga (Cortex Esenbeckiaefebrifugae)was used as a substitute
for CortexAngusturaeand CortexCascarillae.Martius(1843:39) saw its excellent
usefulness against dyspepsia, as a stomachic, and especially against fevers; he,
together with Spix, used it many times (Peckolt, 1899: 337). According to Dra-
gendorff (1898) it was also used as an anticatarrhum.
Esenbeckia grandiflora (variety . .?) ("E. intermedia" of Martius, 1843). The
bark was used like that of E. febrifuga, but its effect was much weaker (Martius,
1843).
Esenbeckia pumila should also have been used as E. febrifuga, according to
Dragendorff(1898) and Peckolt (1899: 338).
The bark of Metrodorea stipularis ("M. pubescens" of local usage = M. pu-
bescens auct non Saint-Hilaire& Tulasne) was used like that of Esenbeckia
febrifuga, and the leaves were used as a sudorific(Peckolt, 1899:338).
The physiological action of some species of Pilocarpus seems to be based
mainly on pilocarpine, its stereoisomers being less active. Pilocarpine stim-
ulates the parasympatheticnervous system. The smooth muscles, therefore,con-
tract, resulting in the following effects: diarrhea, secretion of exocrine glands,
stricture of the pupil, spasm of the accommodationmuscle (myopia), thwarting
of respirationby contractionof bronchialmuscles, vasoconstrictionwith increase
in circulation, decrease of eyeball pressure by dischargingchamberhumor, and
heart weakness (cf. Martindale,1875, who took too much of it, with consequent
vomiting).Because of its side-effect on the heart medicalapplicationis restricted.
It is used in ophthalmology,mainlyagainstglaucoma.Accordingto Pinkhof(1963)
it is also used for early recognitionof lepra. It can be used as an antidoteagainst
atropine. Pilocarpus pennatifolius was originallyused in Paraguay as a masti-
cator, odontalgic, and sialagogue(Baillon, 1878).
Under each species in the systematic part are mentioned uses recorded on
herbariumlabels.

RELATIONSHIPSAND PHYLOGENY
The Englerian system of classification for the Rutaceae is widely accepted
today, albeit with some transferencesby recent authors. Objections to it have
been raised not by taxonomists alone but also by phytochemists (Fish & Water-
man, 1973)and by studies of floral anatomy (Moore, 1936);but see, in contrast,
the morphologicalstudy by Gut (1966). Engler alone, however, has studied the
family as a whole, even if more or less superficially,and subsequentadditionsto
our knowledge of the family are of a fragmentarynature. Therefore, no one is in
a position to make fundamentalwell-foundedmodificationsof the entire family,
but ratheronly minor changes within his "own" taxon. The systematic position
of the Pilocarpinaetherefore,mustbe left untouched,at least until the Cusparieae
and some other tribes of the family have been revised.
The presence of similar species in the Pilocarpinae and in the Cuspariinae
(Pilocarpus vs. Angostura) may be an expression of close relationship.The sim-
ilar appearanceof Esenbeckia and Helietta is also puzzling. These genera belong
to differentsubfamiliesbut their species can only be referredto the correct genus
when fruiting specimens are known. Formerly, many species originally were
Relationshipsand Phylogeny 21

placed in the wrong genus due to the lack of fruits. As mentioned in MOR-
PHOLOGY, the Pilocarpinaediffer mainly from the Cuspariinaein the plano-
convex, not folded, cotyledons, the different shape of anthers and flower buds,
and the degree of coherence of flower parts. Moreover, most Cuspariinaehave
a distinct silification of the leaves, which is absent from the Pilocarpinaeas far
as known (Edman, 1936).
Metrodorea was considered by Hooker (1862) as congeneric with Esenbeckia.
The two genera are easy to distinguishmerely by the sheaths in Metrodorea, but
this characteris consistently accompaniedby opposite leaves and inflorescence
branchlets, and by the valvate aestivation, unguiculatebase, and cladodromous
venation of the petals. AlthoughI do not intend to unite these genera, they must
be regardedas closely related.
Raulinoa differs from the other genera by its spines, the intramarginalleaf
veins, and the slightly zygomorphicflowers. Whether it is more closely related
to Metrodorea as Cowan (1960) says, or to Esenbeckia (subgen. Oppositifolia or
subgen. Lateriflorens)cannot be concluded at present.
Pilocarpus is distinctly different from the other genera in its inflorescences,
occurrence of pinnate leaves, complex petals, non-versatile anthers in some
species, presence of anther glands, the adnate disc, a lesser degree of syncarpy,
the reduction of ovule number, and hairy embryos in some species. This is sup-
ported chemically by the presence of imidazoles, which elsewhere in the higher
plants are only known from Casimiroa (Price, 1963). All these features could
justify the separationof all generabut Pilocarpus from the Pilocarpinae,the many
charactersin common notwithstanding.On morphologicalgroundsone could also
divide Pilocarpus into two subgenera because the simple-leaved species have
non-versatile anthers which are rare elsewhere, a single ovule, and strigillose
embryos. Withouthaving studied the Cuspariinae,however, I consider a subdi-
vision of Pilocarpus prematureat this moment, in contrast with the situation in
Esenbeckia, where earlier workers have already divided up the genus.
Based on the available data (Table I), the subdivision of Esenbeckia was re-
vised. The most remarkablefeature of Esenbeckia sect. Esenbeckia is the swell-
ing of the basal part of the filaments. This does not occur elsewhere in the Ru-
taceae.
The Cusparieaeare considered an advanced tribe, probably derived from an
ancestor in the Zanthoxyleae. This idea was developed by Engler (1931), due to
the reduction of endosperm, folding of the cotyledons, and coherence of floral
parts (see also Gut, 1966:236). Withinthe Cusparieaethe Cuspariinaeare more
advanced, as regards the degree of sympetaly and zygomorphy of the flowers,
but not as regardsthe numberof stamens.
I consider Esenbeckia the most primitivegenus in the subtribe,because of the
paniculateinflorescences, and the various degrees of syncarpy, usually less than
in Metrodorea and Raulinoa, and the lack of specialized chemical compounds
and such morphologicalstructuresas sheaths.
Pilocarpus, in contrast, is highly specializedby the occurrenceof the imidazole
pilocarpine.This is supportedby morphologicalcharactersincludingthe presence
of racemes (reducedpanicles, cf. Stebbins, 1974:262), and the reductionof ovule
number.Simple leaves and the low degree of syncarpyare considered secondary
reversions. It is remarkablethat Jussieu (1825: 402) already was of the opinion
that genera in the Diosmeae, among them Pilocarpus, with a single ovule per
carpel were stationed at the extreme limits of the group. Although he was not
aware of the single ovule in Pilocarpus, it appears that this genus is situated at
the extreme end of the Pilocarpinae.
22 Flora Neotropica

The occurrence of species pairs in the Pilocarpinaedeserves attention. The

best examples are Pilocarpus racemosus-P. goudotianus, Esenbeckia berlandi-
eri-E. pentaphylla, and E. pilocarpoides-E. amazonica. The members of each
pair are geographicallyisolated; a possible ecological isolation could not be con-
cluded from the data available. I have long pondered the status of the pair P.
racemosus-P. goudotianus and have concluded that the differentiationis recent.

DISTRIBUTION
The Pilocarpinae,like the whole tribe, is Neotropical with Esenbeckiarunyonii
alone extending into the United States, just over its southeasternfrontier. It is
notable that while Esenbeckia and Pilocarpus range throughoutthe Neotropics,
they are nearly completely absent from the Amazon basin. The only species
occurringin the upstreampart of the basin is E. amazonica. There is only one
species, Metrodoreaflavida, with an Amazoniandistribution,and this occurs on
non-floodedgrounds.
Esenbeckia has two main distributioncenters, one in Mexico and the other in
SE Brazil, with eight species in each. Between these areas there is a gap which
is occupied by Esenbeckia sect. Esenbeckia (Fig. 4A). Apart from Trinidad&
Tobago, Esenbeckia is very rarein the West Indies. There is only E. pentaphylla
subsp. pentaphylla which is confinedto Jamaica,and one questionablerecord of
E. pilocarpoides from Martinique.
Pilocarpus, in contrast to Esenbeckia, occurs throughoutthe West Indies, but
with one species only. For the rest, its distributionis about the same as that of
Esenbeckia but with most species occurringin SE Brazil (Fig. 4B).
Metrodoreais largely a Braziliangenus with only two localities known outside
Brazil (Fig. 35B).
Raulinoa is endemic to Santa Catarinain Brazil (Cowan & Smith, 1973).
Most species of the Pilocarpinaehave a limiteddistributionsome of them being
known to me by only one or two collections. On the other hand, E. grandiflora
ranges widely on both sides of the Amazon gap, and E. pilocarpoides has a rather
wide distributionwith its two subspecies.

SYSTEMATICTREATMENT
PilocarpinaeEngler(subtribein Rutaceae:Rutoideae:Cusparieae)in Martius,Fl.
bras. 12(2): 129-130. 1 Sep 1874(as "subtrPilocarpeae");Abh. Naturf.
Ges. Halle 13: 146. "1874" (between 5 Jul 1874 and 4 Jul 1875, as
"Subtr Pilocarpeae"); Engler in Engler and Prantl, Nat. Pflanzenf.
3(4): 157. 1896; Dalla-Torreand Harms, Gen siphon. 254. 1902; En-
gler in Engler and Prantl, Nat. Pflanzenf.(ed. 2) 19a: 278. 1931.
Pilocarpeae Bartling, Ord. nat. pl. 388. 1830 (as "genus Pilocarpea"), nom. invalid. ex Art. 33,
pro parte; Spach, Hist. nat. veg. 2: 322. 1834 (as "section"), nom. invalid. ex Art. 33, pro
parte; Lindley, Intr. nat. syst. bot. (ed. 2): 133. 1836, pro parte; Endlicher, Gen. pl. 1152.
1840, pro parte; idem, Ench. bot. 611. 1841, pro parte; Walpers, Repert. 1: 500. 1842, pro
parte; Lindley, Veg. kingd. 471. 1846, pro parte; Tulasne, Ann. Sci. Nat. Bot. ser. 3. 7: 283.
1847; Grisebach, Fl. Brit. W. I. 135. 1859, pro parte.
Shrubs or trees with glands usually visible in all parts; branchlets terete or
obtusely angled, longitudinallywrinkled,lenticellate, the leaf scars heart- to kid-
ney-shaped with 3 trace scars and a hairy bud. Leaves simple, digitately 1-5-
foliolate, or imparipinnate,the leaflets, if any, mostly articulate;petioles, if any,
longitudinallywrinkled, provided or not with narrow membranouswings; vena-
tion brochidodromousto eucamptodromous.Inflorescences + paniculate or ra-
cemose, occasionally reduced in size, with the side-branchletsand pedicels ar-
Systematic Treatment 23

. : ~.;
................................
............................

I*4.1
. I.

EsenbeckG--
subg.g

* sect Esenbeckia I _,/

* sect.Pochypetulae ? '- ' A
v subg.Oppositifo[ia
. subgLeteriflorens I, X

* -

FIG. 4.
Esenbeckia Distribution of (A) and Piloarpus (B)

?? ~ ~ N^ > s/^'
L-
__________Q_V42_*A

~
FIG. 4.Dsrbto ~fEebci ~ A n
7 ioaps()

P4.loccirpus
*_\IG *...'-o
o s c ( a PB.
24 Flora Neotropica

ticulate and longitudinallywrinkled or ribbed below the nodes, the bracts and
bractlets ? minutelyciliate. Flower buds globose; flowers actinomorphicor only
slightly zygomorphicin the perianth;calyx lobed or toothed, or with nearly free
sepals, persistent, ? minutely ciliate; petals free, or occasionally slightly adnate
to other flower organs but only at the very base, not cohering with each other,
spreadingto reflexed at anthesis; stamens all fertile, as many as the petals; fila-
ments free or occasionally adnateto other organsat the very base, implantedjust
below the disc, accumbent in grooves of the disc or sometimes surroundedat
base by it; anthersheart-shaped,dorsifixed,introrse;disc annularto cup-shaped,
? adnate to the ovary; carpels elevated to immersed half their length in the
receptacle, connate but sometimes only at the base, frequently tuberculateand
bringingforth an apophysis; ovules 1 or mostly 2 per loculus, collateral, pendu-
lous, epitropous, hemitropous;style attached adaxially on the carpels; stigma 1.
Capsulesloculicidaland septicidalor divided into loculicidalmericarps,expelling
the seeds on drying;endocarpsmooth or rarely with impressions of the exocarp
nerves, ochre-yellow, hard, lignified, not cartilaginous, the adaxial part pale
brownish, soft, not lignified;funicles each provided with a triangularor ovate,
acute obturatorwhich covers the hilum, but which is poorly developed when the
seed does not grow; seeds 1 or 2 superposed in each loculus, with the chalazal
area frequentlydistinct on the outside of the testa, ventrally near the base of the
seed; hilum distinct on the adaxial side; a small caruncle sometimes present near
the micropyle; cuticular nucellus always remainingin the form of a hyaline bag
with a thick, brown hypostase; granularendospermsometimes for a part remain-
ing; embryo 1-several, shapedlike the seed and only slightly smallerthanit, with
very thick, plano-convex, eared cotyledons.
Type genus. Pilocarpus Vahl. The name is probably derived from the Greek
words "pilos" (felt hat) and "karpos" (fruit), because the parts of the fruit
(mericarps)resemble a hood.
When first treated by Engler (1874a), the Pilocarpinaecomprisedfour genera,
viz. Esenbeckia, Metrodorea, Pilocarpus, and Leptothyrsa Hooker in Bentham
and Hooker. Later on Engler (1896) transferred Leptothyrsa to subtribe
CuspariinaeEngler. The plano-convexcotyledons of Leptothyrsaagree with sub-
tribe Pilocarpinae;however, the non-globose flower buds (caused by the tubular
corolla) and the linear anthers seem to ally the genus with the genera of subtribe
Cuspariinaeas Hooker (1862) had indicated. (The same argumentholds for Ad-
iscanthus Ducke which was attributedto the Pilocarpinaeby Albuquerque(1977:
7) without explanation). Recently Cowan (1960) added Raulinoa to subtribePi-
locarpinae.

Key to the Genera of Pilocarpinae

1. Flowers in a raceme. 4. Pilocarpus.
1. Flowers in a paniculateinflorescencewhich is occasionallyreducedin size.
2. Leaves opposite and providedwith a sheath. 2. Metrodorea.
2. Leaves alternate,opposite, or aggregated,lackinga sheath.
3. Branchletsunarmed;leaves alternate,opposite, or aggregated;flowers usually pen-
tamerous. 1. Esenbeckia.
3. Branchletsusually spiniferous;leaves opposite; flowers tetramerous. 3. Raulinoa.

ESENBECKIA
1. EssenbeckiaKunth in Humboldt, Bonpland and Kunth, Nov. gen. sp. 7: 191
(=246 quartoed. and reprintCramer1963).25 Apr 1825;Adr. Jussieu,
Mem. Mus. Hist. Nat. 12: 486. after 27 Jun, 1825;Kunth, Syn. pi. 4:
Esenbeckia 25

251. 1826;Martius,Nov. gen. sp. pl. 3: 80. 1829;Pohl, P1.Bras. 2: 42.

1830;Spach, Hist. nat. veg. 2: 343. 1834;Meisner, PI. vasc. gen. 1: 63
and 2: 45. 1837; Endlicher, Gen. pl. 1153. 1840; Grisebach, Fl. Brit.
W. I. 135. 1859; Hooker in Bentham and Hooker, Gen. pl. 1: 299.
1862, sectio 2 excl.; Engler in Martius,Fl. bras. 12(2): 139. 1874;En-
gler in Engler and Prantl, Nat. Pflanzenf. 3(4): 159. 1896; Wilson, in
North Amer. Fl. 25: 200. 1911;Englerin Englerand Prantl,Nat. Pflan-
zenf. (ed. 2) 19a: 280. 1931; Macbride, Field Mus. Nat. Hist., Bot.
Ser. 13: 671. 1949;Cowan, Sellowia 12: 87. 1960;Albuquerque,Anais
Acad. Brasil. Ci. 40: 508. 1968;Cowan and Smith in Reitz, Fl. Ilustr.
Cat. 1 (RUTA) 32. 1973.
Esenbeckiasect. Esenbeckia Hooker in Benthamand Hooker, Gen. pl. 1: 300. 1862,non Esen-
beckia Blume, Bijdr. 118. 20 Aug 1825(=Neesia Blume, nom. cons. (Bombacaceae)),nec
Esenbeckia Bridel, Bryol. univ. 2: 753. 1827 (=Garovaglia Endlicher (as "Carovaglia")
(Musci)).
Euodia ("Evodia") Saint-Hilaire,Bull. Sci. Soc. Philom.Paris (1823): 129. Sep 1823,pro parte;
Saint-Hilaire,Fl. Bras. mer. 1: 79. Mar 1824; non J. R. and G. Forster; non "Evodia"
Gaertner.
PolembryumAdr. Jussieu, M6mMus. Hist. Nat. 12: 519. after 27 Jun, 1825;Mem. Rutac. 136.
Dec 1825;G. Don, Gen. hist. 1: 807. 1831;Schott, Rutac. (11). 1834;Lindley, Intr. nat. syst.
bot. (ed. 2): 133. 1836,pro syn.; Steudel, Nomencl. (ed. 2) 2: 367. 1841(as "Polembrium").
Type species. P. castanocarpumAdr. Jussieu.
ColythrumSchott, Rutac. 13. 1834;Lindley, Intr. nat. syst. bot. (ed. 2): 133. 1836;Meisner,P1.
vasc. gen. 1: 63 and2: 45. 1837,pro syn.; Steudel,Nomencl.(ed. 2) 1: 399. 1840.Type species.
C. puberulum Schott.
Kuala Karsten& Trianain Triana,Nuev. jen. 21. 1854;idem in Karsten,Linnaea28: 428. 1857
("1856"); Miillerin Walpers,Ann. 7: 529. 1869;Trianaand Planchon,Ann. Sci. Nat. Bot.
ser. 5. 14: 306. 1872,pro syn. Lectotype species. K. alata Karsten& Trianain Triana.
Branchlets with roundish or spindle-shapedlenticels. Leaves alternateor op-
posite, simpleor 1-5-foliolate;petioles glabrous.Inflorescencespaniculate;bracts
and bractlets usually persistent. Flowers pentamerous or sometimes heptamer-
ous; calyx lobed, the lobes mostly quincuncialand then at anthesis usually con-
nate and overlappingat base; petals minutely ciliate or not; filamentsaccumbent
in usually shallow grooves of the disc; anthers dorsifixedusually at middle, ver-
satile, with protrudingconnective, yellow, glabrous;disc annular,rosette-like, or
cup-shaped, 5- or 10-lobed; ovary 5-locular, the carpels connate at base or
throughout;ovules 2 per loculus; style terete or obtusely 5-angled. Fruits (4-)5-
locular capsules, usually prominentlynerved on the inside of the exocarp, dehis-
cent septicidally along the dorsal commissures and loculicidally from the base
along the ventral sutures up to the apophysis-tip if present, or up to variable
points when apophyses absent; seeds slightly carinate when boiled, strongly so
when dry; chalazal area less shiningand darkerthan the rest of the testa, or this
area not visible on the outside.
Type species. Esenbeckia pilocarpoides Kunth. Esenbeckia was named in hon-
or of C. G. Nees von Esenbeck, the president of the "Leopoldina," and not as
well for his brotherT. F. L. Nees as Cowan and Smith (1973) mention.
Distribution.Mexico to Argentina(Misiones), and in the West Indies. Fig. 4A.
The genus is divided into three subgenera,one of which has two sections. The
main differences are shown in Table I.

Key to the Subgeneraof Esenbeckia

1. Leaves and side branchletsof inflorescencesopposite. Subgen. II. Oppositifolia(spp. 22-24).
1. Leaves essentially alternate,occasionallysuboppositeor crowded;side branchletsof inflo-
rescences rarelyopposite.
26 Flora Neotropica

Table I
Differences Between the Subgeneraof Esenbeckia

Subgen. Subgen. Subgen.

Esenbeckia Oppositifolia Lateriflorens
Phyllotaxy essentiallyalternate opposite alternate
Inflorescences terminal terminal lateral
Protandryoccurring occasionally always not
False midribof sometimes always not
glandulartissue in
calyx lobes
Aestivationof petals imbricate,sometimes imbricateto valvate valvate
(sub)valvate
Nervationof petals mostly actinodromous camptodromoustending unicostate
to parallel to become supra-
basally
actinodromous
Junctureof filaments midway mostly below midway midway
on anthers
Numberof seeds per I or 2 usually 2 1
loculus
Color of chalazalarea black-brown brown black-brown
if any
Granularendosperm absent present absent
remains

2. Inflorescencesterminal. Subgen. I. Esenbeckia(spp. 1-21).

2. Inflorescenceslateral. Subgen. III. Lateriflorens(spp. 25-26).

Key to the Species of Esenbeckial

1. Inflorescenceslateral;leaves (sub)l-foliolate.
2. Petioles winged; inflorescences separatedfrom the foliage leaves; fruits 1.8-2.3 cm
long. 26. E. cowanii.
2. Petioles not winged; inflorescencesalternatingwith foliage leaves; fruits 1.0-1.1 cm
long. 25. E. almawillia.
1. Inflorescences(sub)terminal;leaves simple or 1-5-foliolate.
3. Leaves simple.
4. Flowers 4-5 mm in diam.; petals coriaceous or rarelythick-papery,with hooked
tip; filamentshairyat the base; disc mostly hairy;fruitssmoothand providedwith
apophyses (when wrinkled,with apophyses: see E. panamensis).
5. Leaves spreadingpubescentbelow, the indumentumclearlyperceptibleto the
touch; apex of leaves acuminate;calyx lobes acuminate;petals thin-coria-
ceous to thick-papery. 15. E. cornuta.
5. Leaves appressed-pubescentbelow; apex of leaves obtuse or slightlyacumi-
nate; calyx lobes obtuse or acute; petals coriaceous.
6. Leaf base attenuate;flowers4-5(-6) mm in diam.; disc and ovary pubes-
cent; fruits obovoidal. 14. E. leiocarpa.
6. Leaf base roundedor obtuse; flowers 5-6.5 mm in diam.; disc and ovary
(sub)glabrous;fruits stellately 5-lobed. 16. E. oligantha.
4. Flowers 10.5-15 mm in diam.; petals papery, with roundedapex; filamentsand
disc glabrous;fruits muricateor echinate, withoutapophyses.

The diam. of flowers and the sizes of flower organs to be measuredafter boiling. The length of
the style to be measuredfrom the place of insertionamongthe carpels.
Esenbeckia 27

7. Style 4.2-4.5 mm long; fruits shorterthan 1.5 cm; endocarpextremely thin.

13. E. hartmanii.
7. Style 3.2-3.5 mm long;fruits2-2.5 cm long; endocarp'normal" (thickerthan
above). 12. E. flava.
3. Leaves compound.
8. All leaves 1-foliolate(caution:E. alata and E. pilocarpoides subsp. maurioides,
which sometimes show only 1-foliolateleaves only (at the tips of the branchlets;
3-foliolateleaves downward,at least in E. alata).
9. Petioles not winged;apex of leaflets rounded,obtuse, or slightly acuminate;
filamentswithoutappendage;fruits withoutapophyses.
10. Hairs of shoot tips and leaves ferrugineous-hyaline;
calyx lobes with false
midribconsistingof glandulartissue; petals thickly coriaceous;fruitsglo-
bose with prickles 2.5-5 mm long. 9. E. grandiflora.
10. Hairs of shoot tips and leaves grayish-white;calyx lobes without false
midrib;petals transparent-papery; fruitsdepressed, stellately-lobed,with
prickles of 7-9 mm long, or with irregularblunt tubercles up to ca. 2.5
mm long.
11. Petiolules absent;petals pubescentbelow; fruits stronglyechinate.
10. E. echinoidea.
11. Petiolules ca. 0.7-1 mm long; petals glabrous (?); fruits irregularly
tuberculate. 8. E. nesiotica.
9. Petioles winged;apex of leaflets mostly distinctlyacuminate;filamentsmostly
with basal appendage;fruits with dorsal apophyses.
12. Flowers 5-8 mm in diam.;petals pale yellow to white; ovary pilose; fruits
(sparsely) muricateor muricateto echinate. 17. E. pilocarpoides.
12. Flowers 7-14 mm in diam.; petals violet at base and margin; ovary
(sub)glabrous;fruits slightly tuberculate. 18. E. amazonica.
8. Not all leaves 1-foliolate.
13. Leaves 3-foliolate, sometimesalso 1-5-foliolateleaves present.
14. Leaves (sub)opposite;apex of leaflets acuminate.
15. Flowers 2.5-5 mmin diam.;petals coriaceous;fruitswith apophyses.
24. E. hieronymi.
15. Flowers 4.5-8 mm in diam.; petals papery; fruits with or without
apophyses.
16. Flowers 4.5-5.5 mm in diam.; calyx lobes and petals mostly gla-
brous, but if pubescent than only below; fruits roundedon the
back. 22. E. febrifuga.
16. Flowers 6-8 mm in diam.; calyx lobes pubescent on both sides,
petals pubescentbelow; fruits with dorso-apicalapophyses.
23. E. densiflora.
14. Leaves alternate;apex of leaflets acuminateor not.
17. Petioles distinctlyor obsoletely winged.
18. Apex of leaflets acuminateor caudate;leaflets subglabrous;fila-
ments with basal appendage.
19. Apex of leaflets(slightly)acuminate;calyx lobes 1.5-1.7 mm
long; filamentswith a basal appendagewhich becomes ex-
tremely swollen immediatelyafter flowering, the filaments
appearingas thoughinserted in a pit of the disc. 19. E. alata.
19. Apex of leaflets rostrate-caudate;calyx lobes 1.1-1.5 mm
long; filamentswith a small basal appendage,not insertedin
a pit of the disc. 17b. E. pilocarpoides subsp. maurioides.
18. Apex of leaflets not acuminateor slightlyso; leaflets subglabrous
or pilose; filamentswithoutappendage.
20. Leaves towardsand in the inflorescencesgraduallyreplaced
by 1-foliolateor simple leaves and finallyby bracts;inflores-
cences consistingof I mainbranch;disc cup-shaped.
7. E. pumila.
20. Leaves not replacedin thatmanner;inflorescencesconsisting
of 1-5 main branches, all arisingfrom the same point; disc
rosette-like. 2a. E. berlandieri subsp. berlandieri.
17. Petioles not winged.
21. Leaflets acuminate.
22. Filamentswithoutbasal appendage.
Ic. E. pentaphylla subsp. australensis.
28 Flora Neotropica

22. Filamentswith basal appendageor appearingas thoughbeing

insertedin a pit of the disc.
23. Petals yellow-greenishor obsoletely violet, (sub)-glab-
brous;filamentswithbasalappendagewhichbecomes ex-
tremely swollen immediatelyafter flowering. 19. E. alata.
23. Petals dark purplish, densely villous above; filaments
with appendageless swollen. 21. E. scrotiformis.
21. Leaflets usually not acuminateor slightly so.
24. Calyx lobes (sub)glabrous;petals mostly (sub)glabrous.
25. Filaments with basal appendage. 20. E. warscewiczii.
25. Filamentswithoutbasal appendage.
26. Petals coriaceous,brownish-purplewith lightbrown
margins. 5. E. collina.
26. Petals papery, white or creamish.
27. Leaflet blade abruptlyending at very base; in-
florescences consistingof 1 mainbranch;petals
papery with coriaceous base, not transparent;
carpels at anthesis with apophysis; fruits with
apophyses in the lower half. 3. E. runyonii.
27. Leaflet blade decurrentat very base; inflores-
cences consisting of 1-5 main branches;petals
+ transparent-papery; carpels at anthesis with-
out apophysis; fruits with apophyses in upper
half. 2. E. berlandieri.
24. Calyx lobes clearly pubescentbelow; petals often pubescent
below.
28. Petals darkviolet; filamentswith basal appendage;seeds
with distinct chalazal area. 20. E. warscewiczii.
28. Petals white or creamish;filamentswithoutbasal appen-
dage; seeds rarelywith visible chalazalarea.
29. Flowers 9-10.5 mm in diam.; carpels with pilose tu-
bercles; style pilose; fruits without apophyses, the
loculi roundedon the back. 11. E. macrantha.
29. Flowers 6-9.5 mm in diam.; carpels with glabrous
tubercles;style glabrous;fruits with apophyses, the
loculi not roundedon the back.
30. Inflorescencesconsisting of 1 main branch;ca-
lyx lobes with false midribof glandulartissue;
style 1.4-1.6 mm long. 7. E. pumila.
30. Inflorescencesconsisting of 1-5 main branches
arisingfrom a commonpoint; calyx lobes with-
out false midrib;style 1.5-5 mm long.
31. Leaves particularlybelow soft to the touch
with spreadinghairs; fruits shorterthan 24
mm.
32. Terminal leaflets sessile or shortly
stalked (up to 5 mm); flowers 6-8 mm
in diam.; style 3 mm long.
2. E. berlandieri (subsp. litoralis).
32. Terminal leaflets longer-stalked (5-11
mm), never sessile; flowers 8.5-9 mm
in diam.; style 3-5 mm long. 6. E. irwiniana.
31. Leaves subglabrousor with appressedhairs;
fruits mostly longerthan 24 mm.
33. Fruits 24-30 mm long. 1. E. pentaphylla.
33. Fruits 12-20 mm long. 2. E. berlandieri.
13. Leaves 5-foliolate,sometimesalso 3-foliolateleaves present.
34. Leaves subglabrous;petals strigillose;disc strigillose;fruits with second-
ary apophyses;seeds with a curved beak, chalazalarea not visible.
1. E. pentaphylla (subsp. pentaphylla).
34. Leaves pilose below, occasionallysubglabrous;petals (sub)glabrous;disc
glabrous;fruits withoutsecondaryapophyses.
35. Leaves appressed-pubescentbelow, occasionally subglabrous;flow-
Esenbeckia 29

ers 6.5-9 mmin diam.;carpelsat anthesiswithoutapophyses, yellow-

brownish;fruits 12-20 mm long, with blunt dorsal apophyses which
are shorter than 3 mm. 2. E. berlandieri (subsp. berlandieri).
35. Leaves with spreadinghairs and soft to the touch below; flowers 9-
9.5 mm in diam.; carpels at anthesis with large apophyses, purple;
fruits 20-25 mm long, with dorso-apicalhorns of 10-13 mm.
4. E. feddemae.

I. Esenbeckiasubgen. Esenbeckia.
Leaves essentially alternate,or suboppositeor crowded towards the tips of the
branchlets.Inflorescencesterminal.Flowers protandrousor not; petals imbricate,
occasionally (sub)valvate, the nervation mostly + actinodromous or parallel,
rarely cladodromous; anthers dorsifixed at middle. Seeds 1 or 2 per loculus;
chalazal area black-brown,if any; granularendospermabsent from seeds.
Esenbeckia was first dividedby Grisebach(1859)who recognizedtwo sections:
Euesenbeckia including E. pentaphylla and E. attenuata (=E. grandiflora) and
Polembryum with the one species E. castanocarpa (=E. pilocarpoides). Because
the latter section includes the type species of the genus, the name Polembryum
cannot be maintained.
Hooker (1862) also divided Esenbeckia into two sections: Esenbeckia and
Metrodorea. He incorporated the genus Metrodorea into Esenbeckia, but since
no other authors except Baillon recognized the congenerity of Esenbeckia with
Metrodorea, this classificationhas no practicalvalue.
Engler (1874a) divided Esenbeckia into two sections (Pachypetalae and Hy-
menopetalae) based on the thickness and shape of the apex of the petals together
with simple or dividedleaves. In 1896he omittedthe characterof the leaf division
because two of the four species which he previously includedunder sect. Pachy-
petalae (E. grandiflora and E. intermedia) appeared to have 1-foliolate instead
of simple leaves. It now appears also that two other charactersof sect. Pachy-
petalae (thick petals with acute tip) are not always positively correlated, since
the following species have thickpetals with obtuse tips: E. collina, E. grandiflora,
E. hieronymi, E. pumila, and E. warscewiczii. Esenbeckia cornuta is intermediate
as to petal thickness and has an acuminate tip. Therefore I have emended the
circumscriptionof Engler's sections.
Esenbeckia sect. Hymenopetalae Englerincludes the type species of the genus
and it has to be renamed sect. Esenbeckia. This section comprises all species
with a basal appendageon the filaments,while those of sect. Pachypetalae have
none.
In the present treatmentthe subgenus is divided into two sections.

Key to the Sections of Subgen. Esenbeckia

1. Filamentslacking basal appendages. Ia. Pachypetalae (spp. 1-16).
1. Filamentswith a basal appendageon the back. Ib. Esenbeckia(spp. 17-21).

la. Esenbeckiasubgen. Esenbeckiasect. PachypetalaeEnglerin Martius,Fl. bras.

12(2): 141. 1874, emend. Kaastra.
Sect. EuesenbeckiaGrisebach,Fl. Brit. W. I. 135. 1859.
Petals yellowish except in E. collina which has purplish-brownpetals; filaments
lacking a basal appendage.
Type species. Lectotype, Esenbeckia leiocarpa Engler.
30 Flora Neotropica

Distribution.Mexico to Argentina, most species occurringin Mexico and SE

Brazil, few in northwesternSouth America. Fig. 4A.
1. Esenbeckiapentaphylla(Macfadyen)Grisebach,Fl. Brit. W. I. 135. 1859;Miil-
ler in Walpers, Ann. 7: 529. 1869;Urban, Bot. Jahrb. Syst. 21: 553.
1896;Wilson, in North Amer. Fl. 25: 201. 1911;Fawcett and Rendle,
Fl. Jam. 4: 181, fig. 58. 1920.
GalipeapentaphyllaMacfadyen,Fl. Jam. 1: 196. 1837;Walpers,Repert. 1: 499. 1842.
Esenbeckiapentaphylla auct non Radlkoferin Millspaugh,Publ. Field ColumbianMus., Bot.
Ser. 1: 401. 1898;non Standley, Contr. U.S. Natl. Herb. 23: 536. 1923(not definitelyindi-
cated because misprintedas Zanthoxylumfagara,but intended);Field Mus. Nat. Hist., Bot.
Ser. 3: 308. 1930 = Esenbeckia berlandieri Baillon ex Hemsley subsp. berlandieri.

Tree to 25 m tall; indumentwith hairs to 0.1(-0.2) mm long; branchlets4-8 mm

in diam., greyish-brown,minutelyappressed-pubescent,becoming(sub)glabrous.
Leaves alternate, 3-5-foliolate, sometimes some leaves 1- or 2-foliolate, petiol-
ulate; petiole semiterete, canaliculate, + ribbed, 2-10 cm long, minutely ap-
pressed-puberulous,becoming subglabrous, tubercle present or not; petiolules
0-4 mm long; leaflets usually obovate or elliptic to oblong, 10-24 x 4-10 cm,
lateralleaflets smaller, the base of the terminalleaflet very long-attenuateand 1-
1.3 cm long, the bases of the lowest leaflets slightlyunequaland shorter, at apex
acuminate,obtuse or rounded, or occasionally emarginate,marginthick, subrev-
olute or not but strongly revolute at base, leaflet blades chartaceous, dull green,
subglabrouson both sides but beset with scattered appressed hairs near base,
venation brochidodromous to camptodromous, ? prominent below, midvein
often impressedtowards base above. Inflorescences 10-25(-36) cm wide, shorter
than or equallingthe leaves, consisting of 2-4(-5) terminal, erect, (very widely)
paniculate parts of 14-23 x 7-24 cm, usually with many flowers, hispid-tomen-
tulose; side branchlets alternate, deciduous when sterile; bracts and bractlets
alternate or subopposite, triangular,to 4.5 x 1.5 mm, minutely appressed-pu-
bescent; pedicels 2-6 mm long; flowers 6.8-8 mm in diam., protandrous;calyx
lobes quincuncial, connate and overlapping at base, depressedly ovate, 1-
1.5 x 1.5-2 mm, rounded at apex, coriaceous with thick base and papery mar-
gins, glabrous above, appressed-strigillosebelow, parallel-nerved;petals decid-
uous, cochlear, rarely quincuncial, ovate to subelliptic, 3.3-4 x 1.7-2.2 mm,
roundedat apex, (thickly)papery, semi-transparentor not, yellowish when dried,
creamish, pale yellow, or white when fresh, papillose above, strigillose below,
venation actinodromousto subparallel,the median nerve somewhat thicker; fil-
aments deciduous or persistent, adnateto disc at base, subulate,flattenedat base,
3-4 mm long and 0.4-0.6 mm thick, glabrous; anthers heart-shaped, 1.1-
1.3 x 0.8-0.9 mm, includinga tip 0.1 mm; disc rosette-like, (5-)10-plicate to 1/4
or 2/5 of its thickness, 0.5-0.8 mm high and 0.2-0.8 mm thick, 1.5-2.5 mm in
diam., with very fine glands, glabrousor minutelystrigilloseat tip; carpels adnate
to the disc, completely connate, 0.6 mm high, charged with conical and clavate
glandularprotuberancesof variable size to 0.3-0.4 mm long, the longest placed
medio-dorsally,(sub)glabrousor minutely strigillose; style inserted near base of
carpels, at first ca. 1.5 mm long, later becoming ca. 2.5 mm long, with a free part
at first0.7-1.1 and later 1.5 mm long, 0.3-0.4 mm thick, glabrous;stigmacapitate,
0.2-0.4 x 0.3-0.4 mm. Fruits 1-2 per infructescence, depressed, stellately-lobed,
truncate at tip when closed, 2.3-3 x 3.4-5 cm when dehisced, loculi slightly
irregularly(somewhat angularly)carinate and provided with an apical, obtuse
apophysis 5-7 mm long which upon dehiscence becomes situated at 1/5-1/8 below
the highestpoint, with or withouta secondaryapophysisca. 5 mmlong at 4/5 below
Esenbeckia 31

the tip, charged with numerous shorter, + angular, tubercles giving a shortly
muricate appearance or merely irregularlytuberculate, wrinkled, with glands,
dehiscent septicidallyfrom 5 mm above base to 2 mm below tip, and loculicidally
to the upper, primaryapophysis; seed 1(-2) per loculus, obliquely tear-shaped,
10-14 x 8-10 x 6-8 mm, with a small, curved beak at the apex up to 0.06 mm
long, testa red- to dark brown, linear-colliculatewith interspaces 0.1-0.2 mm
long; chalazalarea visible or not, to 0.6 mm long; hilum0.8-1 mm broad;embryo
1, cotyledons strongly unequal with ears 0.8 mm long, radicle 0.05-0.7 mm, not
shorter than the plumule, projectingbeyond the ears or not.
Distribution.Guatemala,Belize, Panama,Jamaica,and Colombia. Fig. 6.
The species is divided into 3 subspecies.
Key to the Subspecies of Esenbeckia pentaphylla
1. Full grown but not yet maturefruitsglabrous;leaves 3-5-foliolate;disc and carpels strigil-
lose or glabrous.
2. Leaves 3-5-foliolate; apex of leaflets not acuminateor slightly so; flowers 7-8 mm in
diam.; disc and carpels strigillose;fruits with secondaryapophyses. la. subsp. pentaphylla.
2. Leaves 3-foliolate;apex of leaflets usually acuminate;flowers 6.8-7 mm in diam.; disc
and carpels (sub)glabrous;fruits without secondaryapophyses. lb. subsp. belizensis.
1. Full grown but not yet maturefruits minutelypubescent;leaves 3-foliolate;disc and ovary
strigillose. Ic. subsp. australensis.

la. Esenbeckiapentaphylla(Macfadyen)Grisebachsubsp. pentaphylla.

Fig. 5A-B.
Tree 7.5-15 m tall. Leaves 5-(some 3- or 4-)foliolate;leaflets 10-22 x 4-9 cm,
rounded and occasionally emarginateat apex, or obtuse and slightly acuminate.
Inflorescences very widely paniculate with many flowers. Flowers 7-8 mm in
diam., pale yellow, odoriferous;petals to 3.5 x 2 mm, thickly papery, not trans-
parent; filamentsto 3.5 mm long; disc 2-2.5 mm in diam., strigillose at tip with
hyaline hairs 0.1-0.3 mm long; carpels charged with protuberanceswhich are
sparsely minutely strigillose with hyaline hairs 0.1 mm long. Fruits nearly gla-
brous, 2 x 2.6 cm when closed, and 2.3-3 x 4.5-5 cm when dehisced; loculi
provided with a secondary apophysis ca. 5 mm long at ca. 4/ below tip, and with
numerousshorter, + angled tubercles giving a shortly muricateappearance.
Type. Herb. Macfadyen s.n., Jamaica: Rd. from St. Michael's Chapel to
Green Valley Works, and below Moccha Works, Port Royal, Jul, fl & fr (lecto-
type, K-Herb. Hooker 1867,the fl specimen with drawing;isolectotype, K-Herb.
Hooker 1867, fl).
Distribution. Jamaica: woodlands and rocky limestone hills, alt. 160-750 m.
Flowering:Jun-Nov(-Jan).
Specimens examined. JAMAICA.Cornwall:Britton 577 st (NY), 2268 fr (NY); Harris 7057 fr
(F, NY), 9074 fl (NY 2x), 10285fr (BM, C, F, NY, P, US), 12367fl (BM, F, MO, NY, P). Middlesex:
Alexander s.n. fl (K, NY); Britton 3693 fl (NY); Harris 8209 fr (A, BM), 11195 fl (BM, F, NY, US);
Macfadyen s.n. (Dec 1843) fl (K). Surrey: Harris 5664 st (F, NY). County unknown: Harris 5629 fl
(BM, S, Z); Macfadyen s.n. fr (K 2x); March s.n. (1888) fl (GH, GOET 2x, K); Purdie s.n. fl (MPU),
s.n. fl, fr (K); W. M. Wright 12 fl (BM).
Local name. Wild orange(several records).

lb. Esenbeckiapentaphylla (Macfadyen) Grisebach subsp. belizensis (Lundell)

Kaastra, Acta Bot. Neerl. 26: 471. 1977. Fig. 5C-D.
Esenbeckiabelizensis Lundell, Contr. Univ. MichiganHerb. 6: 33. 1941.
Tree with straight bole of 6-25 m tall and trunk 5-50 cm in diam. Leaves
32 Flora Neotropica

3cm

FIG. 5. Esenbeckia pentaphylla. A-B, subsp. pentaphylla. C-D, subsp. belizensis. E-F, subsp.
australensis. A, habit (Harris 12367, NY). B, fruit (Harris 10285, C). C, fruit (Contreras 7358, F).
D, seed (Contreras 7358, F). E, fruit (Stern et al. 1822, MICH). F, indument of fruit (Stern et al.
1822, MICH).
Esenbeckia 33

3-foliolate, some 1- or 2-foliolate, leaflets to 24 x 10 cm, acuminatewith the very

tip blunt, or obtuse, rarelyroundedor emarginate.Inflorescences widely panicu-
late with many flowers. Flowers 6.8-7 mm in diam., creamish, scented; petals to
3.5 x 2.2 mm, papery, semitransparent;filamentsto 3.5 mm long; disc 1.5-2 mm
in diam., glabrous;carpels (sub)glabrous.Fruits 2.6-3 x 3.5 cm when closed and
4.6-5 cm wide when dehisced, when ca. 7 mm long sparsely puberulous with
hairs 0.05 mm long but glabrous when mature, the loculi less sharply edged-
tuberculatethan those of subsp. pentaphylla and without secondary apophyses,
not muricate, merely irregularlytuberculate,ratherbroad and flattenedlaterally
at base with ridges along the margin.
Type. P. H. Gentle 2934, Belize. Stann Creek District: Middlesex, 23 Jul
1939, fl (holotype, MICH).
Distribution.Guatemalaand Belize: in high primaryforest and on ridges, alt.
60-800 m. Collected in flower Jul-Sep.
Specimens examined. GUATEMALA.Peten: Contreras6406 fr (U), 7358 fr (F, NY, S, U), 7370
fr (F, U). Izabal:Record G20 fr (US).
BELIZE. El Cayo District:P. H. Gentle 8897 fl, fr (BM, F, G, MICH, NY, S, UC, US). Stann
Creek District:P. H. Gentle 3069 fl, fr (A, F, MICH, NY, K); Schipp 248 fl (A, BM, F, G, GH, K
2x, MICH, MO 2x, NY, S, UC, US, Z). Toledo District:P. H. Gentle4758 fl (UC, US), 4797 fl (F,
S), 5130 st (DS, G, US); Hummel38 st (K); Schipp S-643 fr (A, BM, F, K, MICH, MO, NY, S, Z).
District unknown: A. P. F. G(entle?) 193 fl (F).

Local names and uses. Belize: candlewood (Hummel 38); verde lucero
(APFG(entle?) 193). Possible use: the wood very hard and close-grained, white
when felled, later on creamish, yellow, and brown.
Palmer 181, a fruitingcollection from near Tampico, Tamaulipas,Mexico, has
characters intermediate between Esenbeckia pentaphylla and E. berlandieri, and
suggests a close relationshipbetween the taxa. The leaves are much like those
of E. pentaphylla subsp. pentaphyllabut at the base are pilose above with spread-
ing hairs like those of E. berlandierisubsp. berlandieri. The calyx is persistent
and seems to be glabrous;a glabrouscalyx has never been observed in E. pen-
taphylla. The fruits resemble those of E. pentaphylla subsp. belizensis, both in
size (2.3-2.7 x 3.8-4.3 cm when dehisced) and in roughness (very irregularly
tuberculate). The broad seeds (9-11 x 7-8 x 5-6 mm) with the shortly curved
beak and without visible chalaza, agree with those of E. pentaphylla. This col-
lection cannot be classified satisfactorilywithoutflowers;however for the present
possibly the best place is in E. pentaphylla.
lc. Esenbeckiapentaphylla(Macfadyen)Grisebach subsp. australensisKaastra,
Acta Bot. Neerl. 26: 471. 1977. (See Addenda). Fig. 5E-F.
Tree with trunk 10-15 m tall and 5-25(-40?) cm in diam.; leaves 3-foliolate,
leaflets acuminateat apex with obtuse acumen, or sometimes obtuse or rounded
at tip. Inflorescences paniculate, loosely flowering. Flowers ca. 8 mm in diam.,
fragrant;petals 3.7-4 x 2-2.2 mm, papery, semitransparent,white when fresh;
filaments 3.5-4 mm long; disc ca. 2 mm in diam., densely strigillose with hairs
0. 1-0.3 mm, like the carpels. Fruits 2.4-2.8 x 2.9 cm when still closed, and 3.4-
4 cm wide when dehisced, densely pilose when immature,becoming glabrous;
seeds shining, chalazal area visible.
Type. Dugand G. 536, Colombia. Atlantico: El Pajar forest, alt. 60-300 m,
29 Mar 1934,fr (holotype, F-726782;isotypes, G, US 2x).
Distribution. Panama and N Colombia (Atlantico and Magdalena).Collected
in flower in Jul.
34 Flora Neotropica

Specimens examined. PANAMA. Los Santos: Vic. of Tonosi, along QuebradaOcho Paso, Stern
et al. 1822fr (MICH).
COLOMBIA.Magdalena:Ariguani,Romero C. 297 fr (COL);mun. Pivijay, Monte Rubio, 9062 fl
(COL).
Local names and use. Colombia: loro (Romero C. 9062). Possible use: the
wood extremely hard, white-yellowish(Dugand G. 536).
The leaves of the present subspecies are like those of subsp. belizensis, while
the fruits are much like those of subsp. pentaphylla.

2. EsenbeckiaberlandieriBaillon ex Hemsley, Biol. centr.-amer., Bot. 1: 170.

1879;Rose, Contr. U.S. Natl. Herb. 5: 111, t. 3, figs. 8-9. 1897;Wil-
son, in North Amer. Fl. 25: 202. 1911; Standley, Contr. U.S. Natl.
Herb. 23: 536. 1923, pro parte.
Shrub or tree, 1-15 m tall with trunk to 20(-25) cm in diam., branchlets2-5
(-10) cm in diam., grayish-brownor grayish-green,hoary or minutelypubescent,
(finally) becoming glabrous. Leaves alternate, 3-5-foliolate, occasionally some
leaves 1-foliolate,with stalked or sessile leaflets; petiole (slightly)canaliculateat
least towards tip, ? ribbed, winged or not, 1-7(-12) mm long and to 3 mm thick,
indument like that on the young branchlets;petiolules, if any, canaliculate, 0-
1 mm long; leaflet blades obovate or elliptic, (3.5-)6-16(-22) x (1.4-)2.5-9 cm,
the lateral ones smaller than the terminalleaflet, attenuate or in sessile leaflets
cuneate, the base to ca. 7 mm long, especially the lateralleaflets unequalat base,
rounded, or obtuse and somewhat shortly acuminateat apex, the very tip often
emarginate,marginthick, ? stronglyrevolute, particularlytowards base, blades
chartaceous, somewhat shining above, dull beneath, nearly glabrous or densely
pilose, venation camptodromous, + prominent, midvein plane or impressed
above. Inflorescences consisting of 1-5 terminalor subterminal,erect, (widely)
paniculatepartialinflorescences which are as long as the leaves or longer and to
26 cm wide in all, usually tomentulose; side branchletsalternateor subopposite;
bracts (narrowlyor broadly)ovate or triangular,1.5-3.5(-5) x 1-2 mm, minutely
pubescent below or (sub)glabrous;pedicels 1-4 mm long; bractlets 2, alternate
or opposite. Flowers 5.5-9 mm in diam., fragrant;calyx lobes quincuncial or
rarely cochlear, very broadly ovate, 0.8-2.2 x 1-2 mm, rounded or obtuse at
apex, coriaceous, base thick, marginsmembranous,venation actinodromousor
parallel;petals imbricate,spreadingto reflexed, elliptic to ovate, 3-4.5 x 1.5-2.4
mm, obtuse or rounded at the apex, ? transparent-papery,white or creamish,
glabrous or papillose above, glabrous or pilose below; filaments subulate, flat-
tened towards base, 2-4 mm long and 0.3-0.8 mm thick, glabrous;anthersovoid
to heart-shaped,(0.8-)1.1-1.5 x (0.5-)0.8-1.4 mm, includinga mucro 0.05-0.20
mm, disc rosette-like or cup-shaped, 10-plicateup to /2 or /3 of its thickness,
later on often 5-lobed, 0.3-0.9 mm high and 0.2-0.5 mm thick, 1.5-2.5 mm in
diam., thick-fleshy, with glands, glabrous;carpels adnate to the disc in the lower
part, free distally, 0.4-0.6 mm high, glabrous, charged with glandularprotuber-
ances or tubercles; style terete or clavate, 0.2-0.3 mm thick, glabrous; stigma
capitate, slightly 5-lobed or not, 0.2-0.4 x 0.3-0.5 mm, with or without 5 large
glands. Fruits depressed, stellately (4-)5-lobed when dehisced, 1.2-2 x 1.7-4.5
cm, the loculi roundedat base, provided with a dorso-apical,blunt apophysis to
3 mm or 4-15 mm, rugose, with glands, glabrous, dehiscent septicidallyfrom 3-
5 mm above base to tip; seed usually I per loculus, obliquely tear-shaped,flat-
tened at or near the base, (7.5-)10-15 x 5-10 x 4.5-7.5 mm, the apex curved or
straightand 2-3 mm long, testa dark or mediumbrown, linear-colliculate,inter-
spaces 0.2-0.3 mm long in parallel fascicles which are criss-crosswise mingled
Esenbeckia 35

A ~
---
--p"1".
. E.pentaphylla
v. E.pentaphylla
(^^/"^tt^^,..^^
subsp.pentophylla 5 ?: ?, o 0
,,
3 ''*. -;-
subsp.belizensis 1- :V \ S /
subsp.australensis
E.pentaphyl.a -, . . . . . . -

1 -I t . _ ,I .

* E.berlandiel
subsp.berlandiern l '
* E.berlondieri
subsp.Ilitoralis
v E
--
t -'-,-_-'-
E.berland-ier
berlandierisubsp
subsp.acapulcensis
acopulcensist - ----,---- 1 ...-
* E.runyonii i s' , S '
' 1)
o E.feddemce ? ( , ; - -
i c

v 'E.colina subsp.conspecta . , J .

cl '

? E.collinasubsp.cotlina :', :
' E.colino subspconspecto ' i
* *E. c
F.nesiotica
nesiotica /t _ '
* E.echinoidea
echinoidei7 - .-
* E.mucronth .

FIG. 6. Distribution of some species of Esenbeckia.

36 Flora Neotropica

with each other; hilum 0.5-4.0 mm broad; embryo 1, cotyledons unequal, ears
0.5-1 mm, radicleusually projectingbeyond the ears and to 1.5 mm long, plumule
2-leafed and 0.2-0.8 mm long.
Distribution.The species ranges from Mexico to Panama(Fig. 6B). Three sub-
species can be distinguished,one allopatricand the other two overlapping.

Key to the Subspecies of Esenbeckia berlandieri

1. Leaves soft to the touch, particularlybelow, with spreadinghairs; perianthpilose with
spreadinghairs. 2b. subsp. litoralis.
1. Leaves (sub)glabrousor minutelyappressed-pubescent,not soft to the touch; perianthgla-
brous or sometimes minutelyappressed-puberulous.
2. Flowers 6.5-9 mm in diam.; carpels only postflorallywith apophyses;fruits depressed,
stellately-lobedwith apophyses to 3 mm long. 2a. subsp. berlandieri.
2. Flowers 5.5-7 mm in diam.; carpels provided with apophyses already in bud; fruits
subglobosewith horn-likeapophyses of 4-15 mm long. 2c. subsp. acapulcensis.

2a. EsenbeckiaberlandieriBaillon ex Hemsley subsp. berlandieri. Fig. 7A-C.

Esenbeckiaovata Brandegee,Univ. Calif. Publ. Bot. 7: 327. 1920;Standley, Contr. U.S. Natl.
Herb. 23: 536. 1923,pro syn. TYPE. Purpus8419, Mexico. Veracruz:Nr. Acasonica, Aug
1919,fl (holotype, UC-200823Herb. Brandegee;isotypes, DS, US).
Esenbeckia yaaxhokob Lundell, Lloydia 4: 50. 1941, syn. nov. TYPE. Steere 2956, Mexico.
QuintanaRoo: Isla de Cozumel, San Miguel, nr. coast, 6-8 Aug 1932,fl (holotype, MICH;
frag. F ex MICH).
Esenbeckiaberlandieriauct non Baillon, Adansonia10: 151. 1871,nom. invalid.
Branchlets hoary to minutely appressed-pubescentwith most hairs 0.1-0.2
(-0.5) mm long, becoming glabrous. Leaves 3-5-foliolate; petiole semiterete, oc-
casionally with wings 0.1-0.2 mm broad, the wings with ears 0(-0.3) mm long,
distal tubercle present; leaflet blades with a (long-)attenuatebase to 1 cm long,
(sub)glabrousabove but minutely puberulousat least at base with hairs 0.2-0.3
mm long, mostly appressed-puberulousbelow particularlytowards base and the
midvein with hairs 0.1-0.3(-0.6) mm long, rarely subglabrousbelow. Partialin-
florescences 20 x 14 cm, hispidulous-tomentulosewith most hairs 0.2-0.4 mm
long; bracts mostly minutelypuberulouson both sides with hairs to 0.2 mm long.
Flowers 6.5-9 mm in diam.; calyx lobes 1-2 x 1-2 mm, glabrous or sometimes
appressed-puberulousbelow with hairs 0.05-0.1 mm long; petals persistent, pa-
pillose above, glabrous below, venation actinodromousto parallel, the median
nerve thicker; filamentspersistent; carpels apically charged with protuberances
varying from 0.1 to 0.6 mm long, postflorallyprovided with an apophysis of ca.
0.8 mm long, glabrous but some of the protuberanceswith a single apical hair;
style (1.5-)2-3.2 mm long, projecting 1-2.2 mm beyond the ovary. Fruits de-
pressed, stellately-lobed, to 3.4 cm broad when dehisced, brown, provided at
13 below tip with a blunt apophysis to ca. 3 mm long, slightly muricateto irreg-
ularly tuberculate,the tubercles to 2 mm long, slightlyirregularlycarinateor not,
exocarp nerves observable externally on the sides; seed with a straight,beaked
apex 2-3 mm long, chalazal area indistinctlybounded, ca. 2-4 x 2-4 mm, some-
what darkerthan the rest of testa when ripe.
Type. Berlandier3125. Mexico. Tamaulipas:Nr. Tampico, Jan 1831, fr (ho-
lotype, K; isotypes, GH, MO-Herb.Engelmann,NY-Torrey Herb., US ex GH).
Distribution.Eastern Mexico, from Tamaulipasto Yucatan. Habitats diverse,
moist or dry, e.g. on rocky, bushy slopes, and woods, primarydeciduousforests,
regenerationareas, thorny woodlands; to 1400 m alt. Flowering Jun-Aug.
Esenbeckia 37

*t

3cm
U,U

?: * ; G : .
'.

FIG. 7. Esenbeckia berlandieri. A-C, subsp. berlandieri. D-E, subsp. litoralis. F-G, subsp.
acapulcensis. A, habit (Ventura A. 3935, MICH). B, fruit (Ventura A. 2843, MICH). C, seed (Ventura
A. 2843, MICH). D, flower bud (King 760, MICH). E, flower (King 760, MICH). F, fruit (Palmer
175, F-3683). G, dehisced fruit (Palmer 175, F-1 162964).
38 Flora Neotropica

Specimens examined. MEXICO. Tamaulipas: Berlandier s.n. (Nov 1830) bud (GH); Crutchfield
& Johnston 5715 fl (MICH);King 4012 fr (F, MICH, NY, UC, US); Palmer 497 fl (GH, US). Vera
Cruz: Chiang 50 fl (GH), 114 fr (F, GH), 294 fr (F, MICH); Crutchfield & Johnston 5686 fl (MICH),
6100 fr (MICH, NY, UC); Dorantes et al. 1250 fl (U), 1325 fl (U); Purpus 6161 fl & fr (BM, F, GH,
MO, NY, UC, US), 8428 fl (GH, MO, NY, S, UC), 8651 fr (GH, MO, NY, UC, US), 8895 fr (GH,
MO, NY, UC, US); Ventura2843 fr (DS, MICH),3935 fl (CR, DS, MICH), 4331 fr (DS, MICH).
Oaxaca: Janzen s.n. (20 Nov 1963) fr (MICH, UC). Chiapas: Miranda 5487 fr (US). Yucatan: Enriquez
330 fr (US); Flores s.n. (1931) fl & fr (F 2x); Gaumer752 fl (A, BM, C, E, F, K, LE, M, MO 2x, NY
2x + photo, S, US 2x, W, WIS), s.n. (comm. GodmanAug 1886)fl (K); Lundell & Lundell7532 fl
(A, F, MICH),7983 fl (MICH).Terr. QuintanaRoo: Cozumel,Lundell& Lundell7831 fl (A, DS, F,
MICH, US); Millspaugh 1475 fr (F, photo NY); Steere 2956 fl (F, MICH). State unknown:Pozo
Zopilote or Paso Sapilote:Liebmann6414 (=probably 6614) fl (US), 6614 fl (C, MO, UC), 15005 fl
(C, F), 15015 fl (C), s.n. (Aug 1841) fl (M). Locality unknown: Liebmann 4198 fr (C 2x), s.n. fr (NY).
Local name. Hokob in Yucatan (Flores s.n. (1931)); yax-hokob (accordingto
Radlkofer(1898)from Gaumer752).
This subspecies is near to Esenbeckiapentaphylla. Evidence of this is e.g. the
occurrence of 5-foliolateleaves, the calyx which appearsto be frequentlypilose,
and the petals occasionally beset with few hairs. Other indications may be the
misidentifications of Palmer 181 (see under E. pentaphylla), and of Gaumer 752
(see Radlkofer, 1898:401).
I agree with Lundell (1941) that the relationshipof E. yaaxhokob is closer to
E. berlandieri than to E. pentaphylla. Differences of E. yaaxhokob from the latter
species are e.g. the (nearly)glabrouspetals, the shorter base of the leaflets, and
the straightapex of the seeds. The leaves of E. berlandieri,however, are, though
mainly 3-foliolate, partly 5-foliolate as well, and the reverse occurs in E. yaax-
hokob. The indumentof the inflorescence is alike in the two species. The spec-
imens of E. yaaxhokob studied by Lundell show the calyx nearly glabrous or
appressed-puberulous,like that of E. berlandieri subsp. berlandieri. Because
there are no distinct differences between the two taxa I decided to reduce E.
yaaxhokob to synonymy under E. berlandieri subsp. berlandieri.
Nonfruitingspecimens of the present subspecies are sometimes muchlike those
of E. runyonii. Some differences are: not abrupt ending of the leaflet base; the
inflorescence of 1-5 partial inflorescences instead of only 1; and calyx lobes,
petals, and filamentsin general narrower,while the style is longer.
The name Esenbeckia berlandieriwas first mentioned by Baillon (Adansonia
10: 151. 1871)in his original description of Picrella (trifoliolata). Although the
name was accompaniedby a diagnosis, it was only mentioned incidentallyand
therefore,underArt. 34(3)of the ICBN, not validly publishedat that time. Hems-
ley validated E. berlandieri 8 years later with a reference to the diagnosis of
Baillon. He saw only Berlandier3125 at K, and this specimen therefore is the
holotype. Baillon based his name on collections of Berlandierand Virlet d'Aoust.
In his type herbarium(P) there are no specimens of E. berlandieri.Virletd'Aoust
1240 (in Baillon's herbarium)and Berlandier 1404 (in the general herbariumat
P), however, seem to have been used for the diagnosis of E. berlandieriBaillon.
They agree completely with the description,but both representHelietta parvifolia
Bentham, the latter being even one of its syntypes. FortunatelyBaillon did not
validly publish his name, therefore there is no need to replace the well known
name E. berlandieri.
Incidentally, on account of the descriptionwith plate 10, it is possible that the
poorly known genus Picrella is also referable 'o Helietta.
2b. EsenbeckiaberlandieriBaillon ex Hemsley subsp. litoralis (Donnell Smith)
Kaastra, Acta Bot. Neerl. 26: 471. 1977. Fig. 7D-E.
Esenbeckia litoralis J. Donnell Smith, Bot. Gaz. (Crawfordsville)23: 242. 22 Apr 1897;Rose,
Contr.U.S. Natl. Herb. 5: 111.27 Aug 1897;J. DonnellSmithin Durandand Pittier,Primit.
Esenbeckia 39

fl. costaric. 2: 74. 1898;Wilson, in North Amer. Fl. 25: 202. 1911;Lundell, Contr. Univ.
MichiganHerb. 6: 34. 1941.
Esenbeckiapilosa Lundell, Contr. Univ. MichiganHerb. 6: 34. 1941, syn. nov. TYPE. Matuda
596, Mexico. Oaxaca:Tehuantepec,26 Jul 1936,fl (holotype, MICH;isotypes, F ex MICH,
US).
Tree to 8.5 m tall with trunk2.5 cm in diam.; wood hard,yellow-brown(Poveda
218); branchletshoary to velvety-tomentulosewith spreadingwhite-hyalinehairs
0.5-0.8 mm long, becoming ? appressed-pubescentand finally glabrous, leaf
scars surroundedwhen young along the upper marginwith sericeous hairs to 0.8
mm long. Leaves (1-)3-foliolate, occasionally some 4- or rarely5-foliolate;petiole
semiterete;leaflet blades with a (shortly)attenuate, or cuneate sessile base to 0.5
cm long, pilose above with spreadinghairs to ca. 0.5 mm long, midvein very
densely pilose, indumentbelow like that above but the main veins ? tomentose,
the hair-cover on both sides but especially beneath soft to the touch, venation
prominent.Partialinflorescences 6-14 x 3-8 cm; branchletsstrongly obtuse-an-
gled, hoary like the vegetative branchlets; bracts beset with few hairs above,
densely pilose below with spreadinghairs ca. 0.5 mm long. Flowers 6-8 mm in
diam., protandrous;calyx lobes 1.7-2.2 x 1.5-1.7 mm, glabrous above or with
few hairs at base, pilose below with spreadinghairs to 0.8 mm long; petals per-
sistent, glabrous above, pilose beneath like the sepals, venation actinodromous
with the median nerve distinctly thicker;filamentsoften deciduous; carpels pro-
vided already in bud with apical apophyses, these 0.4-0.5 mm long, provided
with few glandularand glabrousprotuberancesto 0.2 mm long (to 0.5 mm after
sheddingof the pollen); style 3 mm long, projecting1-2.3 mm beyond the apophy-
ses. Fruits ? stellately-lobed when closed, 1.8-2.2 x 2-2.5 cm, when dehisced
1.5-2 x 2-3.5 cm, the loculi provided /3 from the tip with a subobsolete, blunt
tubercle-likeapophysis 2-3 mm, obsoletely tuberculate;seed with a curved beak
at apex to 0.3 mm long, rarely carinate on the back, testa firm, to 0.4-0.5 mm
thick when mature, chalazal area not visible.
Type. Pittier 34 (Inst. phys.-geogr. nat. CR 2777; edidit J. Donnell Smith 8098
(Pittier & Durand)), Costa Rica. Puntarenas:Along coast of Bahia de Salinas, 3
Jul 1890, fl (lectotype, US-1364873Herb. J. Donnell Smith, photo F, NY, frag.
of lectotype NY; isolectotypes, BR 2x, CR).
Distribution.Mexico (Oaxaca) to Panamabut absent from Belize.
Specimens examined. MEXICO.Oaxaca:King 760 fl (MICH), 1483 bud (MICH, NY, US), 1562
fl (US), 1652A fl (MICH, US); Lathrop 5936 bud (US), 5941 fl (MO); Miranda 8199 fr (US); Webster
et al. 13006 fl (GH, MICH); LI. Williams 9681 fl (F).
GUATEMALA. Zacapa: Standley 74036 fr (F); Steyermark 29345 fl, fr (F).
HONDURAS. Morazan: Molina & Molina 25769 fl (F, NY). El Paraiso: Molina 168 bud (F), 181
bud (F, GH, MO, UC, US), 502 fr (F), 4017 fl (F, GH, US), 7484 bud (F, US); Standley 15577fr (A,
F, US); Standley et al. 548 fr (F, NY, US); Webster et al. 12031 fl (MO, U, US); L. O. Williams &
Molina R. 10228 fl (F, MICH, MO 2x, UC), 11050 fr (F, US). Choluteca: L. O. Williams 14321 bud
(F, GH); L. O. Williams & Molina R. 16716 bud (F).
EL SALVADOR.Chalatenango:Calder6n2462 fr (BM, F, G, US).
NICARAGUA.Nueva Segovia: 0rsted (6?) (Jan 1848)dec. fr (MPU). Chontales:Standley 9363
st (F), 9377 st (F), 9495 bud (F).
COSTARICA. Guanacaste:Howell 10235fl (F, GH, NY, US); Poveda 218 fl (CR).
PANAMA. Panama: Allen 982 fr (F); Bartlett & Lasser 16643 fl (MICH).
Specimensintermediatebetween subsp. berlandieriand subsp. litoralis: GUATEMALA.Zacapa:
Standley 73781 fl (F). Chiquimula: Standley 73711 st (F), 73742 fr (F), 74303 fr (F, G).
HONDURAS. Comayagua:Molina 14338fl (F, NY). El Paraiso:Molina 10079fl (F, US); Webster
et al. 12012 fr (GH, MO, US).

Typical specimens of subsp. litoralis have leaflets soft to the touch especially
underneath,and often almost sessile; the perianthis pilose with spreadinghairs
to 0.8 mm long; the fruits are obsoletely tuberculate.
40 Flora Neotropica

Specimens from Honduras and Guatemala are intermediate between subsp.

berlandieriand subsp. litoralis. The perianthof these specimens is subglabrous,
the leaves are (long-) attenuate and frequently appressed-pubescent,though in
some specimens leaflets occur which are shortly-pilosewith spreadinghairs.
The type of E. pilosa fully agrees with the descriptionof E. berlandierisubsp.
litoralis. Esenbeckiapilosa therefore cannot be maintainedas a separate taxon.
The type of the present subspecies was originally mentioned as Pittier s.n.
(3317 CR). This was an apparentmistake. In 1898J. Donnell Smith correctedthis
by indicating the lectotype as Pittier 2777, number 3317 being Citrus medica
Linnaeus. As a matter of fact I could not trace Pittier 3317 under Esenbeckia.
Thus the lectotype collection is Pittier (in CR with field number34) Inst. phys.-
geogr. nat. CR 2777. This collection was also distributed by J. Donnell Smith
under number8098. The specimen in US bears both numbers,in BR 2 specimens
are present under 8098 only.
The leaves are reportedas having a citrous or ratherdisagreeableodor. 0rsted
6 has smaller apophyses.

2c. EsenbeckiaberlandieriBaillon ex Hemsley subsp. acapulcensis(Rose) Kaas-

tra, Acta Bot. Neerl. 26: 471. 1977. Fig. 7F-G.
Esenbeckiaacapulcensis Rose, Contr. U.S. Natl. Herb. 5: 111, t. 3, figs. 1-7. 1897;Standley,
Contr. U.S. Natl. Herb. 23: 536. 1923, pro syn.
Indumentof hairs 0.1-0.2 mm long; branchletsappressed-puberulous,becom-
ing glabrous. Leaves 3-foliolate; petiole terete, semiterete and canaliculate to-
wards the tip; leaflet blades with long-attenuatebase to ca. 1 cm long, glabrous
on both sides but + appressed-pubescentat base, or quite appressed-pubescent
particularlybelow. Partialinflorescences (5)10-13 x 4-10 cm, appressed-pubes-
cent to somewhat tomentulose, often hoary, the branchlets tomentulose to mi-
nutely velvety towards the tips; bracts minutely pubescent distally above, ap-
pressed-puberulousbelow or (sub)glabrouson both sides. Flowers 5.5-7 mm in
diam.; calyx lobes 0.8-1 x 1.2-1.5 mm, glabrous; petals deciduous, glabrous,
venation imperfectlyacrodromousbut hardlyvisible except for the mediannerve;
filamentsdeciduous; carpels apically provided with apophyses 0.6-0.7 x 0.5-0.6
mm which are glabrousand very densely chargedwith roundishglandulartuber-
cles, the longest tubercles occurringmedio-dorsallyon the loculi; the tubercles
enlargingimmediatelyafter floweringand developing hyaline, spreadinghairs to
0.3 mm long, which graduallydisappear;style 1.1-1.4 mm, free for 0.5-0.6 mm.
Fruits subglobose, depressed and stellately-lobed,ca. 2-3.2 x 2.7-3.5 cm when
closed, when dehisced 1.7-2 x 3.3-4.5 cm, with 5 dorso-apical,erect, obtuse horn-
like apophyses of 4-15 mm long, loculi medio-dorsallystrongly carinate, irregu-
larly (shortly) tuberculate-muricate,the tubercles to 5 mm; seed with a shortly
curved beak at apex, chalazal area not visible.
Type. Palmer 175, Mexico. Guerrero:Acapulco, Dec 1895, fl, fr (lectotype,
US-256910, the flowering part, photo F, NY; isolectotypes, DS, F 3x, GH, K,
MO, NY, UC, US-256908, photo K, NY).
Distribution.Mexico, allopatricwith the other subspecies.
Specimens examined. MEXICO.Chihuahua:Hinton 10924fl, fr (BM, F, G, GH, K, MO, NY, S,
US). Jalisco: McVaugh & Koelz 1672 fr (MICH). Colima:McVaugh23032 fr (MICH). Guerrero:
Herb. Haenke s.n. (PR 28) fl (PR 4x).

Differences of the infrequentlycollected subsp. acapulcensis with the other

subspecies are found in the smallerflowers and shorter style, and particularlyin
Esenbeckia 41

the fruits with the long dorso-apicalhorns. I cannot agree with Standley (1923)
who considered it as synonymous with E. berlandieri(pro sp).

3. EsenbeckiarunyoniiC. V. Morton, J. Wash. Acad. Sci. 20: 136. 1930(as "E.

runyoni"); Engler in Engler and Prantl, Nat. Pflanzenf. (ed. 2) 19a:
282. 1931. Fig. 8.
Large shrub or small round-toppedtree, 4-6 m tall, with trunk 10-25 cm in
diam., bark smooth, gray; indumentwhitish, the hairs to 0.2 mm long; branchlets
2-6 mm in diam., light gray or sometimes darker,minutelypubescent, becoming
glabrous, young shoots hoary, leaf scars with elevated margin.Leaves alternate,
3-foliolate, occasionally some leaves 1-foliolate, with stalked leaflets; petiole
semiterete, flattened or canaliculateabove, slightly ribbed, 1-5(-8) cm long, mi-
nutely pubescent or somewhathoary, commonly with a triangulardistal tubercle;
petiolules canaliculatelike the base of the midveins, 0-4 mm long; leaflet blades
elliptic, 4.5-10(-12) x 2-5 cm, (long-)attenuateand rather abruptly ending at
base, the lateral leaflets smaller, slightly unequal at base, obtuse or rounded at
apex, occasionally emarginate,marginrevolute (strongly so towards the base),
leaflet blades chartaceousor subcoriaceous,shining,minutelypuberulouson both
sides when young but soon becoming subglabrous except for base, venation
camptodromous, main veins ? prominent particularlybelow, the midvein im-
pressed near base above. Inflorescencesterminal,erect, widely paniculate,slight-
ly longer than the leaves, 6-10 x 4-10 cm, consisting of a single primarybranch
with alternate side-branchlets, minutely pubescent and occasionally somewhat
hoary; bracts narrowlytriangularto ovate, mostly ca. 2 x 1 mm but sometimes
to 5 x 2 mm, thick, minutely pubescent below with hairs to 0.3 mm long, the
bracts at the base of the lower side-branchletsoccasionally replaced by small
3-foliolate leaves; pedicels 1-3 mm long; bractlets 2, alternate, occasionally sub-
opposite, subtending a secondary pedicel or not. Flowers 7.5-8.5(-9) mm in
diam., fragrant(Runyon3084); calyx lobes imbricate,often quincuncial,overlap-
ping at anthesis or not, free or connate at base, very broadly ovate, 1-2.8 x 1-
2.6 mm, obtuse to rounded or (sub)acute at apex, coriaceous, thick at base,
glabrous above, (sub)glabrousbeneath, parallelodromousto somewhat imper-
fectly actinodromous;petals deciduous, imbricate,often cochlear, partiallyover-
lapping at anthesis, spreading,ovate, 3-4.3 x 1.5-3 mm, obtuse or occasionally
acutish at apex, papery with thickly coriaceous base, white, cream, or yellow-
green when fresh, papillose above, glabrous below except for a few appressed
hairs near base, venation ? parallel to actinodromous, the median nerve often
somewhat thicker; filamentsdeciduous, slightly adnate to the disc at base, nar-
rowly triangular,1.7-3.5 mm long and 0.5-0.7 mm thick at the base, glabrous;
anthers ? sagittate, (1.2-)1.6-1.7 x (0.6-)0.8-1.1 mm, including the projecting
connective 0.2-0.3 mm, the thecae sometimes divergent at base, somewhat pa-
pillose; disc annular, rosette-like, with 10 slight incurvations to 2/5 or V/ of its
thickness, 0.4-0.7 mm high and 0.3-0.7 mm thick, 2.2-2.5 mm in diam., fleshy,
glabrous; ovary 5(-4)-locular, carpels connate and adnate to the disc in their
lower half, ca. 0.4-0.7 mm high, glabrous, at anthesis provided with an apical,
ovoid apophysis to 0.8 x 0.7 mm which protrudes beyond the disc and bears
both a few protuberances0.1-0.2 mm long and also slightly projectingglands;
style attached ? midway on the carpels, thickened towards tip or not, 1.2-2.2
mm long, projecting0.8-1.6 mm beyond the apophyses, 0.2-0.3 mm thick at tip,
glabrous; stigma capitate, 0.2-0.3 x 0.3-0.5 mm. Fruits mostly 1 per infructes-
cence, with orange odor (Runyon 3084), (3-4-)5-locular, depressed, stellately-
42 Flora Neotropica

I' ?7 .,,. I . "' '

/, . '

. .....,..,..???I
::
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. .
"~

.:':':Z:.'.~'.i~.:. . , ...' ....,~......., . ....-..~

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~~~~~~~~~~~~~~.. . ..:.. ... ....
J
.?.~~~~~~~~~~~~~~~~~~~~~~~~rraasi ?-- .?
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'~~~~.1 ~~:i?
C,flower(Runyon2173 W) D fruit(Bartlett10805, F). E, seed(Runyonsn, summer1943, MO).
..??::

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*. X ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~.'i.
/ '..' ! '* .;

,~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~?
~~?i
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.-l~.
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~~~~~~2~~~~~~~~i':
.?? .;. ?,?.?.

C,~~~'
Aw Iuy 27 W D ri Brlf 00 ? F. ,se Rno s.n. umr14,M

FIG.8. Esenbeckiarunyonai A, habit(Runyon2173, W). B, base of leaflets(Runyon2173, W).

Esenbeckia 43

lobed, flattenedat base and convex above, 2-3 x 3-4 cm, glabrous;the loculi ?
obtusely trigonous,provided +?4 (to /2) from base with a dorsal, blunt apophysis
+1-4 mm long with a broadenedridge below the apophysis, convex at the base
around the stalk, the upper ridge convex, slightly but irregularlytuberculate,
dehiscent septicidallyfrom a few mm above base to the axis; seed 1 per loculus,
very broadly obliquely tear-shapedwith flattenedbase, 9.5-12 x 11-12 x 7.5-9
mm, with a distinctly curved beak of ca. 1.0 mm at the apex, testa dull, dark
reddish-brown,finely reticulate-colliculatewith interspaces of ca. 0.10 x 0.02
mm in parallel fascicules which are irregularlyintermingled;chalazal area not
visible; hilum to 1 mm broad;beak provided with a few dull black spots; embryo
1, thick, cotyledons unequal with ears to 0.5 mm, radicle short, projecting0.5-
0.6 mm beyond the ears, plumuleinconspicuous.
Type. Runyon 177, U.S.A. Texas: CameronCounty, 3 mi. NW of Los Fresnos,
bank of Resaca Viejo, 8 Jul 1929, fl (holotype, US-1438940). Stiles s.n. (Herb.
Runyon 177) from same group of trees, 15 Apr 1929,fr (paratype,US-1436973).
Distribution.U.S.A., lower Rio GrandeValley in extreme SE Texas (rare)and
NE Mexico from Nuevo Leon and TamaulipasS to NW Hidalgo. Xeric scrub
and dense brushwood or tropical deciduous forest; on sandy loam on limestone
hills, and on basalt mesa; alt. 10-1800 m. Flowering May-Jul(-Sep); fruitingall
the year round. According to the originaldescriptionfloweringand fruitingtwice
each year. Fig. 6B.
Specimens examined. U.S.A. Texas: Runyon 177 (4 May 1929) st (US), 3084 fl (BM), s.n. (6 Jul
1930) fl (NY); Schiller 668 st (US), 803 st (K).
MEXICO. Nuevo Le6n: Muller 2085 fl (A, F, MICH, MO, NY, P), 2980 fr (F, GH, MICH, UC,
US). Tamaulipas: Bartlett 10696 fr (DS, F, GH, MICH, US), 10805 fr (DS, F, GH, MICH, US);
Berlandier? s.n. (Dec 1831) fr (G); Crutchfield & Johnston 5178 fr (MICH), 5594 fl (MICH); Crutch-
field et al. 6080 fr (MICH, UC); Fisher 3375 fl (F); Manning & Manning 53404 fl (GH); Palmer 477
fl (GH, MO, K, UC); Sullivan 639 fl (NY); Webster et al. 11232 fl (GH, MO, S). San Luis Potosi:
Pringle s.n. (11 Oct 1890) fr (US 2x, VT); Rzedowski 5688 fr (DS 2x, MICH); Virlet d'Aoust 1884 fl
(P). Hidalgo: Moore & Wood 3848 fl (BM, UC, US, WIS). State unknown: Sargent s.n. (Apr 1887)
fr (A).
CULTIVATED. U.S.A. Texas: Runyon 2173 fl (DS, G, K, NY, P, UC, W), s.n. (summer 1943)
fr (K, MO, NY, UC, US).

Local names and use. Mexico (Tamaulipas)huacac (Manning & Manning

53404), limonillo (Berlandier? s.n. (Dec 1831)). Soap is made from the bark (Man-
ning & Manning 53404).
The species was discovered in 1929by Stiles, who sent a specimen to Runyon,
a photographerin Brownsville(Texas). Runyon went several times to the locality
and sent materialto US. Mortonchose Runyon 177 of 8 Jul 1929as the type. All
the other collections, includingStiles's sample, are collected under Runyon 177
and are not isotypes as they were collected at differenttimes.
This species is frequentlyconfused with E. berlandierisubsp. berlandieri.The
fruits are clearly different, but differentialcharacters can also be found in the
inflorescences and flowers, see under E. berlandieri. The abrupt ending of the
leaflet base is a useful characterof E. runyonii;the only known Esenbeckia that
also shows this feature is E. leiocarpa, a Brazilian species which cannot be
confused with the present one.

4. EsenbeckiafeddemaeKaastra, Acta Bot. Neerl. 26: 471, t. 1. 1977. Fig. 9.

Shrub or treelet 1.5 m tall; branchlets5-6 mm in diam., grayish-greenor gray-
ish-brown, pubescent with spreading,hyaline hairs of ca. 0.5-1.2 mm long, gla-
brescent. Leaves alternate,(3-4-)5-foliolate, leaflets stalked or not; petiole semi-
44 Flora Neotropica

~~? . r 1 I ? \? / I ~~~~~~~~~~....;..
:" .

I I. I?Y?I \ ? \ I- ?I"
? ~ ~ ~~ Y?: . .

;?,? ~ ~ ~ ~ ~ ~ ~~ ~ ~ ~ ~ ~ ~ ~ ~ ~

~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~.:: .. ..

~~~~~~~~~~~~~~~~~~~ .

.....

~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~.
II:
.1. ~ t ~ ~ 4. ,:
;.. ???'?:'
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:. ~~~~~~~~~~~?.. . r.? . ? . ..':,.:,.

'z----~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~

~~~~~~~~~~~~~~~~~~~~. ??

? :. :::ji: 5?r . ..
i~~~~~~~~~~~~~~~~~'
~:?
"'

,
,
r:
, ?. i?r
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:: F
;;~~~~~~~~~~~~

?... ? ..?..
1'" '!:1
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;~~~~
~~~~~~~~~~~~.'r 5r~~~~~~~~~~~~~
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'~~~~~~~~? '~? ?~. ~ ~~~~~~~~~~~~~~~ .

??::~~~~~~~~~~~~~~~~~~~~~~'"...'...': '-.'
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FIG.~~~~~~~~~~~~~~~~~~
9. Esnekafdea., ~'
A,hbt(Fd'a269 y MIH. Rw (ed

tye
j69 IH. , fut(mik12 l
Esenbeckia 45

terete, canaliculate, 3-8 cm long, pubescent with white, hyaline, spreadinghairs

(0.3-)0.4-0.6 mm long; petiolules not sharply separated from the leaflet bases,
0-10 mm; leaflet blades elliptic or slightly obovate, 7.5-18.5 x 4.3-7.7 cm, lat-
eral leaflets smaller: the proximal blades generally 2.5-6 cm long, base (long-)
attenuateand graduallypassing into the petiolule, lateralleaflets unequalat base,
apex obtuse, rounded or slightly acuminateand frequently somewhat recurved,
marginstrongly revolute, leaflet blades subcoriaceous, dull green, pubescent on
both sides like the petiole, soft to the touch particularlybelow, venation camp-
todromous, midvein plane above. Inflorescences much shorter than the leaves,
to 5 x 6 cm, densely crowded with flowers, consisting of 2-3 terminal, erect,
paniculately-branched,thick main branchletswith alternate side-branchlets,pu-
bescent like the leaves; bracts very broadly ovate ca. 2-3 x 3 mm, with slightly
auriculatebase, pubescent on both sides like the leaves but the upper side some-
what shorter and more sparse; pedicels 1-2 mm long; bractlets 2, subopposite.
Flowers 9-9.5 mm in diam.; calyx lobes quincuncial, depressedly ovate, 1-
1.5 x 1.9-2.4 mm, roundedat apex, coriaceous with transparent-paperymargin,
glabrous,venation parallel;petals quincuncial,very widely spreading,ovate, 3.8-
4 x 2.5-2.6 mm, rounded at tip, transparent-papery,creamish both when fresh
and when dried, glabrous, venation actinodromous;filamentspersistent, slightly
adnate to the petals at very base, subulate, 3.3 mm long and 0.7 mm thick,
glabrous; anthers heart-shaped, 1.4-1.6 x 1.2 mm including a mucro 0.1 mm;
disc rosette-like, slightly 10-lobed, 0.8 mm high, 0.6 mm thick, and 2.7 mm in
diam., thick, brownish, glabrous;carpels adnate to the disc and connate only at
base, 0.6 mm high, purple, glabrous, in older flowers each developing a purple
apophysis ca. 0.6 x 0.4 mm densely beset with glands and glandulartubercles to
0.3 mm long; style insertedjust below the tips of the carpels, 1.5 mm long and
0.4 mm thick, free for 1.4 mm; stigma capitate, 0.4 x 0.6 mm. Fruits de-
pressed, stellately 5-lobed, truncateat tip, 20-25 x 25-35 mm when closed, loculi
provided with a dorso-apical, erect horn 10-13 mm long, carinate, slightly wrin-
kled and obsoletely muricate with few prickles 1-2 mm long, the nerves of the
exocarp externally visible, septicidally dehiscent from ca. 5 mm above the base
to the axis; seeds unknown.
Type. Feddema 2699, Mexico. Michoacan: Rancho Chila, ca. 20 km SE of
Coahuayana, and ca. 5 km NW of Aquila, 23 Nov 1963, fl (holotype, MICH;
isotype, MICH).
Distribution. Mexico, Michoacan. Not common in tropical deciduous forest
together with Lysiloma, Lonchocarpus, Bursera, and Apoplanesia. Flowering
and fruitingNov. Fig. 6B.
Specimenexamined. MEXICO.Michoacan:Maguili:Emrick122 fr (F).
This species resembles E. berlandieri. Differences are to be found in the in-
dument of branchlets and leaflets, the hairs being longer and spreading. Other
differences are the shorter inflorescence, the larger flowers, and the massive,
purple apophyses of the ovary.
5. Esenbeckiacollina Brandegee, Univ. Calif. Publ. Bot. 6: 183. 1915;Standley,
Contr. U.S. Natl. Herb. 23: 536. 1923.
Shrub to 3 m tall; branchlets3-6 mm in diam., light grayish-brown,becoming
darker, strongly longitudinallywrinkled, the young branchlets hoary with ap-
pressed whitish hairs 0.1-0.3 mm long, becoming glabrous. Leaves 3-foliolate,
those in the inflorescences 1-foliolate,the leaflets petiolulate; petiole semiterete,
46 Flora Neotropica

adaxially flattenedor slightly canaliculate,slightly thickened at base or not, 1.4-

4.5 cm or shorter, + densely pubescent with hairs to 0.2 mm long, terminal
tubercle present or not; petiolules winged by the decurrentbases of the leaflets,
4-10 mm long, those of lateralleaflets 1-6 mm; leaflet blades (slightly narrowly)
elliptic to (slightly narrowly)obovate, 3.5-10 x 2-4.5(-6) cm, attenuate at base
and equal or not, obtuse and emarginate,or roundedat apex, marginsubrevolute,
chartaceous, dull grayish-green,paler beneath, venation camptodromous,prom-
inent on both sides, midveinplane or prominulentabove. Inflorescencesno longer
than the leaves, hoary or pubescent with hairs 0.1-0.2 mm long; side-branchlets
and pedicels alternateor occasionally subopposite;bractsvery broadlytriangular,
indumentlike that of the branchlets, or subglabrous;bractlets 1 or 2, alternate.
Flowers 8.5-11 mm in diam.; calyx lobes quincuncial,roundedat apex, varying
in size in one flower to 1.5 x 2.5 mm, coriaceous but the marginspapery and
lighterin color thanthe rest of the lobes, glabrouson both sides, venationparallel;
petals persistent, cochlear or quincuncial,very widely spreading,elliptic, obtuse
or rounded, coriaceous with papery margins, purplish-brownwith light yellow-
brown marginswhen dried, glabrouson both sides, venation actinodromous,the
mediannerve thickerand forminga distinctrib on the back of the petals; filaments
persistent, subulate, triangularin cross-section, flattened especially towards the
widened base, the adaxial side with a median, obtuse angle, 3.2-3.5 mm long and
0.6-0.9 mm broadat base, fleshy, swollen, purplish-brownwhen dried, glabrous;
anthers narrowly heart-shaped, somewhat papillose particularlytowards apex;
disc annular, 10-lobed,0.3-0.8 mm high, 0.2-0.5 mm thick at tip, 2.3-2.6 mm in
diam., fleshy, purplish,glabrous;carpels adnate to the disc, hardly immersedin
the receptacle, 0.4-0.7 mm high, provided with a dorso-apical, irregular,solid,
light brown-purplish,glabrous apophysis; style persistent, inserted near base of
the carpels, clavate or terete, ? obtuse-angled,purplish-brownwhen dried, gla-
brous; stigma clavate or capitate, 5-lobed, 0.3-0.4 x 0.5-0.8 mm, developing a
large yellow gland in each lobe. Fruits 2-3 per infructescence, depressed, stel-
lately-lobed due to 5 dorsal horn-likeapophyses 3-4 mm long inserted 1/, below
tip of the dehisced loculi, 1.2-1.5 x 2.4-2.8 cm, to 2 cm broad when not yet
dehisced, nearlyflat at apex when still closed, the apophyses later somewhatbent
downwardsand outwardsby dehiscence of the loculi, densely glandular-punctate,
glabrous;loculi roundedat base, the nerves of the exocarp externallyvisible, the
exocarp wrinkledand weakly muricate; seed I per loculus observed, obliquely
tear-shaped, 10.3 x 5.7 mm, the oblique base slightlyflattened,long-beaked,tes-
ta darkreddish-brown,linear-reticulate-colliculate, the interspacesto 0.20 x 0.02
mm, chalazal area not visible, hilum to 0.6 mm broad; embryo 1, cotyledons
unequal, radicle projecting0.3 mm beyond the ears which are 1 mm long.

Key to the Subspecies of Esenbeckia collina

1. Leaflets above appressed-puberulous;
flowers 8.5-9 mm in diam. 5a. subsp. collina.
1. Leaflets above spreading-puberulous;
flowers 10.5-11 mm in diam. Sb. subsp. conspecta.

5a. Esenbeckiacollina Brandegee subsp. collina. Fig. 10.

Low compact shrub90-120 cm; branchletsangled, strongly cracked when old,
knotted, crooked. Leaves alternate,crowded at tips of branchlets;lateralleaflets
somewhat smaller than the terminal one, appressed-puberulouson both sides
with hairs to ca. 0.2 mm long, the midvein more densely hairy, especially above.
Inflorescences terminal, 4-12 x 1-2 cm, consisting of 1 to few, erect, narrowly
paniculate, often racemiformpartialinflorescences;pedicels 5-7 mm long. Flow-
Esenbeckia 47

'.

FIG. 10. Esenbeckia collina subsp. collina. A, habit (Purpus 7140, type, GH). B, flower (Purpus
7140, type, GH). C, fruit (Fisher 35497, MO). D, dehisced fruit (Fisher 35497, NY). E, seed (Fisher
35497, NY).
48 Flora Neotropica

ers 8.5-9 mm in diam.; calyx lobes heart-shaped, their bases distinctly eared
except in the smaller lobes; petals 4 x 2.5-2.8 mm; anthers 1 x 0.8 x 0.4 mm;
apophyses of the ovary ca. 0.9 x 0.7 mm, smallerin bud, chargedwith tubercles
and projectingglands; style 2 x 0.5 mm, the free part 1.6 mm long.
Type. Purpus 7140, Mexico. Oaxaca:Cerrodel Picacho, Jul 1914,fl (holotype,
UC-173396Herb. Brandegee;isotypes, BM, F, GH, MO, NY, US).
Distribution.Rare in Oaxaca, Mexico. Once reportedfrom 90 m alt. Collected
in flower Jul-Aug. Fig. 6C.
Specimenexamined.MEXICO.Oaxaca:Ixtepec, Fisher 35497 fl, fr (F, MO, NY).

5b. EsenbeckiacollinaBrandegeesubsp. conspectaKaastra,Acta Bot. Neerl. 26:

473. 1977. Fig. 11.
Slender shrub 3 m tall; branchletsterete. Leaves alternateor subopposite, the
petioles ridged, the lateral leaflets smaller than the terminalone; leaflets puber-
ulous on both sides with hairs 0.3 mm; these ? spreadingand slightly curved
above, appressed below. The leafy, terminalinflorescences composed of several
erect or spreadingpanicles 6-7 x 4 cm leafy at base; pedicels 3-3.5 mm long.
Flowers 10.5-11 mm in diam., scentless; calyx lobes very broadlyor depressedly
ovate; petals 4.8-5 x 2.5 mm; anthers in outline 2 x 1.3-1.5 mm, including a
mucronatetip 0.1 mm; apophyses of the carpels 1.2-1.3 x 0.8 mm, reachingthis
size already in bud, provided with few tubercles 0.1-0.2 mm long; style 3.2 mm
long and 0.3-0.4 mm thick, the free part 2.3 mm long. Fruits unknown.
Type. Turner2086, Mexico. Michoacan:2 mi. NW of La Placita, 45 air mi. S
of Colima 5 Jul 1950, fl (holotype, MICH).
Distribution.Known only from the type locality. On red silty-clay soil among
limestone boulders on hillside. Fig. 6C.
This subspecies differsfrom subsp. collina in the less compact growth with the
leaves not crowded at the tips of the branchletsand in the more prominentve-
nation of the leaflets. Differences are also found in the indumentand in the size
of the flowers, which seem to be darkerviolet. Finally, the apophyses are larger
and smoother.
The subspecies differs from E. warscewiczii in the filaments, the prominent
venation of the leaves, the glabrous perianth, and the longer apophyses, style,
and anthers.

6. EsenbeckiairwinianaKaastra, Acta Bot. Neerl. 26: 473, t. 2. 1977. Fig. 12.

Few-branchedshrub, or tree, 1-4 m tall; branchletsdark grayish- to reddish-
brown, pubescent with spreadinghairs0.2 mm, becomingglabrousin age. Leaves
alternate, 3-foliolate, with stalked leaflets; petiole semiterete, canaliculate, 2-6
cm long, densely pubescent or somewhat velvety with spreading hairs 0.1-0.3
mm long, tubercle present to nearly obsolete; petiolule of the terminalleaflet 5--
11 mm long, those of the lateral ones 1-5 mm; leaflet blades obovate or elliptic,
4-11 x 2.2-5 cm or the lateral leaflets smaller, rounded, obtuse, or very shortly
attenuateat base, the lateral ones unequalat base, emarginateor obtuse at apex,
marginthick and revolute towards base, chartaceous, green and somewhat shin-
ing above, dull and pale beneath, pubescent above with spreadinghairs 0.1-0.2
mm long, the midvein densely so, pubescent to tomentose below with curved,
soft hairs to 0.6-0.7 mm long, the midvein densely so, venation camptodromous,
prominenton both sides, the midvein impressed above. Inflorescences terminal,
erect, paniculate, 10-25 x 10-25 cm, many- but rather loosely-flowered, made
Esenbeckia 49

i
.

A
FIG. 11. Esenbeckia collina subsp. conspecta (Turner 2086, type, MICH). A, habit. B, leaf ve-
nation.
50 Flora Neotropica

.
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~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~..
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?:. ~~~~~~~~~~C:] ?:'.? . ' '. ?..i? .l ..-a ? :. .' .Z
'

i? ? ? .-../.
i.~~
.' ' : ~ ~
1?

~? .
.. -~ ... .?
'
,.'
.~~~~~~~~~~~~~~~~~~~~'
.,;:?; :? :

1..
.. .~~~~~~~~~~~ . ''

-?'"-Ti~~~~~~~~~~~~~~~~,;
?
,.......'.":.'..:.'.
..
.,

'
I-.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
'

'.; .;~~~~~~~~~~~
.
'::.??,:, . ' ':,?
q? "
:":.:?;"...?
''', .???::r.:?:?i~3... ?
.'?'..

i~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~.'
.....

FI, ... i~..f~. irinan. .,f..t ,35,... Iwn ta.: :ye . B, .lwe (ri a..

23451 typ, NY. C,frui (Anersonet a. 880, U.ves:.~. D,seedshowig : adacen. . (A

derson~~~~~~~~~~~~~~~~~~~~~~~~
et. U) .'..

derson et al. 8850, U).

Esenbeckia 51

up of 1-2 main branchlets 10-25 cm bearingfew, alternate, secondary branchlets

5-10 cm long, densely pubescent with spreadinghairs 0.05-0.2 mm long; branch-
lets of higher order mostly falling off immediately after anthesis like the bracts
and bractlets; bracts triangular,+0.7-1 x 0.4-0.5 mm, thickly coriaceous, hair-
cover on the back like that of the branchlets; pedicels 2-3 mm long; bractlets
alternate. Flowers 8.5-9 mm in diam.; calyx lobes quincuncial,broadly ovate to
subcircular, ca. 1 x 1 mm, obtuse, very thickly coriaceous, pubescent on the
back with spreadinghairs 0.1 mm long, nerves parallel, false midribobsolete or
absent; petals persistent, cochlear or quincuncial,very widely spreading,later on
reflexed, elliptic or slightly obovate, 4-4.2 x 2-2.1 mm, obtuse or rounded at
apex, + transparent-papery,reportedly white, sulphur-yellowishwhen dried,
minutely velvety-papillose above, pubescent below with spreadinghairs 0.1-0.2
mm long; venation actinodromouswith the median nerve slightly thicker; fila-
ments persistent, subulate with flattenedbase, ca. 4 mm long and 0.4 mm thick,
glabrous;anthers heart-shaped,0.8 x 0.6 mm includinga mucro 0.1 mm, finely
brownish-papillosewhen dried; disc cup-shaped, slightly lobed, 0.5 mm high,
0.2-0.3 mm thick, and 1.5-2 mm in diam., as high as the ovary including its
tubercles, fleshy, glabrous; carpels connate with each other and adnate to the
disc, ca. 0.4 mm high, with tubercles0.3 mm, glabrous;in age developingyellow-
brown tubercles in the lower part and few long, dark (purplish?)ones at tip,
becoming hoary with extremely short, spreadinghairs; style inserted near the
base of the carpels, ca. 3-5 mm long and 0.2-0.3 mm thick, free for 2.8 mm,
glabrous; stigma capitate, 0.1-0.2 x 0.4 mm. Fruits ca. 4 per infructescence,
depressedly globose, stellately 5-lobed, 1.5-2 x 2.5(-3.5 when dehisced) cm,
hoary; loculi irregularlymuricatewith prickles to 3 mm, dorso-apicallyprovided
with a slightly furrowedapophysis 4-6 mm long which points upwardswhen the
fruit is still closed, septicidallydehiscing from ca. 5 mm above the base to 2 mm
below tip; seeds tear-shaped,slightly flattenednear base, 8 x 4.8 x 3.8 mm, the
apex with a sortly curved beak 0.5 mm, the testa mediumbrown, finely reticulate-
colliculate with interspaces 0.05-0.10 mm; chalazal area not seen; hilum to 0.7
mm broad; embryo 1 per seed, with unequal cotyledons, punctate, radicle pro-
jecting 0.6 mm beyond the ears, plumule0.1 mm long.
Type. Irwin et al. 23415, Brazil. Minas Gerais: Serrado Espinhago,N of Grao
Mogol, 17 Feb 1969, fl (holotype, NY; isotype, U).
Distribution. Serra do Espinhaqo, Minas Gerais, Brazil. Reported as locally
common in sandy cerrado and open woods on rocky slopes; one collection from
near a stream;alt. 950-1150 m. Flowering in Feb. Fig. 15A.
Specimen examined. BRAZIL. Minas Gerais: Serra do Espinhaco, 9 km by road SW of Mendanha,
Anderson et al. 8850 fr (NY n.v., U).

This species is close to E. pumila. Vegetativelyit is distinguishedby the slender

petioles and the stalked leaflets.
According to the label of the type the stamens should be violet. The upper
tubercles of the ovary and possibly also the papillae of the anthers might be
violet.

7. Esenbeckia pumila Pohl, P1. Bras. 2: 44, t. 128. 1830; Martius, Nov. gen. sp.
pl. 3: 84. 1831 (as "E. pusilla"); Dietrich, Syn. pl. 1: 831. 1839; Engler
in Martius, Fl. bras. 12(2): 143. 1874. Fig. 13.
Esenbeckia latifolia Martius, Nov. gen. sp. pl. 3: 84, t. 234. 1831, syn. nov.; Dietrich, Syn. pl.
1: 831. 1839; Walpers, Repert. 1: 501. 1842. TYPE. Martius s.n., Brazil. Minas Gerais:
Nr. boundary with Goias, Chapada do Parana, Sep 1818, fl but now st (holotype, M).
52 Flora Neotropica

:, . 'e
*9 :

..6'
.c...c n

FIG. 13. Esenbeckia pumila. A, habit with analyses (type, Poh, P. Bras. 2: t. 128. 1830). B, fruit
(Liitzelburg19121295,M).
Esenbeckia 53

Colythrumpumilum(Pohl) Schott, Rutac. 14. 1834.

Colythrumlatifolium(Martius)Schott, Rutac. 14. 1834.
ColythrumpuberulumSchott, Rutac. 13, t. 7. 1834;Englerin Martius,Fl. bras. 12(2):143. 1874,
pro syn. TYPE. Schott, Rutac. t. 7. 1834 (lectotype).
Esenbeckia choisyoides Schlechtendal,Linnaea 27: 529. 1855("1854"), syn. nov. TYPE. Un-
known.
EsenbeckialeucophyllaTurczaninow,Bull. Soc. Imp. NaturalistesMoscou31(1):440. 1858,syn.
nov.; Miiller in Walpers, Ann. 7: 529. 1869. TYPE. Blanchet 2779, Brazil. Bahia: Serra
Jacobina,(1838?)fl (holotype,if any, n.v.; isotypes (with localities SerraA9uruaand/orRio
S. Francisco), BM, BR-Herb. F6e, BR-Herb. Martius,E-Herb. Arnott, F ex G, G-Herb.
Moricand2x, GH ex Herb. Bentham,K-Herb. Hooker, photo NS 2836 made by NY, NY;
LE, NY ex W, OXF, P, P-Herb.Drake, P-Herb.Drakeex Herb. Franqueville,W).
Esenbeckiapumila Pohl var. genuina Englerin Martius,Fl. bras. 12(2):143. 1874,nom. invalid.
ex Art. 24, ICBN.
Esenbeckiapumila Pohl var. latifolia (Martius)Englerin Martius,Fl. bras. 12(2):144. 1874,syn.
nov.
Esenbeckiapumila Pohl var. leucophylla(Turczaninow)Englerin Martius,Fl. bras. 12(2): 144.
1874, syn. nov.
Subshrubwith clumps of several branches inserted on a woody base, 25-100
cm tall, sometimesarborescentto 3 m; branchlets3-5 mm in diam., brownturning
red, + tomentose when young with grayish hairs to 0.5 mm long. Leaves alter-
nate, 3-foliolate except for the uppermost, occasionally some 2- or 5-foliolate,
towardsand in the inflorescencegraduallybeing replacedby 1-foliolateand simple
leaves, with usually sessile leaflets; petiole semiterete, (slightly) canaliculate,
obsoletely winged or ribbed, 1-4(-7.3) cm long and to 3 mm thick, the wings to
0.3(-1) mm broad, usually without auricles, indumentlike that of the branchlets
or less dense; leaflet blades elliptic or (narrowly) obovate, those in the inflo-
rescence narrowlyoblong or narrowlyelliptic, 6-14.5(-17) x 2-7 cm, the lateral
leaflets and the leaflets of the uppermostleaves smaller, usually (narrowly)cu-
neate at base, sometimes obtuse or somewhat shortly attenuateto 3 mm, slightly
unequalat base, at apex roundedor obtuse and slightly acuminatewith emargin-
ate tip, occasionally subacute, marginthick and + revolute, blades chartaceous
or subcoriaceous, dull, paler beneath than above, subglabrousto pilose above
with hairs to 0.5 mm long, minutely pubescent to tomentose below with hairs
0.5-1 mm long, venation camptodromous,prominentparticularlybelow, midvein
somewhat impressed above. Inflorescences terminal, many-flowered,consisting
of 1 main branch and few alternate secondary branchlets subtended by a 3-fo-
liolate leaf or, towards the tip, an 1-foliolateor simple leaf or a bract, ? densely
tomentose with hairs to 0.5 mm long, primaryand secondary branchlets erect,
paniculately-branched, 5-13 x 2-12 cm, with alternate or subopposite side-
branchlets, bracts (broadly)ovate, ca. 1-3 x 0.7-2 mm, or longer and narrower
when intermediatebetween 1-foliolateleaf and bract, the indumentlike that of
the branchletsbut often less dense, provided with a false glandularmidrib;ped-
icels 1-4 mm long; bractlets 2, alternate or subopposite. Flowers 6-7.5(-9.5)
mm in diam.; calyx lobes quincuncial, broadly or depressedly ovate, 1-1.3
(-2.2) x 1.3-1.6(-2) mm, rounded at apex, coriaceous, often brown-pigmented
with yellow marginswhen dried, minutelypubescent below, parallel-nerved,with
a false midrib;petals cochlear or quincuncial,adnateto the disc, spreading,ovate
or elliptic, 3-3.5(-4) x 1.9-2(-2.6) mm, rounded or obtuse at apex, papery or
rarelycoriaceous with paperymargins,often brown-pigmentedwhen driedexcept
for the yellow margins,creamishwhen fresh, papilloseabove, sparsely or densely
minutelypubescent below with hairs 0.05-0.1 mm long, venation actinodromous,
the nerves branched towards the margin;filaments subulate, flattened towards
the base, 1.8-2.5 mm long and 0.4-0.6 mm thick, glabrous;anthersheart-shaped,
0.9-1.2 x 0.7-0.9 mm includinga tip 0.05-0.1 mm, becoming pigmentedon the
54 Flora Neotropica

back and at tip, papillose; disc cup-shaped, with 5(-10) slight incurvations,0.5-
0.8 mm high, 0.2-0.4 mm thick, ca. 1.5-2.3(-2.8) mm in diam., coriaceous, gla-
brous; carpels adnate to the disc in the lower part, 0.5-0.7 mm high, densely
chargedwith conical or finger-likeglandularprotuberances0.5 mmlong, glabrous;
style inserted 1/3 to 12 from base of the carpels, 1.4-1.6 mm long and 0.1-0.4 mm
thick, projecting ca. 0.7-1 mm beyond the ovary including its protuberances,
glabrous;stigmacapitate, 0.2-0.3 x 0.2-0.4 mm. Fruits depressed, stellately-glo-
bose with flattened top when still closed, 1.5-2.5 x 2-3 cm(-4 cm when de-
hisced), sparsely minutelypuberulouswith hairs 0.05-0.1 mm long; loculi dorso-
apically trigonous, muricatewith prickles and tubercles to 4 mm long, wrinkled,
dehiscing septicidallyfrom up to 5 mm above the base and to 2 mm from the axis,
the exocarp rather thick; seed obliquely tear-shaped, 10.5-12 x 5.7-6 x 5 mm,
flattenednear the base, blunt and obsoletely curved or straightat the apex, testa
0.4 mm thick, chestnut with shininggranules, granulate-pusticulate,the granules
ca. 0.05 mm long; chalazal area not seen; hilum 1.5 mm broad; embryo 1, apex
acute, cotyledons unequal, radicle ca. 1 mm long and projecting0.5 mm beyond
the ears.
Type. Pohl 750 delta = 739 Herb. Bras., Brazil. Goias: Serra dos Crystaes,
Nov-Dec 1819, fl (holotype, W, the lowest specimen on the sheet, from which
the drawingin Pohl was made; isotypes, K-Herb. Hooker, photo NS 2862 made
by NY, NY; K-Herb. Benthamex W).
Distribution.Brazil: Piaui, Mato Grosso, Goias, Distrito Federal, Bahia, and
Minas Gerais. Chieflyin cerradoand grassy campos but also in forest and gallery
margins;alt. to 1000m. Commonin some areas, otherwise infrequent.Flowering
Sep-Dec. Fig. 15A.
Specimens examined. BRAZIL. Piaui: Lutzelburg1912/295fr (M). Mato Grosso: Prance et al.
18946 fr (U). Mato Grosso or Goias: Weddell 2967 fl (P 3x). Goias: Anderson et al. 7924 fl (U),
Burchell 6190 fl (K, P); Duarte 9423 fl (NY); Gardner 3028, Oct 1839, fl (K), 1841, fl (K), 3828 fl
(CGE); Glaziou 20798 bud (BR, C, G, K, LE, MPU, P 2x, R, S); Irwin et al. 10788 fl (U), 10826a fr
(U), 12459a fr (U), 14400 fr (U), 34444 fr (U); Riedel 2511 fl (K, LE 2x); Smith & Macedo (in Macedo)
4652 fl (S). Distrito Federal: Castellanos 21831 fl (R); Heringer 8742 fl (NY, SP); Irwin & Soderstrom
5993 fr (NY); Irwin et al. 8236 fl (U), 9710 fl (U), 12175 fr (U), 26481 fr (U), 26647 fr (U); Pereira
4752 = Pabst 5077 fl (F, M), 9003 fr (LP). Bahia: Blanchet 2780 fl (G, photo MO, NY), s.n. (Rio San
Francisco) fl (P); Liitzelburg 1912/284 fr (M). Minas Gerais: Anderson et al. 9041 fr (U); Belem 1929
fl (NY); Belem & Barroso 4002 fl (NY). State unknown: Pohl s.n. (1839 ex W) fl (BR).

Uses. The bark possibly used like that of Esenbeckiafebrifuga (see PHYTO-
CHEMISTRY).
This species is distinguishedby its flat and broad petioles.
Through specimens received from NY (Irwin et al., Anderson et al.) and col-
lections of Harley et al. (K) several fruits came to my disposal;fruitswere seldom
collected before. Ripe seeds however, are still very rare (Lutzelburg1912/284).
By studying these collections it became clear that var. leucophylla (=Colythrum
puberulum Schott) is untenable. Several collections show a gradient from the
typical tomentose indumentof var. leucophylla to the subglabroushair-coverof
var. pumila: e.g. Irwin et al. 14400, 12175; Pereira & Pabst 4752; Smith &
Macedo (in Macedo) 4652.
Esenbeckiachoisyoides is probablynot different.The ample descriptionagrees
fully with tomentose specimens of E. pumila. No type is known, either at HAL
or elsewhere.
The type of E. latifolia was indicated as from "in confiniis Prov. Minarum et
Goyaz, in campis editis versus Vdo do Parandn, Sept." The specimen at M
collected by Martius in: "M. G. et Goy. Chapada do Parandn [=Parana] Sept.
Esenbeckia 55

in campis elevatis," is the type of E. latifolia from which the drawing in the
protologue was apparently made. Unfortunately the inflorescence has disap-
peared. Thus no conclusions can be drawn about the size of the flowers which
should be smaller, according to Martius, than those of E. pumila. Some small
differences in the vegetative parts are of minorimportance.Therefore,I decided
to reduce E. latifolia to synonymy of E. pumila. Engler made a variety of it and
added the diagnostic character "foliis atque ramulis floriferis omnino glabris."
The leaves however, are not fully glabrous:there are some appressedhairs pres-
ent as in E. pumila. Engler did not see the type of E. latifolia, but only used the
originaldescription with drawing.
Glaziou 20798 has some 5-foliolateleaves.
8. Esenbeckianesiotica Standley in Ferris, Contr. Dudley Herb. 1: 73, t. 1, fig.
6. 1927;Engler in Engler and Prantl, Nat. Pflanzenf. (ed. 2) 19a: 281.
1931. Fig. 14.
Usually tall tree with smooth, greenish-brownand mottled bark;branchletsca.
5 mm in diam., brownish-gray,densely appressed-pubescentwhen young with
grayish hairs 0.2-0.4 mm long but soon becoming glabrous. Leaves alternateor
subopposite, crowded at tips of the branchlets, 1-foliolate with stalked leaflet;
petiole terete, slender, 14-22 mm long, minutely pubescent with mostly ap-
pressed, white-hyalinehairs 0.1-0.2 mm long, but densely strigillosetowards the
leaf-joint with hairs to 0.6 mm long; petiolule ca. 0.7-1 mm long; leaflet blade
elliptic or slightly obovate, 5.5-11 x 3-5.5 cm, broadlycuneate or obtuse at base
and at extreme base very shortly attenuate and often slightly unequal, at apex
rounded and slightly emarginate,the marginplane, the blade chartaceous, dull
green, glabrous or nearly so on both sides except for the minutely pubescent
midvein, or appressed-pubescentat base with hairs 0.1-0.2 mm long, venation
camptodromous,prominenton both sides, midveinplane above. Infructescences
terminal,erect, paniculate,as long as the leaves, to 9 x 5 cm, minutelypubescent
with hairs 0.1-0.2 mm long; side-branchletsalternate;bracts (broadly)ovate, to
ca. 1 x 0.8 mm, minutelypubescent below; fruit stalks ca. 12 mm long; bractlets
1-3, alternate. Flowers at anthesis unknown (all data taken from young fruiting
stages); calyx lobes very broadly or depressedly ovate, ca. I x 1.3-1.5 mm,
minutelypubescent with appressed hairs 0.1-0.2 mm long; petals persistent, ca.
4 mm long, transparent-papery,probablyglabrous;filamentspersistent. Fruits 2-
6 per infructescence, depressed, stellately-lobed, 18-20 x 28-35 mm both closed
and dehisced, with glands, glabrous;loculi somewhat blunt-angleddorsally half-
way, densely beset with irregular,blunt densely foveolate and irregularlywrin-
kled tubercles to 2.5 mm long, dehiscent septicidally from the lower half to the
axis, and loculicidally to ca. /3 from below tip; seed (only 1 seen of which only
the testa remained)probably 1 per loculus, obliquely subglobose with rounded
apex, ca. 9 x 8 x 8 mm, testa shiningdarkbrown, reticulate-colliculatewith the
interspaces linear 0.2 mm long, or rounded 0.05 mm in diam.; chalazal area not
visible; hilum 3.5 mm broad; embryo unknown.
Type. Ferris 5699, Mexico. Nayarit: Las Tres Marias, Isla Maria Madre,
west side, 24 Oct 1925, fr (holotype, DS-145714; isotypes, US-1265963, US-
1265994).
Distribution.Known only from the type locality, where it is reportedly com-
mon.
The leaves are undoubtedly 1-foliolatebut this is somewhat difficultto observe
in the present specimens.
56 Flora Neotropica

: , :. . '. .' :
..~~~~~~~~~~: < ?..
t::11i'..:
..
... .. . :-
.;. ..::'. ;; '
, : 0':- ,:::.': $,t;,.,0f ......f: ,, ff :: 0 }t,7;.0 , 0 .' ?

~~ ... i:
:~~..
FIG.?14. :.... ."...,,ic: ?
?zB*r
! i;i
Esenbeckia ;:?i i:ii
A, habit (Ferris5699,type, DS).
nesiotica.
~
.
~
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
::~~~~~~~~~~~~~~~~~~~~~ ',
. . . ? ' . ?

:: i: ~ ~ ~~~ ~ :i~. ~~'

~i
~..,r?' :::j

'~~~~~~~~~~. "'?1~ A"

'
~~~: ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~":
?
::?. '~ '

~~~
!i % :
?; : ..

~~: ??
%.~ ~ ~ :?y..~?:;
~ ~~~~~~i :'

;
' .i. ':?,.
?~:
.

z ;~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
? ~ ~~ ~ ~E..'~;;~~ ~ '
?.:' ?? ..I
I ?:?F: 'i .,...

,~ ~ ~ ~ ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~? ,.

?
?r??~~~~~~~~~~~~~~~~~
.:' .: ..1?
' ,...? .
~ ~ ~~~~~~~~~~~
,. ? /

~~';
' ' ?? R '???: :???~~~~~~
, I .... ..i ?, .;':. .
?;? .' ?:
..I~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
? .

\ ~~~~~~
,.?.~
;. .!
'.:
? ~ ~ ?I: 1....::
? :i;?'
: : : i?'.~:
i:
'i :-i:~;::::{
i
'

? '
'' ?:~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
.:
.?
?.
.'.: ?
:
~~~~~~~~~~~~~~~~~~~~~~~~~~~
?
.':..:. : . . .
..
~..Z
. ,

?~~ i ... ..?

!i::::!?:.i,;cI '~
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~!
FIG. 14. Esenbeckia
nesiotica. A, habit (Ferris
5699, type,DS).~~~~~~~~~~
Esenbeckia 57

9. EsenbeckiagrandifloraMartius, Nov. gen. sp. pl. 3: 85. 1831;Dietrich, Syn.

pl. 1: 831. 1839; Martius, Flora 24(2). Beibl. 2: 29. 1841; Engler in
Martius, Fl. bras. 12(2): 146. 1874;Macbride, Field Mus. Nat. Hist.,
Bot. Ser. 13: 672. 1949; Cowan, Sellowia 12: 88, t. 3, fig. a-f. 1960;
Cowan and Smith in Reitz, Fl. Ilustr. Cat. 1 (RUTA) 33, t. 8, fig. a-f,
and t. 9. 1973.
Shrub or small tree to 7(-10) m tall with trunk3.5-8 cm diam. but to 20 cm in
Venezuela; barkfull of cracks, indumentof ferrugineous-hyalinehairs to 0.3 mm
long; branchlets 2-5 mm diam., dark reddish-brown,later on grayish, glabrate
but younger branchlets (dispersedly) strigillose. Leaves alternate or some
(sub)opposite, often crowded at tips of branchlets, 1-foliolate,the leaflet sessile;
petiole terete, slightly canaliculatewhen mature,becomingtransverselycracked,
with a turgidjoint just proximallyof the leaflet, glabrousor minutelypuberulous,
leaflet-blade(narrowly)obovate to (narrowly)elliptic, at base narrowlycuneate
to attenuate, apex obtuse, shortly subacuminateor somewhat roundishor rarely
rounded, margin(sub)revolute, ? rigid-coriaceous,dull or occasionally shining,
glabrous above, sparsely minutely puberulous beneath when young, venation
brochidodromous,main veins prominent beneath, midvein proximally slightly
impressed above. Inflorescences terminal,or axillaryat tips of branchlets,erect,
(narrowly)paniculate, usually few-flowered, densely, or rarely sparsely, strigil-
lose; side-branchletsall or only those of higher order opposite, shorter than the
leaves; bracts and bractlets alternateor opposite, (broadly)triangular,concave-
conduplicate, coriaceous, papillose towards the base above, strigillose below,
with a prominentfalse midriband parallelnerves; pedicels 1.5-5 mm long. Flow-
ers 7.5-14 mm in diam., calyx lobes quincuncial,very broadly ovate to subcir-
cular, obtuse or rounded, very thickly coriaceous, glabrous above, the nerves
externally not visible, parallel and slightly branched towards the tip, provided
with a median false midrib on the back; petals mostly deciduous, contorted to
the right or rarely cochlear, very widely spreading, ovate to elliptic, obtuse,
nerves externally not visible, subparalleland slightly branchingtowards the tip,
the median nerve somewhat thicker, usually without false midrib;filamentssub-
ulate, somewhat complanate, 2-3.5 mm long, some glands present, dorsally near
base with some hairs to 0.1 mm long, or glabrous; anthers droppingoff early,
heart-shaped, 0.7-1.5 x 0.6-1.2 mm, including a mucronate tip 0.1 mm; disc
annular,forming a ? distinct pentangle, the angles sulcate, the sides obsoletely
or scarcely lobed, 0.7-1.5 mm high and ca. 0.5 mm thick above, 2.5-4 mm in
diam., fleshy, swollen towards the tip, glabrous, sometimes with granules, rarely
somewhat pilose; carpels depressedly globose, 5-lobed with shallow broad
grooves, completely connate and often with their lower part immersed in the
receptacle, glabrous, very densely provided with clavate to finger-like, blunt-
pointed protuberanceswhich are finely granulateand glandularat anthesis, 1 or
2 of the longest protuberances standing midway on the carpels; style inserted
midway on the carpelsjust above an elevated part of the receptacle, to 2 mm long
and 0.4-0.5 mm thick, with hyalinehairs nearbase to 0.3 mmlong; stigmacapitate
or clavate, ? distinctly 5-lobed, 0.5-0.8 x 0.6-1 mm, with 5 large glands which
sometimes later disappearleaving the stigma 5-pitted. Fruits (4-)5-locular, glo-
bose, 15-35 (when dehisced to 40) mm in diam., echinate with broadly based,
mostly slightly curved pyramidalprickles to 3.5-4(-5) mm long, becoming gla-
brous when mature, dehiscent septicidally from 3-4 mm above base mostly not
up to the axis, and loculicidallyto 1/6 or '/3 from base; endocarpbearingimpres-
sions of the nerves of the exocarp; seeds 1 or 2 per loculus, superposed,obliquely
58 Flora Neotropica

tear-shaped,9-13(-15) x 5.5-7.5 mm, truncate, or at base either flattenedwhen

there are 2, ventrally obliquely flattened when only 1, apex very shortly beaked
or somewhat truncate, testa dull brown, finely reticulate with interspaces to ca.
0.1 mm; chalazal area roundish, contracted towards apex, to 5 or 7 mm long,
black; hilum to 0.7-1.5 mm broad, gradually contracted towards apex; black
caruncle present above the micropyle;embryos up to 8, one fully developed, the
others smalleror vestigial and crowded aroundthe tip of the large one, this very
thick, cotyledons unequal,punctate,radicleto 0.7 mmlong and projectingbeyond
ears of cotyledons.
Distribution.Venezuela, Trinidad,Surinam,Peru, Brazil (rare in the Amazon
basin), Paraguayand Argentina(Misiones). Fig. 15B.

Key to the Subspecies of Esenbeckia grandiflora

1. Petioles 4-25 mm long; flowers 8-14 mm in diam.; perianth hairy. 9a. subsp. grandiflora.
1. Petioles 2-5(-8) mm long; flowers 7.5-8 mm in diam.; perianth glabrous.
9b. subsp. brevipetiolata.

9a. EsenbeckiagrandifloraMartiussubsp. grandiflora.

Polembryum castanocarpum Jussieu, Mem. Mus. Hist. Nat. 12: 541, t. 15, fig. 49. 1825; Mem.
Rut. 136, fig. 49. Dec 1825 (reprinted from the first); G. Don, Gen. hist. 1: 807. 1831 (as "P.
castaneaecarpon" and erroneously attributed to Saint-Hilaire); Steudel, Nomencl. (ed. 2)
2: 367. 1841; Engler in Engler and Prantl, Nat. Pflanzenf. 3(4): 159. 1896, pro syn. TYPE.
Mem. Mus. Hist. Nat. 12: t. 28, fig. 49 (lectotype).

Petioles 4-25 cm long. Leaflets 3-22 x 1.2-11 cm or smaller.Inflorescences3-

7 cm long with side-branchlets2-3 cm. Flowers 8-14 mm in diam.; calyx lobes 1-
2 x 1.4-3 mm, strigillose below with hairs to 0.3 mm long; petals 3.4-6 x 2.2-
4.3 mm, thinlyor thicklycoriaceous,white to yellowish-creamwithgreenish,some-
times rose-tinge, papillose above, strigillosebelow with hairs to 0.4 mm; anthers
cucullate and much swollen, 1-1.5 x 0.7-1.2 x 0.6-0.8 mm, the tip with a yel-
lowish gland at maturity;disc sulcate to 4/10 of the thickness, very rarelyminutely
pilose; protuberancesof the carpels sparsely pilose with hairs to 0.3 mm long.

Key to the Varieties of Esenbeckia grandiflora subsp. grandiflora

1. Leaflets 5-22 x 1.7-11 cm; inflorescences usually 3-5 cm long with side-branchlets not over
2 cm; calyx lobes 1.3-2 x 1.8-3 mm; protuberances of the carpels to 0.8 or 1.5 mm long
9a-1. var. grandiflora.
1. Leaflets 3-9 x 1.2-3.2 cm or smaller; inflorescences to 7(-20) cm long with side-branchlets
to 3(-10) cm; calyx lobes 1-1.3 x 1.4-1.8 mm; protuberances of the carpels to 0.4 or 0.8
mm long. 9a-2. var. intermedia.

9a-1. Esenbeckia grandiflora Martius subsp. and var. grandiflora. Fig. 16.
Polembryumjussieui Schott, Rutac. 11, t. 6. 1834; Steudel, Nomencl. (ed. 2) 2: 367. 1841; Engler
in Martius, Fl. bras. 12(2): 147. 1874, pro syn. TYPE. Schott 749 delta = 4754 Herb. Bras.,
Brazil. Rio de Janeiro: Tijuca, fl (lectotype, W; isolectotype, NY ex W; both used for
drawing in Schott); Schott 749 (isotypes?), K 2x, one without collector or nr).
Colythrum grandiflorum (Martius) Steudel, Nomencl. (ed. 2) 1: 399. 1840.
Esenbeckia attenuata (Grisebach, Fl. Brit. W. I. 135. 1859; Miiller in Walpers, Ann. 7: 528. 1869;
Urban, Bot. Jahrb. Syst. 21: 553. 1896, syn. nov. TYPE. Cruger s.n., Trinidad. (St.
George, El) Tucuche, 28 May 1848, fl, fr (lectotype, K; isolectotype, GH).
Esenbeckia fasciculata Barbosa Rodrigues, Revista Engenharia 5: 1. cum icon. 28 Jul 1883, syn.
nov. TYPE. Brazil. Rio de Janeiro: Serra do Mar, nr. Rodeio, 360 m, Feb-May, fl (n.v.).
Esenbeckia grandiflora Martius var. macrophylla Chodat & Hassler, Bull. Herb. Boissier, ser.
2. 4: 1285. 1904, syn. nov. TYPE. Hassler 6857, Paraguay. Nr. Chololo6in basin of Rio Y-aci,
Esenbeckia 59

v E.pumila '/ /- ~ / / ", L I J '., I

/ / :. (

7I60 50 50 40

* E,--grndiflor var
gr-ndif Lor- .

v E grandiflorovar intermedia
ci . B
-
* E.grandiflorasubsp brevipetiolata-- 30

FIG. 15. Distribution of some species of Esenbeckia.

60 Flora Neotropica

FIG. 16. Esenbeckia grandiflora var. grandiflora. A, habit (F. C. Hoehne SP 28424, M). B, flower
(Hatschbach 17991, MICH). C, fruit (Leite 567, NY). D, seed (Leite 567, NY).
Esenbeckia 61

Dec 1900, fl, fr (lectotype, G-Herb. Chodat & Hassler; isolectotypes, BM, G-Herb. Delessert,
K, NY, P, S, UC, W).
Esenbeckia obovalifolia Pittier, Bol. Soc. Venez. Ci. Nat. 9: 122. 1944, syn. nov. TYPE. Delgado
298 ("2981," see note), Venezuela. D. F.: Between Las Flores and Papel6n, Selvas del
Avila, rd. from Ronda, Jul 1940, bud, fr (holotype, VEN-5776; isotype, US).
Esenbeckia grandiflora Martius var. peruviana Macbride, Field Mus. Nat. Hist., Bot. Ser. 13:
673. 1849, syn. nov. TYPE. Klug 3722, Peru. San Matin: Nr. Moyobamba, Zepelacio, Jul
1934, fl & fr (holotype, F-753378; isotypes, BM, GH, K, MO, NY, S, US).
Esenbeckia rigida Cowan, Bol. Mus. Nac. Rio de Janeiro, Bot. 27: 1. 1961, syn. nov. TYPE.
Segadas-Vianna et al. (Restinga I) 913, Brazil. Rio de Janeiro: Casimiroa de Abreu Co.,
nr. Barra de Sio Joao, 3 Sep 1953, fr (holotype, US-2279053; isotypes, R, n.v.).

Leaflets 5-16(-22) x 1.7-6.5(-11) cm. Inflorescences narrowly paniculate, usu-

ally ca. 3-5 x 2-4 cm with side-branchlets to 2 cm, bracts and bractlets 1.5-
2.5 x 1.1-1.8 mm. Flowers perfumed; calyx lobes 1.2-2 x 1.8-3 mm; petals 4.5-
6 x 2.2-4.3 mm, thickly coriaceous; carpels to 2 x 3.5 mm with protuberances
to 0.8 mm or some to 1.5 mm long.
Type. Martius 472, Brazil. Minas Gerais: Nr. Ouro Preto ("Villa Rica"), 1817-
1820, fl (lectotype, M, photo 19225 made by F at F, MO, NY).
Distribution. Venezuela and Peru in mountain- and rain forests, and Brazil in
rain forest, scrub, and capoeira. Reported from lateritic soil and once from thin
soil overlying calcareous rock. Also in restinga along the Brazilian coast; to 1550
m. Flowering in Venezuela Dec-Jan; in Peru Jul (1 record); SE Brazil (Aug-)Sep-
Mar(-May).
Specimens examined. VENEZUELA. Aragua: Pittier 15435 fr (US, VEN); Steyermark et al.
106496 fr (VEN). Terr. Amacuro: Blanco 507 fr (NY 2x, US, VEN). Nuevo Esparta: Gines 3577 fr
(US). Bolivar: Lasser 1483 fr (NY, VEN); Maguire 35808 fr (NY, US, VEN); Rusby & Squires s.n.
(Ciudad Bolivar, 1896) fr (NY); Steyermark 88527 fr (MO, NY 2x, U, UC, US, VEN, W).
TRINIDAD & TOBAGO. Trinidad. St. George: Finlay TRIN 3064 fr (TRIN, US); Purdie 65 fr (K);
Herb. TRIN 3853 fr (TRIN, G). County unknown: Broadway TRIN 5249 fr (NY, TRIN); arr. Hart
TRIN 1399 fl (TRIN, US), TRIN 1411 fr (TRIN, US); Kerr s.n. (Jan 1863) fl (K); Purdie s.n. fl & fr
(GOET, GH).
SURINAM. Brokopondo: LBB 3220 fr (BR, L, NY, U 2x, US). Marowijne: Lindeman et al. 493
fr (U 7x).
BRAZIL. Para: A. S. Silva et al. 17 fr (U) (not mapped). Ceara: Allemdo 279 fr (R 5x); Ducke MG
1549 fr (photo, NY); MG 2011 fl & fr (photo, NY); Herb. Saldanha 7584 st (R). Acre: Ule 5806 fl
(G, HBG, K, L). Mato Grosso: Manso 248 fl (BR). Bahia: Belem & Pinheiro 2548 fl* (NY); Duarte
6841 fr* (NY); Harley et al. 15253 fl (K 9x, U); 17558 fl (K 9x, U). Distrito Federal: Pires 58085 fr
(U); Prance & Silva 59058 fr (K, NY, P, S, U). Minas Gerais: Martius s.n. (Villa Rica) (BR, K, M);
Regnell III 373 (Feb 1847) fl (C 2x, K, LE, M, P, R, S), s.n. (4 May 1869) fr (S 2x); Vidal 1712 fr
(R 4x); Widgren s.n. (1845) fl (S). Espirito Santo: J. G. Kuhlmann 301 fr (NY). Sao Paulo: Anonymus
CGGSP 113 fr (C); Gaudichaud 752 fl (P); F. C. Hoehne SP 251 fr (SP, US); SP 1335 fl (SP), SP
1492 fl (SP), SP 2847 fl (SP), SP 28424 fl (F, M, NY 2x, P, S, SP, US), SP 31430 fl (SP); W. Hoehne
SP 30869 fr (SP); de Jonghe 479 fr (CGE); Koscynski 144 fl (SP); M. Kuhlmann 755 fr (SP), 1162 fr
(SP, US); Langsdorff s.n. (Dec 1825, Ypanema) fl & fr (LE); Lofgren CGGSP 1363 fr (P), CGGSP
4417 fl (SP); Manso 23 fl (W); Martius coll. lign. 12 st (M), 614 fr (M 2x); Mosen 3968 fr (P, S 2x),
4059 fl (LD, S); Novaes CGGSP 3305 fl (SP), SP 1950 fl (SP); Pereira 7097 fr (LP, M); Sellow B
2171, c 2168 bud (S); Smith & McWilliams 15389 fl (MICH); Usteri 40a fr (K); Vechi CPEF 282 fl
(R 2x). Rio de Janeiro: Glaziou 10458 fl (BR, C 2x, G, K, LE, P 2x, S), 11856 fl* (C, K, P, R);
Glaziou & Schwacke (in Glaziou) 18172 fl (C, K, LE, P); Glaziou et al. (in Glaziou) 8330 fr* (C, K,
P); Luschnath s.n. (Herb. Martius 464) fl (BR); Lutz 1423 fl & fr* (R 9x); (Manso) 247 (Herb. Martius
464) fl (BR); Herb. Martius 464 (pro parte) fl (BM, F, G 2x, HAL, K, LE, M, MO 2x, NY 2x, P, photo
at NY); Nadeaud s.n. fr (P 2x); Pereira et al. 3968 fr* (M); Pissis 12 fl (P 2x); Pulle & Lutz (in
Lutz) 1094 fr* (BM, R, U); Riedel 84 fl/fr (US); Schwacke R 71066 fl (R). Parana: Dusen 9478 fr (S),
13215 fr (S), 13788 fl (GH, S 2x), 14946 fr (S), 14964 fl (S), 16554 fl/fr (F, GH, K, MICH, NY, P, S),
s.n. (Valta Grande, 1908-1912) st (S), s.n. (Patrim6nio, 1908-1912) fr (F, GH, K, L, MICH, MO,
NY, P, S), s.n. (14 Jul 1911) st (GH, S), s.n. (6 Oct 1911) st (GH, NY, S); Hatschbach 3795 fl (US),
7703 fl & fr (L, NY), 8656 fl (US), 9446 fl (US), 17991 fl (MICH), 18587 fl (UC), 28378 fr (S, UC);
Hatschbach & Moreira 9411 fl (US); Hatschbach et al. 13248B fl (U, UC), 13908 fl & fr (U); Jonsson
62 Flora Neotropica

245a fr (GH, NY, S); Lindeman & de Haas 2378 fr (K, NY, U); Schwacke II 164 fl (R); L. B. Smith
et al. 14447 fl (GH, MO, NY, P, R, UC, US, WIS). Santa Catarina: Dusen 11788 fr (F, GH, K, L,
MICH, MO, NY, P, S), 17798 fr (S); Ehrhardt 13 fr (HBG 2x); Gevieski 60 fr (US); Inst. Malariologia
44 (17 Feb 1950) fl (S); Klein 493 fr (NY, S, UC), 600 fr (S, US), 761 st (US), 785 fr (S, US), 1073
fl (US), 1256 fl (US), 1564 fl (BR, K, M, US), 7175 fl (US); IM 44 (9 Apr 1951) fl & fr (S), IM 44 (24
Apr 1951) fr (S, US); Reitz 3141 fl & fr (S, UC, US), 6044 fl (F, US); Reitz & Klein 1116 fl (US), 2004
fr (NY, UC), 4688 fr (G), 4805 fr (P, UC), 6198 fl (BR, G, K, M, US, Z), 9040 fr (Z), 9461 fl (US),
9595 fr (G, US), 9633 fl (BR, GH, K, L, NY, P, U, UC, WIS), 12492 fl (UC, US); Smith & Klein
7389 fr (NY, RB); Smith & Reitz 12931 fr (C, F, LE, R, US); Ule s.n. (Jul 1891) fr (HBG). Rio
Grande do Sul: Camargo 61595 fl (S); Dutra R 71508 fl (R); Gaudichaud 16 fr (P); Krapovickas et al.
22959 fl (ZT); Leite 567 fr (NY), 2582 fr (A); Lindeman et al. ICN 21599 fl/fr (U); Malme It. Regn.
I 222 st (S 2x), It. Regn. II 867 bud (S); Rambo 413 fl (SP), 29104 fr (F), 30942 fl (S), 41841 fr (BR,
G), 42012 fl/fr (MO), 42733 fl (F, P, W). State unknown: Muller 164 (S. Brazil 1880) fl (P); Riedel s.n.
fl (LE); Riedel & Langsdorff 857 fr (LE); Sellow 1645 (?) (R 71065) fr* (R), 2171 bud (S), 5495 fl (K,
LE, R), s.n. (Herb. Hooker 1867 ex W) fl (K).
PARAGUAY. Caaguazu: Balansa 2519 fl (G), 2519a fr (G); Hassler 5129 fl (G 2x). Alto Parana:
Hassler 512G fl (BM). La Cordillera: Fiebrig 765 fl (E, F, G); Hassler 4019 fr (G), 12217 fr (A, BM,
E, G 2x). Paraguari: Balansa 3261 fr (P), 3262 fr (BM, C, G, K, L, P).
ARGENTINA. Misiones: Cabrera et al. 74 fr (LP); Devoto BAB 90738 fr (BAB 2x); Krapovickas
& Cristobal 13697 fr (UC); Meaca 31/138 fr (F); Meyer 5489 fr (UC); Milano & Buceta BAB 90812
fr (BAB); F. M. Rodriguez 495 fr (F, UC).
CULTIVATED. Hort. Butantan 1492 fl (GH); Hort. K s.n. (May 1857) fl (K), s.n. (1859) fl/fr (K).

Local names and uses. Trinidad:Gasparil(BroadwayTRIN5249); gaspari(llo)

colorado (Cruger s.n., El Tucuche; Purdie 65); confused with E. pilocarpoides?
Ceara: cocao (?) (Ducke MG 2011). Sao Paulo: apochi-taguara(=apojitaguara)
(Ignotus 12 in Herb. Martius); caputuna (Mosen 3968); carrapateirao (Regnell III
373 in P-Herb. Glaziou);pau-cutia,canela-(de-)cutia(Lifgren CGGSP4417; Ko-
scinsky 144); guaxupita(several records). Santa Catarina:cutia (several records);
cutia-amarela(Reitz 6044). Argentina, Misiones: canelero-del-monte(Milano &
Buceta BAB 90812). Uses: Accordingto Cowan and Smith(1973),used for helves
in Santa Catarina.According to Peckolt (1899: 338), the wood (of "E. fascicu-
lata") resembles boxwood, and its export was permittedonly to Portugal,being
"'madeirade lei." It was once reportedas source of the "famous walkingstick"
in Trinidad,but confusion with E. pilocarpoides may have taken place (Criiger
s.n., El Tucuche). The bark was used like that of E. febrifuga (see PHYTO-
CHEMISTRY).
Collections from the eastern Braziliancoasts (restinga, mata costeira), except
for Smith & MacWilliams15389, are distinguishedby their more rigid-coriaceous,
somewhat shiningleaflets, with apex often more or less conduplicateand some-
times rounded. The marginis more strongly revolute, the base broadly cuneate
or shortlyattenuate,and the leafletblade is usually subglabrousbeneath. Whereas
the leaflets of var. grandifloraare usually (not always!) obovate, the leaflets of
the restinga collections are mostly elliptic, though slightly obovate leaflets also
occur. These specimens undoubtedly belong to Polembryumjussieui Schott,
which is synonymous with Esenbeckia grandiflora Martius var. grandiflora.
Schott himself (1834) already added to his P. jussieui as observatio II: "An
Esenbeckia grandiflora Martii hujus generis?. " I agree with Engler in transferring
P. jussieui to E. grandiflora.In my opinion the restingacollections are ecological
variants the leaves of which have become adapted to the special climatological
conditions. The thickness of the leaves is the main differentialcharacterand this
is not a reliablebasis for a separatetaxon. The specimens concernedare indicated
in the list of studied specimens by an asterisk.
Esenbeckiaattenuata differs not at all from the present variety. The specimens
so identifiedwere all collected in Trinidadduringthe last part of the 19thcentury.
Esenbeckia 63

When Grisebach published his E. attenuata (1859), E. grandiflora was known

only from SE Brazil.
The comprehensive description of E. fasciculata Barbosa Rodrigues agrees
well with mine of var. grandiflora.Unfortunatelythe originalpublicationis very
rare and the copy I traced (at Kew) lacks the plate. The type is unknownin any
herbarium.
The type of E. rigida Cowan is certainlyidentical with the restingacollections
and therefore with Polembryum jussieui (=E. grandiflora var. grandiflora). With
regardto differencespointed out by Cowan, the (only 1) fruit is indeed somewhat
depressed-globose,but only on one side, due to incompletedevelopmentof some
of the loculi. The type specimen is glabrous, but the shoot apex appears to be
strigillose and therefore it does not differ as such from many specimens of var.
grandiflora.The fruitingstalk of the type is 6 mm long. This is 1 mm longer than
the flowering pedicels of var. grandiflora. However, it is quite normal in Pilo-
carpinae for pedicels to grow during maturationof the fruit, as the sepals do.
Cowan's description of the leaflet-blade "widely elliptic 10-12 x 5.5-6.5 cm"
agrees with my descriptionof var. grandiflorain conformitywith the SADT-chart
as "elliptic."
Esenbeckia grandiflora var. macrophylla Chodat & Hassler does not deserve
the status of a separate taxon. The leaflets of the syntypes are similarto those
of var. grandiflora.
The specimen Delgado 298 in VEN is annotated by Pittier as the type of E.
obovalifolia. Pittier, in his originaldescription, gives the collecting date as July
1940.Othercollections show that Delgado must have collected no. 298, not 2981,
at that time. Thereforeit is clear that the designationof the type of E. obovalifolia
Pittier as no. 2981 is a mistake. The descriptiondiffers in the size of the petals,
but Pittierfailed to mentionthat he studiedbuds only. The size of the calyx lobes
is erroneously given as 2.5 instead of 1.5 mm; the stigma is 5-lobed only (as
appears from the dissected flowers which were drawn and are still present). As
the type of E. obovalifolia fully agrees with E. grandiflora var. grandiflora, I
reduce E. obovalifolia to synonymy.
The type of var. peruviana Macbridewas collected from a ratherrobust plant
with some leaflets to 18 x 8.5 cm. The flowers, 13 mm in diam., are like those
of var. grandiflora.The fruits have ratherlargepricklesand the (immature)seeds
are somewhat larger. These differences are, in my opinion, not sufficientto sup-
port varietal status.
The inflorescences are usually small. Klein 1564, from Santa Catarina, has
inflorescencesto 11 x 5 cm with side-branchletsto 3 cm. In other charactersthis
specimen does not differ. Some specimens from Bahia also show inflorescences
10 cm long.

9a-2. EsenbeckiagrandifloraMartiussubsp. grandifloravar. intermedia(Martius

ex Engler in Martius)Kaastra, Acta Bot. Neerl. 26: 475. 1977.
Fig. 17.
Esenbeckia intermedia Martius ex Engler in Martius, Fl. bras. 12(2): 147, t. 32, fig. 2. 1874;
Engler in Engler and Prantl, Nat. Pflanzenf. 3(4): 159, fig. 94 D-J. 1896; Glaziou, Bull. Soc.
Bot. France 52. Mem. 3: 85. 1905 (as E. "intermededia").

Leaflets usually ca. 3-9 x 1.2-3.2 cm or smaller. Inflorescences to 7(-20) cm

long with side-branchletsto 3(-10) cm; bracts and bractlets somewhat swollen,
to 1-1.5 mm long; flowers more numerous than in var. grandiflora. Calyx lobes
64 Flora Neotropica

. ...

.
FIG* 17 Esenbeckia grand.flora var. . .nte .type CHerb Warming

FIG. 17. Esenbeckiagrandifora var. intermedia.A, habit (Glaziou2525, type, C-Herb. Warming
Esenbeckia 65

1-1.3 x 1.4-1.8 mm; petals 3.4-4.8 x 2.2-2.8 mm, thinly coriaceous; carpels to
0.5 mm high with protuberances to 0.4 mm and some to 0.8 mm long.
Type. Glaziou 2525, Brazil. Rio de Janeiro: Corcovado, 28 Jan 1868, fl
(lectotype, C-Herb. Warming 2432/1; isolectotypes, BR 2x, C-Herb. Warming
2432/2, F ex C, MO ex P, P, P-Herb. Glaziou, P ex Caen, U ex P ex Caen).
Distribution. Confined to Mts. Corcovado, Tijuca, and Pedro Bonita, in or near
Rio de Janeiro, Brazil. Flowering from Nov-Jan(-Mar).
Specimens examined. BRAZIL. Rio de Janeiro: Ackermann s.n. (Herb. Martius 1065) fl (BR);
Brade 10643 fl (GH, R 5x); Burchell 1423 fl (K, P); Glaziou 678 st (BR, C, P 2x), 1390 fr (BR, C, P),
3918 fl (C 2x, F, K, P 3x, R); Herb. Harvey s.n. (after 6 Aug 1858) fr (DS); Luschnath s.n. (Feb
1833) fl (LE); Herb. Martius 464 (p.p.) fl (GH, L 2x, LE, M, W), 1065 fl (BM; photos F, MO; HAL,
K, L 2x, LE 2x, M 2x, NY, P, W); Mello Filho 1107 fl & fr (R 14x, UC); Miers s.n. (Tijuca) fl (BM);
Mosen 2445 bud (S 2x); Nadeaud s.n. (30 May 1862) fr (P 3x); Riedel s.n. (without locality) fl/fr (LE);
Schenk Herb. Bras. 2330 fl (C); Herb. Warming 245111 fr (C).

Collection intermediate between var. grandiflora and var. intermedia: BRA-

ZIL. Rio de Janeiro: Corcovado: J. G. Kuhlmann RB 15384 fl (BM, F, NY).
The following collections could not be identified to variety because only fruits
were present: BRAZIL. Rio de Janeiro: Miers 3333x fr (BM). Locality unknown:
Olboze(?) 486 fr (BM).
The specimens belonging to var. intermedia are smaller in some aspects of
leaves and flowers than those of var. grandiflora. Other characters, like the in-
dument of the leaves and the diameter of the flowers, are like those of var.
grandiflora but somewhat on the small side, though not so different as Engler
(1874a) reports. Due to the local, limited distribution and the minor differences
of a quantitative nature E. intermedia should better be regarded as a variety,
under E. grandiflora.
Martius (1843: 39) published E. intermedia as a nomen nudum mentioning
"Herb. Flor. Bras. n. 1065. Apogi-tagoara Bras. (S. Paulo)." In the library of
the Institute for Systematic Botany at Utrecht a copy exists of Martius's Her-
barium florae brasiliensis (1837-1841) with a lithographed appendage called
"Martii herbarium florae brasiliensis III." This is a list of Martius's herbarium,
numbers 723-1310. Number 1065 mentions: "Esenbeckia intermedia M. (Eadem
n. 464. E. grandiflorae mixtae" (see below!). All the specimens of Martius herb.
1065 which I observed indeed belong to var. intermedia. However, because
"Apogi-tagoara" and "S. Paulo" were mentioned I suppose Martius exchanged
no. 1065 with Ignotus 12 in herb. Martius s.n. (M), which originated according
to the label from Sao Paulo, near Ipanema. On this label is mentioned "apochi-
taguara"; the specimen belongs to var. grandiflora. It is therefore not clear to
which taxon Martius referred in his Systema materiae medicae vegetabilis bras-
iliensis (1843), whether var. grandiflora or var. intermedia.
The collection Martius 464 appears to be a mixture of specimens belonging to
the two varieties. According to Martius's Herbarium florae brasiliensis this col-
lection was made "ad Macahe," but as Martius was never there himself (Urban,
1906: 103) he must have received the specimens through other persons and dis-
tributed them all under one number. In view of the geographic range of var.
intermedia I suppose those specimens which belong to this variety were not
collected near Macahe but near Rio de Janeiro (town).
J. G. Kuhlmann 15384 seems to be intermediate between the varieties. The
flowers have a hairy disc and style.
Mello Filho 1107 has inflorescences to 20 cm long, with side-branchlets to 10
cm.
66 Flora Neotropica

9b. Esenbeckia grandiflora Martius subsp. brevipetiolata Kaastra, Acta Bot.

Neerl. 26: 475. 1977.
Shrub 1-3 m tall. Petioles 2-5(-8) mm long; leaflet blade 4-9.5 x 2.2-5 cm or
smaller, the veins prominent on both sides. Inflorescences 3-5 x 2 cm; bracts ca.
0.8 x 0.1 mm. Flowers 7.5-8 mm in diam., green, glabrous; calyx lobes 0.5-
0.6 x 1-1.4 mm; petals 3.5 x 2 mm, coriaceous; anthers not cucullate, in outline
0.7 x 0.6 mm, not much swollen; disc with grooves to l/6 of its thickness; carpels
ca. 0.7 mm high, with tubercles 0.2-0.3 mm. Fruits provided with angulate-py-
ramidal, ca. 2.5 mm long spines, less echinate than in subsp. grandiflora.
Type. Belem & Magelhdes 914, Brazil. Bahia: Maraii, 25 Apr 1965, fl (holotype,
NY ex UB).
Distribution. NE Bahia, near Marau, and NE Pernambuco, Brazil. Restinga,
sometimes transitional to coastal forests. Collected once in flower in Apr.
Specimensexamined.BRAZIL. Bahia:Marau,Belem 1848fr (NY ex UB); Pereira 9601(& Pabst
8490) fr (R). Pernambuco or Paraiba: Tambe, Ducke & Andrade Lima 48 fr (R); Mordes EAN 2165
fl (NY, P, S, UC, all ex EAN).

This subspecies resembles the restinga collections of var. grandiflora. Differ-

ences can be found in the shorter petioles and in the smaller flowers with glabrous
perianth. The similar consistence of the leaves may be due to similar ecological
conditions.

10. Esenbeckia echinoidea Standley & Steyermark, Publ. Field Mus. Nat. Hist.,
Bot. Ser. 23: 164. 1944. Fig. 18.
Small tree, 5-7 m tall; indument of grayish-white hairs 0.1-0.3 mm long;
branchlets 4-6 mm in diam., grayish- to dark brown, densely minutely appressed-
pubescent, soon becoming glabrous. Leaves alternate, crowded at tip of branch-
lets, 1-foliolate, the leaflet sessile; petiole terete or somewhat flattened and slight-
ly canaliculate above, 1.2-6.5 cm long, minutely appressed-pubescent, tubercle
present or not; leaflet blade (broadly) elliptic, 6-28 x 3-15 cm, very shortly at-
tenuate or nearly rounded at base, obtuse or rounded at apex, often some emar-
ginate or slightly acuminate with a short blunt tip to 5 mm long, margin (slightly)
revolute, the blade chartaceous, dull green, paler beneath, subglabrous or mi-
nutely puberulous on both sides, rarely densely pilose beneath when young with
spreading hairs soft to the touch, venation camptodromous, ? prominent on both
sides, midvein plane above. Inflorescences axillary at tip of branchlets, erect,
paniculate, shorter than the leaves, ca. (2-)5-6 x (1-)2 cm, few-flowered, mi-
nutely pubescent; side-branchlets few, subopposite; bracts conduplicate, ca.
1.5 x 0.6 mm, minutely pubescent on the back; pedicels 1.5-2 mm long; bractlet
1. Flowers 8-8.5 mm in diam., protandrous; calyx lobes quincuncial, broadly or
depressedly ovate, obtuse, 1.6-2.5 x 1.9-2.4 mm, thinly coriaceous with papery
margins, pubescent below with hairs 0.1 mm long, venation actinodromous with
the median nerve distinctly thicker; petals deciduous, quincuncial, very widely
spreading, ovate, obtuse, 3-4 x 1.9-2.4 mm, transparent-papery, creamish-yel-
low, pubescent below with hairs 0.2-0.3 mm long, venation actinodromous, the
median nerve distinctly thicker; filaments deciduous, shortly subulate with a flat-
tened base, 2.5-3.5 mm long and 0.5 mm thick, glabrous; anthers heart-shaped,
1.7 x 1.2 mm, including a tip 0.2 mm, papillose; disc annular, 10-lobed, with
incurvations to half-way, 0.5 mm high and 2.5 mm in diam., fleshy, beset with
few protuberances to 0.5 mm long, glabrous; carpels adnate to the disc at base,
0.5 mm high, with protuberances to 0.4 mm long, glabrous; style attached near
Esenbeckia 67

'S '

FIG. 18. Esenbeckia echinoidea. A, habit (Standley74456 type F) B flower Pittier 113, US).
68 Flora Neotropica

base of the carpels, 1.6 mm long and 0.4 mm thick, the free part projecting0.7
mm beyond the protuberancesof the ovary and beset with hairs 0.3 mm long;
stigma clavate when immature,capitate when mature, 0.3 x 0.5 mm. Fruits de-
pressedly globose, stellately-lobed, 25-35 x 35-40 mm when closed, loculi
roundedon the back, echinate with very hard, pyramidal,longitudinallywrinkled
prickles to 7 or 9 mm long and 2-3 mm broad at base when mature, somewhat
sharp at the tip, and moderatelypilose with hairs 0.1-0.3 mm; fruit already de-
hiscent septicidally on the back when not yet maturefrom a few mm above the
base up to the axis, the loculi later splittingloculicidallyto half-way on the back;
seed probably 1 per loculus, obliquelytear-shapedca. 12 x 6 mm, with a slightly
curved beak 1 mm long at apex, testa dark brown, reticulate-colliculatewith
linear interspaces 0.1-0.2 mm long and rounded ones 0.05 mm in diam.; embryo
unknown.
Type Standley 74456, Guatemala. Chiquimula:Between Chiquimulaand Za-
capa, between Ramirez and Cumbrede Chiquimula,15 Oct 1940, fr (holotype,
F, photo 49602 made by F, F; isotypes, A, G, NY, US, all ex F).
Distribution.Guatemala.Rocky hillsides, gullies, shaded or bushy places and
forests. Flowering Apr-May. Fig. 6C.
Specimens examined. GUATEMALA. Guatemala: Locality unknown, Aguilar 95 fr (F); S slope
of Lake Amatitlan, Pittier 113 fl, fl/fr (US); nr. Amatitlan, Standley 61395 st (F). El Progreso: Nr.
Barranquillo, Steyermark 46452 fl (F, US 2x). Zacapa: Between Agua Blanca and Cumbre de Chi-
quimula, Standley 74413 st (F).

The fruits of this species superficiallyresemble those of the Mexican E. macr-

antha; the prickles, however, are more wrinkled and narrower, and pilose in-
stead of glabrous. The two species are otherwise well distinct from each other.
The present species differs from E. grandiflorain the color of the hairs which
gives a grayish-whiteappearanceto the terminalleaf buds, whereas the compa-
rable hairs of E. grandifloraare brownish. Otherdifferencesof E. echinoidea can
be found in the bases of the leaflets, the thickness of calyx lobes and petals, the
largerfruits with longer prickles which are not truly globose though they seem
to be so when viewed from a certain distance.

11. EsenbeckiamacranthaRose, Contr. U.S. Natl. Herb. 5: 111, t. 2. 1897;Wil-

son, in North Amer. Fl. 25: 201. 1911; Standley, Contr. U.S. Natl.
Herb. 23: 536. 1923. Fig. 19.
Tree to 7 m tall; branchlets4-10 mm in diam., light to mediumgrayish-brown,
later on darkgray-brown,minutelypubescent with hairs 0.2 mm long, becoming
glabrous.Leaves alternate,(2-)3-foliolatewith stalkedleaflets;petiole semiterete,
(hardly)canaliculatetowards the tip, 2.5-7 cm long and to 3 mm thick, hoary to
tomentose or minutely pubescent, the hairs 0.2-0.5 mm long, tubercle usually
present; petiolules of lateralleaflets 0-1 mm long, of terminalone 2-6(-10) mm;
leaflet blades mostly elliptic or obovate, 6.5-18 x 3-9 cm or the lateral ones
shorter, (shortly) attenuate and especially the lateral ones unequal at base, at
apex obtuse or rounded, often slightly acuminate, occasionally emarginateor
acutish, margin thick, often revolute, the blade subcoriaceous or chartaceous,
(sparsely) pubescent above with hairs 0.2-0.4 mm long, + tomentose below with
hairs 0.5-0.6 mm long, venation camptodromous,prominent on both sides or
plane in chartaceousleaves, the midvein impressed above. Inflorescencesat tips
of branchlets,consisting of up to 4 erect, paniculate,short- or long-stalkedpartial
inflorescences 5 x 6-7 cm, tomentose with spreading hairs 0.2-0.6 mm, the
many-floweredside-branchletsalternate;bracts caducous, narrowlyovate to nar-
rowly triangular,somewhat concave, to 2.8 x 0.8-1 mm, beset with some ap-
Esenbeckia 69

FIG. 19. Esenbeckia macrantha. A, habit (Nelson 1831, type, GH). B, flower (Nelson 1831, type,
GH). C, fruit (C. E. Smith et al. 3721, US). D, fruit (Conzatti 4031, US). E, fruit (Conzatti 4106,
US). F, seed (Conzatti 4106, US). G, embryo (Conzatti 4106, US).
70 Flora Neotropica

pressed hairs at base above, densely pilose below with spreadinghairs 0.5 mm
long; pedicels ca. 2 mm long; bractlets 2, opposite or alternate, some subtending
secondarypeduncles.Flowers9-10.5 mmin diam.;calyx lobes cochlearto (sub)sep-
arate, depressedly ovate, rounded at apex, 2.4 x 3.4 mm, coriaceous, thick at
base, glabrousabove, densely pubescent below with spreading,hyaline hairs 0.3
mm long, venation parallel,the nerves branched;petals persistent, cochlear, very
widely spreadingand often reflexed, adnate to the disc, elliptic, 4.2-5 x 2.2-3.2
mm, obtuse, papery, yellowish, glands inconspicuous, densely pubescent below
with spreading,hyaline hairs 0.2-0.3 mm long, venation actinodromous,the me-
dian nerve thicker;filamentsoften persistent, subulate,terete but in cross-section
transverselynarrowlytriangulartowardsbase, 2.5-3.5 mm long and 0.5 mm thick,
glabrous; anthers heart-shaped,including a mucro 0.2 mm, ca. 1.7 x 1.2 mm;
disc annular,rosette-like, 10-plicate to '/3 or 2/5 of its thickness, 0.5-0.8 x 0.7-
0.8 mm and 2.7-3 mm in diam., very thickly fleshy, charged with small glands,
glabrous;carpels adnate to the disc but free distally, 0.5 mm high, providedwith
an apicalapophysis0.2-0.3 mm, very densely beset with finger-likeprotuberances
to 0.3 mm long, and sparsely pilose with hairs 0.3 mm long; style inserted near
base of carpels, 3 mm long, the free part 2 mm long and 0.3-0.4 mm thick, beset
with few spreadinghairs 0.3 mm long; stigma capitate, 0.2-0.3 x 0.3-0.5 mm.
Fruits 2-3 per whole infructescence, depressedly globose, 2-3 x 3 cm (-5 cm
when dehisced), glabrous;loculi roundedon the back, very densely chargedwith
numerous + conical protuberancesvarying in size to ca. 10 x 7 mm, dehiscent
septicidally from ? 6 mm above base to the tip and loculicidally to 1/3 on the
back; seeds obliquely tear-shaped, ventrally obliquely flattened at base, ca.
16 x 10-13 x 11 mm, with a short curved beak 0.5-1 mm long, testa firm, 0.5
mm thick, medium to dark brown, colliculate with roundish interspaces 0.05-
0.10 x 0.05 mm but those on the back of the seed narrowly elliptic and up to
0.20 x 0.05 mm, the interspaces not swollen or only slightly so; chalazalarea not
visible; hilum narrowlyobtrullate,to 4 mm broad; a distinct caruncle above and
beside the micropyle;embryo 1 per seed, cotyledons unequal, radicle as long as
the ears and projectingca. 1 mm beyond them, plumule 2-leafed, 0.8 mm long.
Type Nelson 1831, Mexico. Oaxaca: 6 mi. above Dominguillo,30 Oct 1897, fl
(holotype, US-49400, photo madeby NY, NY with fragmentof holotype; isotype,
GH ex US).
Distribution. Mexico, Puebla and Oaxaca. In understorey on gravelly soil (I
record);alt. 1200-2100m. Collected in flower Oct-Nov. Fig. 6C.
Specimens examined. MEXICO.Puebla: Coxcatlain,Purpus4196 fl/fr (GH, NY, UC); Tehuacian
region, above Calipan,C. E. Smith et al. 3721 fr (F, G, US). Oaxaca: Cuicatlan,Portillo, Conzatti
4031 fr (US); Frotitlan,Cuesta de San Bernardino,Conzatti4106 fr (US).
This species has remarkablelarge fruits with usually robust protuberancesand
large seeds. The fruits of Conzatti4031 are rathersmooth with tuberclesto 3 mm
long, and more deeply stellately-lobed with the loculi rounded to slightly trian-
gular on the back.

12. Esenbeckiaflava Brandegee, Zoe 1: 378, t. 12. Feb 1891;Rose, Contr. U.S.
Natl. Herb. 5: 111. 1897;Wilson, in North Amer. Fl. 25: 201. 1911;
Standley, Contr. U.S. Natl. Herb. 23: 536. 1923. Fig. 21.
Shrub or small tree to 8 m tall with trunk 20 cm in diam.; bark usually light
gray and smooth, crown spreadingwith crooked branches;branchlets4-8 mm in
diam., grayish-brown,strongly longitudinallywrinkled,grayish-white-tomentose
with hairs 0.5 mm long, becoming glabrous, leaf scars with swollen margin.
Esenbeckia 71

Leaves alternate, simple; petiole semiterete, 0.8-2.5 cm, tomentose with hairs
0.6 mm long; blade elliptic or oblong, rarelyovate, (2.8-)4-12(-19.3) x (1.7-)1.8-
6(-8.8) cm but in the inflorescences smaller, at base (very) shortly attenuate,
cuneate, or rounded, slightly unequal, at apex emarginateto retuse, or rarely
obtuse, marginslightly revolute or not, the blade chartaceousto subcoriaceous,
dullgrayish-greenon both sides, tomentoseor pilose, with spreading? curvedhairs
0.3-0.8 mm long, soft to the touch, becoming less densely hairy above, venation
distinctly camptodromousand finally prominent, costa plane above. Inflores-
cences densely flowered consisting of terminal and axillary, erect, paniculate,
partial inflorescences 3-9 x 3-6 cm, indumentlike that of the young vegetative
branches, side-branchletsalternate and subtended by a bract or a small leaf;
bracts caducous, narrowly triangular, ? conduplicate, to 4 x 1 mm, glabrous
above, densely pilose below with spreadinghairs 0.5 mm long; pedicels to 3 mm
long; bractlets 2, alternate. Flowers 10.5-15 mm in diam., with "sickish-sweet"
odor; calyx lobes quincuncial,broadly or depressedly ovate, 1-2 x 1.7-2.5 mm,
rounded at tip, thickly papery, marginmembranous,(dispersedly) pilose below
with spreading hairs to 0.5 mm long, venation parallel but the nerves distally
branched;petals persistent, quincuncialor cochlear, adnate to the disc at base,
spreading,broadly elliptic, 5-6.5 x 3.7-5 mm, roundedat the apex, transparent-
papery, yellowish when dried, creamish-whitewhen fresh, the indumentlike that
of the calyx but less dense, venationactinodromous,the nerves branchedtowards
the margin;filamentspersistent, adnate at base to the disc, subulate, 3-5.5 mm
long and 0.7 mm thick, glabrous;anthers ovoid, (1.3-)1.6 x 1.2 mm includinga
slightly incurved tip 0.1-0.2 mm, papillose; disc annular,rosette-like, irregularly
10-lobedto 1/3of its thickness, 0.8 mm high and 1.6 mm thick, 3-4.2 mm in diam.,
thickly fleshy, glabrous; carpels to 1 mm high, provided with numerousfinger-
like outgrowthsvarying to 0.5 mm long, glabrous;style inserted on the proximal
half of the carpels, 3.2-3.5 mm long, 0.2-0.4 mm thick, the free part 1.5-2.5 mm,
glabrous; stigma capitate, obsoletely 5-lobed, 0.3-0.5 x 0.4-0.7 mm. Fruits 2-3
per infructescence, stellately-globose,depressed, 2-2.5 x 3-4 cm, to 6 cm broad
when dehisced;loculi roundedon the back, (mostly strongly)muricateto echinate
with rathersharp prickles varyingin length to 6 or 10 mm, charged with glands,
glabrous, incompletely dehiscing septicidally from 2 mm above base to 2 mm
from tip, and loculicidally to ? 1/5 above base, the fruit walls strongly spreading
finally; exocarp with ? reticulatelybranchednerves; seed usually 1 per loculus,
obliquely subglobose, 10-14 x 11-14 x 9-10 mm, with a blunt or shortly beaked
apex 0.05 mm, testa dark brown, shining, linear-colliculatewith the interspaces
(up to 0.20 x 0.05 mm) parallel together in patches alternatingwith granulate
patches; chalazal area not visible; hilum to 3 mm broad, permanentlycovered
with the axial part of the endocarp;embryo l, cotyledons unequal with ears 0.8
mm long, radicle 1.3 mm long projecting0-0.5 mm beyond the ears.
Type. Brandegee 89, Mexico. Baja California:San Jose del Cabo, Sep 1890,
fl & fr (lectotype, UC-109534-Herb.Brandegee).
Distribution.Mexico, Baja California.Canyons, gravelly slopes, andfrequently
along brooklets; alt. 100-1050m. Flowering Sep-Nov. Fig. 20A.
Specimens examined. MEXICO. Baja California: Herb. Bailey 234 st (F); Brandegee s.n. (1890)
fl & fr (US), s.n. (Sep 1891) fl & fr (A 4x, DS, GH, NY), s.n. (17 Oct 1899) fl (NY, UC), s.n. (1901)
fr (NY), s.n. (Nov 1902) fl (US); Carter & Chisaki 3613 fr (DS, MICH, UC, US); Carter & Ferris
3439 st (DS, UC); Carter & Kellogg 3275 fl/fr (DS, K, MICH, US); Carter et al. 501 fr (DS, MICH,
UC, US); Gentry 11233 fl/fr (MICH), 12343 fr (MICH); Hastings & Turner 64-207 fl (DS), 64-252 fr
(DS); Johnston 4087 bud (GH, K, UC, US); Jones 22355 fr (F, MO), 27433 fl (DS), 27433a fl (DS,
MO, NY, UC); Mason 1878 fr (K, US); Moran 3806 fl/fr (DS, UC), 3881 dec. fr (DS, UC), 6899 fr
(DS, K), 9344 st (UC), 11774 st (DS, UC), 11793 dec. fr (DS); Nelson & Goldman 7258 fr (BM, US);
72 Flora Neotropica

3 - 115 110_________ 105 __5

100 95_ 0
___-

3'

* E.hartmonii -

-----7---"--r-T?----}--r*--.; ?C--:-:

* E.cornuta-
v E. oligantho , /
|
FIG. 20. Distribution of some species of Esenbeckia.
Esenbeckia 73

.1e

1Oc189NY.C,fut(oa689DS. D eicdfut(ign 61 Y........ (Wggin

FIG 2. Esenbeckia flava. A, habit (Brandegee sn., Sep 1891, GH) B flower (Brandegee s.n.,
17 Oct 1899, NY). C, fruit (Moran 6899, DS). D, dehisced fruit (Wiggins 5671, NY). E, seed (Wiggins
5671, NY).
74 Flora Neotropica

Peters 94 st (UC); Rose 16345 dec. fr (US); Rzedowski 26572 fl (MICH); Shreve 7241 fr (F); Whitehead
841 st (DS); Wiggins 5590 fr (A, DS, MICH, NY, UC, US), 5671 fr (DS, GH, MICH, NY, UC, US),
11503 st (DS), 14543 fr (DS, K), 15015 fr (DS), 15425 fr (DS, UC); Xantus s.n. (Cape S. Lucas, 1859/
1860) fr (GH).
Local names and uses. Palo amarillo(several records);palo morio (Bailey234).
According to the original description the wood was often used for poles in the
constructionof houses.
The species is usually collected in the fruitingstage. The fruits are very large,
especially when nearly ripe, their size relatedto the incomplete septicidalrupture
which results in widely patent loculi. The fruits are rarely 6-locular.
In Brandegee s.n. (Nov 1902) one of the flowers observed is male, with a
rudimentaryovary. The disc in this flower has five projecting, triangularpoints
covering the ovary and the rudimentarystyle.
13. EsenbeckiahartmaniiRobinson & Fernald, Proc. Amer. Acad. Arts 30: 115.
1894;Rose, Contr. U.S. Natl. Herb. 5: 111. 1887; Wilson, in North
Amer. Fl. 25: 200. 1911; Standley, Contr. U.S. Natl. Herb. 23: 536.
1923; Engler in Engler and Prantl, Nat. Pflanzenf. (ed. 2) 19a: 281.
1931. Fig. 22.
Usually a small, stiff-branchedpolypodial shrub, or a treelet to 5 m tall with
stem 15-25 cm in diam., with light gray bark; branchlets3-7 mm in diam., dark
grayish, minutely pubescent with hairs 0.2-0.3(-0.5) mm long, becoming gla-
brous. Leaves alternate,simple;petiole semiterete, 0.2-0.8(-1.2) cm long, dense-
ly minutelypubescent with hairs0.2-0.5 mm long; blade (narrowly)(sub)obovate,
1-7(-8) x 0.5-2.7(-4) cm, at base cuneate or acutish and slightlyunequal,at apex
emarginate,rounded, or retuse, the marginflat or subrevolute, the blade subco-
riaceous, dull green on both sides, above subglabrousor puberulousbut minutely
pubescent with hairs 0.3 mm long on the lower part of the costa and at base,
pubescent below with hairs to 0.4(-0.6) mm long, venation camptodromous,
prominenton both sides, costa plane above. Inflorescences terminal, erect, pa-
niculate, 2-5(-8) x 2-5(-6) cm, densely minutely pubescent with hairs 0.2-0.3
mm; side-branchlets alternate; lower bracts persisting into anthesis, leaf-like,
narrowly(sub)obovate, to 3 x I mm, minutely pubescent; upper bracts like the
2 bractlets deciduous before anthesis, elliptic, shorter than the lower ones; ped-
icels to 8 mm long. Flowers 12-14 mm in diam.; calyx lobes quincuncial,broadly
to depressedlyovate, 1.5-1.8 x 1.7-2.2 mm, roundedat tip, coriaceous, minutely
pubescent below with hairs0.2-0.5 mm long or nearlyglabrous,venationparallel,
the nerves branched towards the margin;petals persistent, cochlear, adnate to,
the disc at base, very widely spreading, elliptic or (sub)obovate, 4.5-5.5 x 3-4
mm, roundedat tip, papery with thin margin,white or somewhat yellowish when
fresh, pale yellowish when boiled, papillose above, the indumentbelow like that
on calyx, venation actinodromous, the nerves branching towards the margin:;
filamentspersistent, adnate to the disc at base, subulate, 4-5 mm long and 0.5--
0.7 mm thick, glabrous; anthers heart-shapedto ovoid, (1.4-)1.6-1.7 x 0.9-1.1
mm, includinga tip 0.2 mm, papillose;disc rosette-like, with + 10 irregular,slight
incurvations, 0.4-1 mm high and 3-3.5 mm in diam., thickly fleshy, glabrous;
carpels proximallyadnate to the disc, 0.4-0.6 mm high, densely beset with thick
conical to pear-shapedprotuberances0.3-2 mm long occasionally chargedat tip
with a hyaline hair to 0.3 mm long; style inserted on the lower half of the carpels,
contorted when dry, 4.2-4.5 mm long, the free part 3.2-4 mm long and 0.4-0.5
mm thick, glabrous; stigma capitate, slightly and irregularlylobed or 5-lobed,
Esenbeckia 75

FIG. 22. Esenbeckia hartmanii. A, habit (Goldman 246, GH). B, branch (Palmer 1801, US). C,
flower (Gentry 2254, A). D, fruit (Gentry 6773, MICH). E, seed (Gentry 6773, MICH).
76 Flora Neotropica

0.3 x 0.3-0.5 mm. Fruits to ?3 per infructescence; I or more loculi frequently

not developed, depressed, stellately-globose, 1.2-1.4 x 2-2.5 mm, to 3 cm broad
when dehisced; loculi globose, rounded on the back, glabrous, muricate with
short, rather sharp broad-based prickles 1-3 mm long, septicidally dehiscent from
2 mm above base to 2 mm from tip, and loculicidally dehiscent to 2-3 mm above
base, eventually widely spreading, the nerves on the inside of the exocarp retic-
ulate; endocarp very thin, often irregularly ruptured upon dehiscence; seed 1 per
loculus, obliquely subglobose 8-9 x 8-10 x 8-10 mm, with a straight, blunt or
very short mucronate apex, testa 0.5 mm thick, dark brown, shining, linear-col-
liculate 0.2-0.3 x 0.05 mm, and colliculate 0.05 mm in diam., the interspaces
parallel together in patches; chalazal area not visible; hilum 2.5-3 mm broad;
embryo 1, cotyledons unequal with ears to 0.5 mm, radicle 0.5 mm long not
projecting beyond the ears, plumule 0.2 mm long.
Type. C. V. Hartman 240, Mexico. Sonora: La Tinaja, 19 Nov 1890, fr (ho-
lotype, GH, photo, DS; isotypes, K, NY 2x, US).
Distribution. Mexico, Sonora and Sinaloa. Dry areas in littoral dunes, lava
fields, coastal plains on heavy clay soil, granitic slopes, thickets and thorn forests,
sometimes transitional to desert; alt. 30-1100 m. Flowering Jun-Jul. Fig. 20A.
Specimens examined. MEXICO. Sonora: Gentry 2254 fl (A, F, MO), 6773 fr (MICH), 14384 fr
(US); Hastings & Turner 65-88 fr (DS), 65-174 fl/fr (DS), 69-176 fr (DS); Mason & Brewer 1731 fr
(DS); Shreve 6765 st (F, MO); White 2921 fl (GH, MICH); Wiggins 7459 fr (A, DS, F, MICH, UC,
US, VT). Sinaloa: Gentry 7068 fl (F, MICH); Goldman 246 fr (F, GH, US); Ortega 1212 fl (US),
4204 fl (US), 4621 fl (US), 6348 fl/fr (DS, M, US), 6398 fr (DS, GH, US); Palmer 1801 fr (C, F, GH,
NY 2x, S, US 2x); Rose et al. 13609 fr (GH, NY, US), 14735 fr (NY, US).

Local names. Sonora: sa.mota (White 2921). Sinaloa: crucecilla (?) (Ortega
1212).
The endocarp is distinctly thinner than in other Esenbeckia species, but is still
cartilaginous.
The lower side of the leaves of Rose 14735 is somewhat tomentose.

14. Esenbeckia leiocarpa Engler in Martius, Fl. bras. 12(2): 145, t. 32, fig. 1. 1874;
Engler in Engler and Prantl, Nat. Pflanzenf. 3(4): 159, fig. 94C.
1896. Fig. 23.
Tree 5-15 m tall with trunk to 30 cm in diam., indument with hairs to 0.2 mm
long; branchlets 4-5 mm in diam., light grayish-brown; hoary with minute, seri-
ceous, hyaline hairs, becoming glabrous in age, the elevated margin of the leaf
scars very densely beset with sericeous hairs. Leaves subopposite or alternate,
simple; petiole flattened and often slightly canaliculate, 9-30 mm long, hair-cover
like that on the branchlets; blade (narrowly) elliptic, 4.5-18 x 1.7-8 cm, attenuate
at base, the very base abruptly ending or sometimes minutely subauriculate,
obtuse or somewhat shortly acuminate at apex, margin recurved, the blade char-
taceous, (dispersedly) appressed-pubescent on both sides, venation ? campto-
dromous, costa plane above with a rib which vanishes in the sulcus of the petiole.
Inflorescences terminal, leafy panicles with numerous flowers, 15-20 x 20-30
cm, consisting of several axillary, erect, very widely paniculate partial inflores-
cences mostly longer than the leaves and ca. 8-17 cm long, with alternate or
subopposite side-branchlets 4-5(-7) cm; hair-cover like that of non-flowering
branchlets but the hairs more spreading; bracts caducous; pedicels 1.5-2 mm,
provided at their bases with 1 narrowly trigonous bractlet, + semiterete in cross-
section 0. 1-1 x 0.05-0.3 mm, sericeous-pubescent on all sides. Flowers 4-5(-6)
Esenbeckia 77

..I ~ ~B

ON^IIrF <^N

FIG. 23. Esenbeckia leiocarpa. A, habit (Novaes 517, US). B, base of leaf (Novaes 517, US).
C, flower (Novaes 517, US). D, fruit with Dl, interior face (W. Hoehne FFO 2995, F). E, seed (W.
Hoehne FFO 2995, F). F,F, embryHoehne
embryo
(W. FFO 2995, F).
78 Flora Neotropica

mm in diam.; calyx lobes quincuncial, not or scarcely overlappingat anthesis,

(very) broadly ovate, 0.7-1.3 x 0.8-1.3 mm, ? acute, the very tip shortly acu-
minate, the blade coriaceous, appressed-silky-pubescenton both sides with hairs
to 0.1 mm long, often sparsely so above, nerves ? parallelbut somewhatbranch-
ing distally; petals deciduous, cochlear to subvalvate, widely spreading,elliptic
to ovate, 2.1-2.3 x 0.9-1.5 mm, acuminateat apex, softly coriaceous, thickened
towards apex and the very tip somewhatuncinateup to 0.2 mm, creamish-white,
glands hardly visible, papillose above, shortly ciliate at the margin, silky ap-
pressed-pubescentbelow with hairs to 0.1 mm long, venation + parallelwith the
primary nerve thicker than the 2-4 secondary ones and visible externally, all
principalnerves branchingbefore reachingthe tip; filamentsdeciduous, subulate
to trigonous with a flattened base, 1.5-2 mm long, provided with glands and at
base on abaxialside with silky, appressedhairs to 0.2 mm long; anthersdorsifixed
13 to 12 from the base, heart-shaped,including a tip of 0.05-0.2 mm ca. 0.6-
0.7 x 0.6-0.7 mm, papillose; disc annular, 10-lobed, with grooves to 1/3 of its
thickness, 0.4 mm high, 0.4 mm thick (as high as the ovary) and 1.2-1.8 mm in
diam., greatly thickened upwards, with glands, silky-pubescentwith hairs to 0.2
mm long; carpels adnate to the disc, connate, provided with a free apophysis
0.2-0.3 mm high and very densely silky-pubescentwith hairs 0.2 mm long; style
inserted near base of carpels, 0.6-0.9 mm long and 0.1-0.2 mm thick, the free
projecting part 0.4-0.5 mm, glabrous; stigma clavate-capitate, slightly 5-lobed,
0.2-0.3 x 0.3-0.4 mm. Fruits to ca. 5 per vegetative branch, subobovoid, 2-
2.5 x 2 cm when still closed, hoary like the branchlets;loculi smooth, provided
V?from tip with an obsolete, blunt apophysis to 1 or 2 mm long, dehiscent im-
mediately when ripe loculicidally and septicidally, from the tip to the base, the
fruit then soon deciduous; nerves of the exocarp hardly visible internally
but prominent externally; seeds mostly 2 per loculus, obliquely tear-shaped,
6(-10) x (6-)7.5-8 mm and (3.5-)4.5 mm thick, (somewhat obliquely) flattened
at base, apex blunt, testa brown, shining, finely irregularlygranulate-reticulate.,
chalazal area elliptic to roundish, 4-6 x 3-3.5 mm, black; hilum 0.8 mm broad;
embryo 1 per seed, with thick, unequal finely punctate cotyledons, radicle pro-
jecting beyond the ears, plumule with 2 leaf initials.
Type. Helmreichen29, Brazil. Sao Paulo, fl (lectotype, M ex W, the specimen
labeled 429; isolectotypes, LE ex W (with indistinct number),NY ex W, US ex
W, W).
Distribution.Brazil, Mato Grosso to Bahia and Sao Paulo. In forest and ca-
poeirao. Flowering Nov-Jan(-Mar). Fig. 20B.
Specimens examined. BRAZIL. Mato Grosso: Herb. Martius 1069 fl (K, M, W 2x). Bahia: Belem
& Pinheiro 2815 fl (NY, US). Espirito Santo: J. G. Kuhlmann 6613 fl (BM, F, MO, NY 2x). Sao
Paulo: F. C. Hoehne CPEF II 163 fl (R); W. Hoehne FFO 2995 fr (F); Hunger Filho s.n. (Navarro
de Andrade, Jul 1928) fr (P); M. Kuhlmann 849 fr (SP), 4508 fr (SP); Lofgren CGGSP 4416 st (SP);
Manso (in Riedel) s.n. (1835) fl (LE); Martius s.n. (nr. Ypanema, Jan) fl (M 2x); Novaes 517 fl (SI'P,
US); CGGSP 3216 fl (SP); Vecchi CPEF 163 fl (P, R, SP). Rio de Janeiro: Glaziou 675 fl (BR 2x, P),
1392 fl & fr (BR 3x, C 2x, P 2x), 3917 fl (C, K, photo, NY; P 3x, R, VT), 10460 fl (C 2x, LE, P);
Horto Florestal RB 100583 fr (BM, F), 100585 fl (BM); Nunes EFCB 17 fl (R 2x); SP 20923 fl & fr
(SP); Riedel 35 (?) fr (LE), s.n. (nr. Rio de Janeiro) fl & fr (BM, M); Schott & Pohl D 999 = Herb.
Bras. 56118 fl (F, K, NY, US, W). State unknown: Anonymus 1938 (Herb. Hooker, 1867 ex W) fl
(K); Helmreichen s.n. fl (BR 2x).
CULTIVATED. In Sao Paulo and Rio de Janeiro: Ducke RB 8556 fl (S, U, US); Faria 11 fl (SP);
Gehrt SP 29999 fl (SP); Handro s.n. (20 Jan 1944) fl (MICH, SP); Hunger Filho EFSP 301 fl (SP).

Local names. Sao Paulo and Rio de Janeiro:guarantan(=hard wood) (several

records). Espirito Santo: guarataia-vermelha(J. G. Kuhlmann6613).
This species is close to Esenbeckia cornuta, which see.
Esenbeckia 79

Glaziou 1392 (C) is the only syntype that was annotated by Engler as Esen-
beckia leiocarpa. All other specimens were annotated as "E. laevicarpa." The
latter, unpublishedname was probablychangedby Englerbecause it is composed
of mixed Latin and Greek stems.
In Riedel s.n. (M) embryos with 3 cotyledons were observed.

15. Esenbeckiacornuta Engler in Martius, Fl. bras. 12(2): 146. 1874; Engler in
Engler and Prantl, Nat. Pflanzenf. 3(4): 159. 1896; Macbride, Field
Mus. Nat. Hist., Bot. Ser. 13: 672. 1949. Fig. 24.
Shrub or tree?; branchlets4-5 mm in diam., hoary with silky pubescence of
ca. 0.2 mm long. Leaves alternate, simple; petiole subterete, slightly flattened
distally, 6-18 mm long, pubescence like that on the branchlets;blade elliptic, 4.5-
18 x 1.7-8.5 cm, acute to obtuse at base, ? acuminate at apex with an obtuse
acumen to ca. 1 cm, marginrecurved towards base, the blade chartaceous, mi-
nutely pubescent above with silky hairs to 0.4 mm long, spreadingtowards base
and appressed distally, silky-puberulousbelow with the same hairs as above and
also densely pilose with thickerhairs to 0.3 mm long, the vesture below distinctly
perceptibleto the touch, venation ? camptodromous,costa impressedabove. Inflo-
rescences with numerousflowers at tips of the branchlets,ca. 11-16 x 10-18 cm,
the hair-coverlike that of the young vegetative branchlets, composed of several
axillary, erect, (very) widely paniculate partial inflorescences ca. 8 x 5 cm;
branchletsalternateor subopposite;pedicels 1.5-2 mm long; bracts and bractlets
caducous. Flowers 4.5-5 mm in diam.; calyx lobes quincuncial, tending to be-
come separate, ovate, acuminate, 0.7-1 x 0.5-0.7 mm, coriaceous, with glands
hardly observable, appressed-pubescenton both sides with hairs 0.05-0.1 mm
long, strigillose below, especially along the margins, with silky-hyalinehairs to
0.3 mm long, venation ? parallel;petals imbricate,widely spreading,elliptic, ca.
2.1-2.4 x 1.1 mm, acuminateand thickenedat apex, the very tip uncinatefor up
to 0.2 mm, thickly papery to thinly coriaceous, probably yellowish when fresh,
papillose above, pilose below with thick, silky hairsto 0.2 mm long, glandshardly
visible, venation hyphodromous,only the primarynerve observable, the (usually
2) secondary nerves faint and invisible; filaments subulate with flattened base,
1.5 mm long, provided at base with few appressed hairs 0.1 mm long, papillose;
anthers broadly heart-shaped,includinga tip 0.05 mm, 0.6 x 0.6 mm, papillose;
disc annular, 10-plicate,to 0.4 mm high, ca. 0.2 mm thick (as high as the ovary
with its apophyses), 1.2 mm in diam., dark-pigmented,glabrous;carpels adnate
to the disc, connate with each other, 0.2-0.3 mm high, provided with a free
apophysis 0.2-0.3 mm and at once very densely silky with hairs 0.2 mm long and
chargedwith 1-3 tuberclesof 0. 1-0.2 mm; style insertednear base of the carpels,
0.9 x 0.1-0.2 mm, the free part exserted 0.4 mm beyond the apophyses, some-
what thickened towards the tip; stigma clavate-capitate, 0.2 x 0.2 mm. Fruits
(according to Engler, 1874a, translated):"capsules 5-locular, carpels subtrigo-
nous, at last becomingglabrous, provided with a rathertall, ascending horn mid-
way on the back. Capsules lignified, carpels 1.75 cm long and I cm broad, de-
hiscent ventrally nearly to the base, and on the back half-way to the horn which
is 0.5 cm long"; young fruits observed by the present author ca. 9 mm high,
hoary with appressed, shining hairs 0.1-0.2 mm long.
Type. Warscewiczs.n., Peru. Cajamarca:Nr. Jaen de Bracamoras,fl & young
fr (B, destroyed, photo 12512made by F, F, MO, NY; lectotype, K-Herb. Ben-
tham from D. Hanbury;isolectotype, NY ex W).
Distribution.Known only from the type locality.
80 Flora Neotropica

FIG. 24. Esenbeckia cornuta. A, habit (Warscewicz s.n., type, NY). B, indument of lower side
of leaf (Warscewicz s.n., type, NY). C, flower bud (Warscewicz s.n., type, NY). D, flower (War-
scewicz s.n., type, NY). E, young fruit (Warscewicz s.n., type, K).
Esenbeckia 81

This species strongly resembles Esenbeckia leiocarpa, as Engler(1874a)point-

ed out, but the fruit is differentas is expressed by the epithets. The fruits of E.
leiocarpa are probably larger and provided with apophyses up to 1 or 2 mm
instead of 5 mm. They are hoary and do not become glabrous. The present study
has revealed a sufficientnumberof differentcharactersto distinguishE. cornuta
as a separate species, apart from the different distribution.These can be found
in the shape of leaves and calyx lobes, in the indumentof the lower surface of
the leaves and of the disc. Furthermorethe petals are differentin thickness, and
the ovary of E. cornuta is provided with tubercles (initials of the apophyses?).
The photo of the type in B shows flowers but no fruits. Probably a decayed
(?) fruit once existed, but was destroyed. ThereforeI have selected a lectotype
out of the isotypes. This has young fruits on which the characteristichorns are
already evident.
16. EsenbeckiaoliganthaKaastra,Acta Bot. Neerl. 26: 475, t. 3. 1977. Fig. 25.
Shrub or tree 1.3-3 m tall, appressed-pubescentwith creamish hairs 0.05-0.4
mm long; branchlets 3-7 mm in diam., grayish-brown,reddish-browntowards
the tip, hoary, becomingglabrous,the elevated marginof young leaf scars densely
pubescent. Leaves alternate,simple;petiole slightlyflattenedadaxiallyand some-
times subcanaliculate,2-15 mm long and ca. 1 mm thick, the indumentlike that
of young branchlets; blade elliptic or obovate, 4-11 x 2.5-8.5 cm, rounded to
obtuse and unequal at base, obtuse or subacuminateat apex with the very tip
emarginateor mucronate,marginplane or subrevolute, the blade subcoriaceous,
green, somewhat more yellowish beneath, shining, (sparsely) appressed-pubes-
cent on both sides, venation ? camptodromous,costa plane above and developing
a ridge. Inflorescences terminal,leafy few-floweredpanicles consisting of erect,
subpaniculateaxillary partialinflorescences 1-2 x to 1 cm, growing to ca. 4 cm
long when still flowering, in fruit to 1.5-7 cm long, appressed-pubescentwith
hairs 0.1 mm long; side-branchletsalternate;bracts caducous, to 0.3 mm long;
pedicels 2-4 mm long; bractlet 1 or obsolete. Flowers ca. 5-6.5 mm in diam., 5-
6(-7)-merous; calyx lobes separate, broadly triangular,0.7-0.9 x 0.5-0.7 mm,
acutish to acuminate, coriaceous, appressed-pubescentthroughoutbelow and at
base above with hairs 0.1 mm long, nerves parallel and branchingtowards the
margin;petals deciduous, valvate, widely spreading,ovate-elliptic, 2.5-2.8 x 1-
1.7 mm, acuminate at apex with tip uncinately inflexed through 0.4-0.5 mm,
coriaceous, thicker towards the tip, yellow, papillose above especially towards
the margin, sparsely strigillose below, venation parallel and branchingtowards
the tip, the median nerve thicker;filamentspersistent, subulate, flattenedproxi-
mally, 1.5-1.7 mm long and 0.2-0.3 mm thick, glabrous;anthers broadly heart-
shaped, ca. 0.8-0.9 x 0.8 mm including the mucronatetip 0.05 mm, papillose;
disc annular,5-6(-7)-lobed, 1.5-1.7 mm in diam., each lobe lobulatewith shallow
grooves, ca. 0.5-0.6 mm high (not higher than the ovary), with thickened upper
margin, glabrous, with glands; carpels adnate to the disc, connate, 0.8-1.2 mm
high, provided with an apical apophysis ca. 0.2 mm high and chargedwith gland-
like tubercles ca. 0.1 mm, subglabrous;style inserted in the upper part of the
carpels, ca. 0.7 mm long and 0.2 mm thick, projectingby 0.5 mm beyond the
apophyses, glabrous; stigma capitate, ca. 0.2 x 0.2 mm. Partialinfructescences
several, each in the axil of a leaf, with 1-3 fruits each; fruits depressed, stellately
(4-)5-6-lobed, 1.4-1.9 x 2.5-3.2 cm when still closed, hoary with appressedhairs
0.1-0.2 mm; loculi obsoletely tuberculate,providedwith a dorsal bluntapophysis
to 3 mm ?1/3 from the tip, septicidallydehiscent from 3 mm above the base to I
mm from the tip; seeds (only 1, immature,seen) ca. 10 x 7 mm; embryos 2.
82 Flora Neotropica

FIG. 25. Esenbeckia oligantha. A habit Irwin et al. 31543, U). B, inflorescence Anderson et al.
36546, type, U). C, flower (Anderson et al. 36546, type, U). D, fruit (Anderson et al. 36545, U).
Esenbeckia 83

Type. Anderson et al. 36546, Brazil. Bahia: Espigao Mestre, 23 km W of Bar-

reiras, 3 Mar 1972, fl (holotype, U ex NY; isotypes, NY 9x to be distributed).
Distribution.Espigao Mestre in western Bahia, Brazil. As large parts of Bahia
are not yet explored botanically, more collections are to be expected. Rocky or
sandy cerrado and cerradao;alt. 600-700 m. Collected in flower and in fruit in
March. Fig. 20B.
Specimens examined. BRAZIL. Bahia: Espigao Mestre, type locality, Anderson et al. 36545 fr
(NY 8x, U); basin of Rio das Ondas, upper slopes of Espigao Mestre, 32 km W of Barreiras, Irwin et al.
31543 fr (NY 7x, U).
This species has much in common with Esenbeckia leiocarpa and E. cornuta
but is easily distinguished.Strikingdifferencesare: the indumentwhich is cream-
ish with the hairs less shining, not hyaline; the broaderleaves with rounded or
obtuse base; the quite different inflorescence and aestivation. The disc and the
ovary show charactersin common with one or both other species. The fruits are
very differentfrom those of E. leiocarpa but are probablymore similarto those
of E. cornuta.

lb. Esenbeckiasubgen. Esenbeckiasect. Esenbeckia.

Esenbeckia sect. Polembryum Grisebach, Fl. Brit. W. I. 135. 1859. Type species. Esenbeckia
castanocarpa Grisebach (=E. pilocarpoides Kunth).
Esenbeckia sect. Hymenopetalae Engler in Martius, Fl. bras. 12(2): 140. 1874. Type species.
Esenbeckia pilocarpoides Kunth.

Petals purplish or violet, in Esenbeckia alata faintly so, but yellowish in E.

pilocarpoides; filaments with a basal appendageon the back which reaches be-
yond the grooves of the disc.
Type species. Esenbeckia pilocarpoides Kunth.
Distribution.This section has its main area of distributionin northwesternS
America, especially in Peru and Colombia. Esenbeckiapilocarpoides is the only
species which, with its subsp. maurioides, ranges furtherafield. Fig. 4A.
17. EsenbeckiapilocarpoidesKunth in Humboldt, Bonpland and Kunth, Nov.
gen. sp. 7: 192(folio ed., = p. 248 quartoed. and reprintCramer1963),
t. 655. 25 Apr 1825;Kunth, Syn. pl. 4: 251. 16Jan 1826;Schott, Rutac.
9, t. 5. 1834;Englerin Martius,Fl. bras. 12(2): 144. 1874;Urban, Bot.
Jahrb.Syst. 21: 554. 1896, syn. uno excl.; Wilson, in North Amer. Fl.
25: 202. 1911, syn. uno excl.; Pittier, Trab. Mus. Com. Venez. 7: 339.
1930;Macbride,Field Mus. Nat. Hist., Bot. Ser. 13: 673. 1949.
Shrubor smalltree 2-5(-7.5) m tall with trunk4-5(-12?) cm in diam.;branchlets
2-4 mm in diam., dark grayish-brown,grayish-greenwhen young, glabrousbut
appressed-puberulouswhen young with hairs to 1.5 mm long. Leaves alternate
or subopposite, often crowded at tips of branches, l(-3)-foliolate, with sessile
leaflets; petiole ? terete, (slightly) winged, 8-40(-60) mm long, the base slightly
tumid or not but rather thick in fast-grown shoots, tubercle usually absent, gla-
brous or puberulouswith straighthairs 0.1-0.2 mm long; leaflet blade rathervari-
able in outline, (narrowly)ellptic to (narrowly)(sub)obovate,6-25.5 x 2.1-8 cm,
(mainly shortly) attenuateor occasionally subcuneateat base, acuminateat apex
with obtuse acumen 4-15(-30) mm long, or occasionally retuse, the marginsub-
revolute, the blade chartaceous-subcoriaceous,dull and deep green above when
fresh, paler beneath, glabrouson both sides, rarely subglabrousat the basal part
of the midvein, venation brochydodromousto sometimes slightly camptodro-
84 Flora Neotropica

mous, prominent on both sides, midvein prominent above. Inflorescences ter-

minalor axillarynear tips of the branchlets,erect, narrowlypaniculate,not longer
than the leaves, ca. 5-16 x 2-8 cm, densely minutelypubescent with hairs 0.1-
0.5 mm long; side-branchletsalternate, 1-7 mm long; bracts (very broadly)ovate,
concave to cucullate, 0.6-2.5 x 0.5-1.2(-2) mm, subglabrous;pedicels 1-4 mm
long; bractlets 2, subopposite or alternate, one or both sometimes subtendinga
secondary pedicel. Flowers 5-8 mm in diam., scentless; calyx lobes quincuncial,
adnate to the petals at very base, subcircular, 1.1-1.5 x 1.2-2 mm, rounded or
obtuse at apex, softly coriaceous, ciliate or fimbriate,facially glabrous, with 8-
10 parallel nerves branchinghere and there towards the tip; petals persistent,
imbricate,mostly quincuncial,very widely spreading,elliptic to subcircular,2.6-
4.4 x 1.8-3.6 mm, roundishor obtuse, the marginsomewhat wavy, chartaceous
or subcoriaceous, pale yellow to white, papillose above, glabrous beneath, the
8-10 nerves like those of the calyx lobes but ? diverging;filamentssubulate,0.7-
1.3 mm, glabrous,furnishedat base with an abaxialsubglobularknob 0.1-0.2 mm
high and sometimes protrudingradiallyfrom the disc; anthers heart-shaped,ca.
0.4-0.5 mm long, including a mucronate tip 0.1 mm easily seen in bud only,
papillose especially dorsally; disc annular, 5-partedby deep grooves to /2 of its
thickness, 2-3 mm in diam., fleshy, the parts +2-lobed, 0.4-0.5 mm high and 0.3-
0.5 mm thick, chargedwith some glands and also sometimes with some tubercles,
carpels adnate to the disc and connate in the lower half, globose in the upper
half, ca. 0.3 mm high, furnished with numerous ovoid protuberancesto 0.1-0.2
mm long and densely beset with spreading, thick hairs to 0.2 mm long; style
inserted ca. midway on the carpels, 0.5-1 mm long (nearly completely free), 0.1-
0.3 mm thick, glabrous;stigma capitate, slightly 5-lobed, 0.2-0.4 x 0.2-0.4 mm.
Fruits depressed, stellately-lobed,(10-)15-20 x 20-30 mm, with glands, sparsely
appressed-pubescentwith hairs 0.1 mm long; loculi provided 2/3 from the base
with an apophysis, muricateor sparsely muricatewith tubercles or occasionally
with blunt spines to 5 mm long, weakly curved and shrunken when mature,
dehiscent septicidally from apex down to 3-4 mm above the base; seed 1 per
loculus, obliquely tear-shaped,8.5-11 x 5-6.5 mm, the ventral side flattenedat
the base, the apex with a short, curved beak, the dull brown testa linear-collicu-
late with interspaces of ca. 0.1-0.2 mm, but particularlynear the chalaza also
roundish-colliculatewith interspacesof the same length, those shiningand darker
than the rest of the testa; chalazal area ovate, 2.5-4 x 1.8-2.2 mm, darkbrown-
black; hilum 0.5-1.2 mm broad; micropyle with a dark brown-blackcaruncle;
embryo 1 or more, usually 2, 1 much taller than the other(s), cotyledons unequal,
thick, with 2 ears and a conical radicle projecting beyond them, plumule very
small.
Distribution.Martinique(?), Colombia,Venezuela, Trinidadand Tobago, Guy-
ana, Surinam,and SW Brazil; a subspecies in coastal Brazil (Maranhao,Bahia,
and Espirito Santo).
Key to the Subspecies of Esenbeckia pilocarpoides
1. Leaves 1-foliolate; petioles shortly eared; apex of leaflets acuminate, straight or laterally
curved. 17a. subsp. pilocarpoides.
1. Leaves varying from 3- to 1-foliolate; petioles long-eared, the ears to 3 or 4 mm; apex of
leaflets often rostrate-caudate, laterally curved. 17b. subsp. maurioides.

17a. Esenbeckia pilocarpoides Kunth subsp. pilocarpoides. Fig. 27.

Pilocarpus humboldtii Sprengel in Syst. veg. ed. 16. 4(2): 126. 1827, nom. illeg. ex Art. 55 ICBN;
G. Don, Gen. hist. 1: 795. 1831, pro syn.; Holmes, Pharm. J. Trans. ser. 4. 1: 541. 1895.
Esenbeckia castanocarpa Grisebach, Fl. Brit. W. I. 135. 1859; Engler in Martius, Fl. bras. 12(2):
 Esenbeckia 85

144. 1874;Urban, Bot. Jahrb.Syst. 21: 554. 1896,pro syn. TYPE. Sieber34, Trinidad,1825,
fl & fr (lectotype, K; isolectotypes, C (s.n.), E, G-Herb. Moricand,G-Herb. DC-Delessert,
L 2x, LE 2x, M ex Landishuth,W drawnin Schott, Rutac. t. 5. 1834).
Esenbeckiavenezuelensis Englerin Englerand Prantl,Nat. Pflanzenf.3(4): 159. Mar 1896, syn.
nov.; Engler, Bot. Jahrb. Syst. 21. Beibl. 54: 27. 12 May 1896;Pittier, Trab. Mus. Corn.
Venez. 7: 339. 1930; Engler in Engler and Prantl, Nat. Pflanzenf. (ed. 2) 19a: 282. 1931.
TYPE. Otto 1015, Venezuela. Bolivar:Nr. Upata, 24 Nov 1840,fl (holotype, B, destroyed,
photo 12518made by F, F, MO).
Leaves 1-foliolate; petiole slightly winged, the wings shortly eared at tip and
usually 1-2 mm broad; apex of leaflet acuminate 4-15(-30) mm long, the very tip
obtuse or sometimes retuse.
Type. Humboldt & Bonpland 276, Venezuela. Sucre: Nr. Cumana, near Que-
tepe, early Sep 1799, fl (holotype, P-Herb. Humboldt & Bonpland, photo 36820
made by F, F, MO).
Distribution. As given for the species; in Brazil only in Para (not mapped) and
Mato Grosso. Fig. 26A. Light forest on poor rocky soil, exposed granitic out-
crops, etc; alt. 0-1100 m. Flowering May-Jun, in Venezuela chiefly Jul-Nov.
Specimens examined. WINDWARDISLANDS. Martinique:Plee 652 fl (P, P-type Herb. Baillon;
cultivated?).
COLOMBIA.Cauca:Goudot2 (Apr 1844)st (P).
VENEZUELA. Trujillo: Pittier 5799 bud (VEN), 10823 fl (NY). Falc6n: Lasser & Foldats 3027 fl
(VEN); Pittier 6377 fl (G, NY, US, VEN). Lara: Funck & Schlim 675 fl (G 3x, MPU, P), 678 bis fl
(G, P); Mocquerys s.n. (1893-1894) fl & fr (P 3x); R. F. Smith V 886 fl & fr (VEN). Yaracuy: Agostini
et al. 1750 fl (U, VEN 2x); Bernardi 6925 fr (VEN); Brito 14 fr (VEN). Carabobo: Pittier 8961 fl (G,
GH, NY, US, VEN). Aragua: Pittier & Nakchenovick 15554 fl (VEN). Nueva Esparta: Benitez de
Rojas 1633 fr (U); Gines 3355 fl (US), 3770 fl (US); Miller & Johnston 224 fl (A, BM, GH, MO, NY,
US). Sucre: Aristeguieta 5370 fl (VEN); Aristeguieta & Agostini 4736 fl (K, NY, U, US, VEN);
Broadway 384 fl (GH, NY, US); Funck 26 fl (BR 4x, G, NY); Steyermark 62839 fr (F, NY, US,
VEN), 107802fl (VEN). Bolivar:Blanco 376 fr (NY, VEN); Otto 1015fl (photo, F, MO);Steyermark
57682 fl (F), 87967 fr (GH, NY, U, US, VEN), 88198 fr (NY, US, VEN). State unknown:El Motin:
Croizat 592 fl (NY), 592A fr (NY 2x).
TRINIDADAND TOBAGO.Trinidad.St. George: Britton2716 fr (GH, NY, US); Brittonet al.
2161 fr (GH, K, NY, TRIN 2x, US); Criiger s.n. (Bocas Islands) fl (GH); Finlay TRIN 3064 fl (F,
TRIN, US); Purdie 65 fl (K). Caroni: Arr. Hart TRIN 1324 fr (K, TRIN). Victoria: Marshall TRIN
12248 fr (K 3x, NY, TRIN); Swabey TRIN 12525 fr (K, TRIN). County unknown: Criiger s.n. (28.47)
fl (K); Finlay TRIN 3063 fl, fr (F, TRIN, US); Arr. Hart TRIN 3063 fr (TRIN); Purdie 64 fl, fr (K),
s.n. fl, fr (CGE, GH, GOET, P). Tobago:Broadway3843 fl (A, BM, C, G 3x, K 2x, M, MO 2x, NY
2x, P 2x, S 2x, U), 4192 fl (L, Z), 4257 fr (BM, E, F, G 2x, NY 2x). Islandunknown:R. 0. Williams
TRIN 12169 fl (TRIN).
GUYANA. Essequibo: A. C. Smith 2361 fr (A, K, NY, US). County unknown: Schomburgk 153.5
fl (K).
SURINAM. Nickerie: Irwin et al. (in Maguire) 55309 fr (K, NY, S, US); Stahel 361 fl (U). Ma-
rowijne:Cowan & Lindeman39049-Afl (NY), 39194 fl (F, GH, K, NY, P, RB, U, US).
BRAZIL. Para: Berg et al. 749 fr (U). Mato Grosso: Berg et al. (in Prance) 18590 fl (U).

Local names and use. Trinidad: gasparee or gaspari(llo) (Purdie 64, 65; Schwa-
bey 12525; Arr. Hart TRIN 1324). This species supplied good walking sticks while
the "swizzle stick largely originates from this tree" (Purdie 64, Marshall 12248,
Williams 12169).
Esenbeckia venezuelensis appears to be synonymous with the present species.
According to the description of Engler there is a single, slight difference in the
petioles which should be densely pilose instead of puberulous. The type as far as
it can be observed on the photo resembles E. pilocarpoides (the holotype is lost).
Steyermark 57682, collected near the type locality, identified by Steyermark as
E. venezuelensis, belongs to E. pilocarpoides too, as does Blanco 376 from about
the same place. Steyermark 57682 has rather strongly tumid petiole bases, such
as occasionally occur, especially in fast-grown shoots, in this species (A. C. Smith
2361; Broadway 3843 (MO)).
86 Flora Neotropica

, i? z.j * E.pilocorpoidessubsp.pilocarpoides
\
| o? I * E.pilocarpoidessubsp.maurioides
t o E.amozonico
k.<, vL . .
.T'' . o E. ? QlatQi
.0.. _ E. warscewiczii
FIG. 26 Dsrbto o s spce E. scrotiformis
of-J

* E febrifuga7
* E densiflora ()
v E hieronymi

o... ..2 .

FIG. 26. Distribution of some species of Esenbeckia.

Esenbeckia 87

t 0.2

FIG. 27. Esenbeckia pilocarpoides subsp. pilocarpoides. A, habit with analyses (Sieber 34, W,
originallydrawnby Obererin Schott, Rutac. t. 5. 1834).B, papilloseupperside of petal (Pittier10823,
NY). C, fruit (Britton et al. 2161, NY). D, seed (Britton et al. 2161, NY). E, ovule with obturator
(Pittier 8961, NY). F, developing seed (Steyermark 87967, NY).
88 Flora Neotropica

The fruits of A. C. Smith 2361, Croizat 592 A, and Steyermark 87967 have a
somewhatechinate appearance,the tuberclesbeing replacedby blunt spines. The
last collection differs further from all others in having one 2-foliolate leaf and
wings of the petioles only ca. 0.4 mm broad.

17b. EsenbeckiapilocarpoidesKunth subsp. maurioides(Martius)Kaastra, Acta

Bot. Neerl. 26: 477. 1977.
EsenbeckiamaurioidesMartius,Nov. gen. sp. pl. 3: 82, t. 232. 1831;Dietrich, Syn. pl. 1: 831.
1839;Walpers,Repert. 1: 501. 1842.
Colythrummaurioides(Martius)Schott, Rutac. 14. 1834.
Esenbeckiapilocarpoides Kunth var. maurioides (Martius)Engler in Martius,Fl. bras. 12(2):
144. 1874.
Shrub to 6 m tall with trunk 5-8 cm in diam. Leaves varying from 3- to
1-foliolate, the 1-foliolateones frequently 2-partedor 2-cleft, or the leaflets of a
2-foliolate leaf ? connate; petiole provided with wings ca. 0.6-1.3 mm broad
endingin blunt ears 3-4 x 1-1.5 mm or shorter;apex of leaflets acuminate,most-
ly rostrate-caudatewith a usually blunt, laterally curved acumen (2-)7-25 mm
long; lateral leaflets sometimes greatly reduced in size with a slightly unequal
base, curved and with an admedialmidvein.
Type. Martiuss.n., Brazil. Bahia:Above Serrado Mar, nr. Almada(=Ilheus),
Dec (1818), fl (lost; lectotype, Martius,Nov. gen. sp. pl. 3: t. 232 and t. 233 pro
parte. 1831).
Distribution.Brazil, along the coasts of Maranhao,Bahia, and Espirito Santo.
In Maranhaoin high forest on terrafirme. Collected in flower in Apr. Fig. 26A.
Specimens examined. BRAZIL. Maranhao:MaracassumeRiver region, CandidoMendes, Fr6es
(in Krukoff) 1710 fl (A, G 2x, K, MICH, MO, NY, P, S, U); Rio Pindare, between Rapoza and
Monqao,Fr6es (in Krukoff)11666fl & fr (A, F, K, MO, NY, S, U); Island of Sao Luiz, Arutahy,
Fr6es (in Krukoff) 11797 fl (A, F, K, MO, NY, S). Espirito Santo: Itapemirim, Wied-Neuwied s.n.
(Itap(emirim)) fl (BR, M).
Local names and use. Maranhao: boragica (Froes (in Krukoff) 11666, 11797);
pipoca (Froes (in Krukoff)1710). The wood used by Indians in preparingcertain
parts of arrows (Froes (in Krukoff) 11666).
The originallabels of Wied-Neuwieds.n. mention "Itap." After studying Ur-
ban in Fl. bras. 1(1) (1906), several maps, and indexes, I drew the conclusion that
Itap. stands for Itapemirim.AfterwardsI came across the specimen in BR-Herb.
Martius, on which Martiushad attached a label with: "Itapemirim.Communic.
Pr. Vidensis 1829." Itapemirim, situated in coastal Espirito Santo, therefore
seems to be the locality. The collection was not identifiedby Martius, only by
Engler, and the specimen in M also by Schultes. It agrees very well with Martius'
description, and the drawing, t. 232 and 233 pro parte in Martius,Nov. gen. sp.
pl. 3, seems to be based on it. The date of acquisitionby Martiusis in accordance
with the completion of the publication(after 29 June 1829and before ca. March
1831; cf. Stafleu (1967: 297). Therefore it is not impossible that Wied-Neuwied
s.n. actually is the type of this subspecies.
Froes (in Krukoff)11666 shows only long-eared 1-foliolateleaves. The absence
of 2-3-foliolate leaves is perhaps due to incomplete sampling;cf. the specimen
of Froes (in Krukoff)1710 in A, which shows 1-foliolateleaves only, while du-
plicates in other herbariahave 1-3-foliolate leaves. In other characters Fr6es
(in Krukoff)11666 is not different, and the collection belongs therefore to this
subspecies.
Esenbeckia 89

18. EsenbeckiaamazonicaKaastra, Acta Bot. Neerl. 26: 477, t. 4. 1977.Fig. 28.

Esenbeckia cornuta sensu Albuquerque,Acta Amazonica6 (3: Supl.): 23. Sep 1976(received
at Utrecht 29 Nov 1977),non Englerin Martius,syn. nov.
Small tree or shrub3-6 m tall; branchlets3-5 mm in diam., grayish-brown,the
younger ones greenish, glabrousbut younger ones minutelypubescent with hairs
to 0.2 or 0.3 mm long. Leaves alternate, occasionally some subopposite, 1-fo-
liolate, the leaflet sessile; petiole ? terete, narrowlyor rarely strongly winged,
11-50 mm long, with a terminaltubercle ca. 0.2-0.3 mm long, mostly glabrous
or minutely puberuloustowards the tip with hairs 0.1-0.2 mm long, wings at tip
to 0.6(-0.8) mm broad and mostly not auriculate; leaflet blade obovate, 8-
31 x 2.8-12.5 cm, base mostly cuneate or somewhatvery shortlyattenuate,apex
(sub)acuminate,the acumen obtuse or acute and 6-9(-16) mm long, marginsub-
revolute, the blade chartaceous, pale green beneath, glabrous on both sides or
beset towardsthe base with few appressedhairs0.1-0.2 mmlong, venationbroch-
idodromous-camptodromous,midvein usually prominent above. Inflorescences
1-few, terminalor axillary near the tips of branchlets, erect, ca. as long as the
leaves, 14-27 x 6-15 cm, paniculately-branchedwith alternate,long side-branch-
lets, minutely pubescent with hairs to 0.1 mm long; bracts (broadly)ovate, con-
cave-conduplicate, 0.5-3.5 x 0.4-1 mm; pedicels 2-4 mm long; bractlets some-
times subtendingsecondary pedicels. Flowers (7-)8-14 mm in diam.; calyx lobes
quincuncial,very broadly ovate to heart-shaped, 1.1-1.6 x 1.5-2.3 mm, obtuse,
chartaceousto subcoriaceous,violet-tinged,at anthesis subfimbriate,glabrouson
both sides or minutely pubescent below with hairs 0.05-0.1 mm long; petals
persistent, imbricate, very widely spreading, ovate, 4.2-6 x 2-3.5 mm, obtuse,
chartaceous,greenish-whitesuffused with violet or darkred on the upper side at
base and margin, this subundulateand sometimes ciliate, minutely pubescent
above with hairs 0.05-0.1 mm long, glabrous or minutely appressed-pubescent
below with hairs 0.1-0.2 mm, with 6-10 parallel primarynerves; filamentsper-
sistent, subulate, to 1.9 mm long, glabrous, provided with a basal, abaxial, ?
globular appendage protrudingradially from the disc; anthers heart-shaped,in-
cluding a mucro 0.05-0.1 mm, 0.5-0.9 x 0.4-0.8 mm, papillose mainly on the
back; disc annular, 5-parted, 0.5-1 mm high and 2-3.5 mm in diam., each part
slightly2-lobedor not, fleshy, violet, papilloseandbeset with some tuberclesto ca.
0.2 mm; carpels adnate to the disc in their lower half, scarcely connate with each
other, ca. as high as the disc, in the upper half provided with a small radial
apophysis, beset with few or many tubercles to 0.2 mm long, mostly glabrousor
with few spreadinghairs to 0.2 mm long; style inserted near base of the carpels,
0.6-1.1 mm long and ca. 0.2-0.4 mm thick, projectingup to 0.7 mm beyond the
ovary, glabrous;stigma clavate or capitate, shallowly 5-lobed, 0.2-0.3 mm high.
Fruits depressed, stellately 5-lobed, 1.5-2.5 x 1-2.2 cm when closed,
(sub)glabrous,with scattered glands; loculi connate in the lower part only, pro-
vided with an apohysis /2 to % from the base, slightly tuberculate,the nerves of
the exocarp prominenton the outer surface, septicidally dehiscent at first from
some distance above the base to 1 mm from the tip, finallydehiscent loculicidally;
seed (only 1, immatureone studied) 1 per locule, obliquelytear-shaped,ca. 7 x 5
mm, light brown-orange,finely colliculate, chalazal area not observed; embryo
1, with thick unequal cotyledons, punctate, radicle shortly projectingbeyond the
ears, plumulevery small.
Type. Ducke 1266, Brazil. Amazonas: Esperanga,nr. mouth of Rio Javary, 28
Sep 1942, fl & fr (holotype, US-2592220;isotypes, A, F, NY, R 3x, UC).
90 Flora Neotropica

filament (Ducke 1266, type, A). C, fruit (Ducke 1266, type, NY).
Esenbeckia 91

Distribution.Upper basin of the Rio Amazonas in Colombia(Amazonas),Peru

(Loreto, Huanuco), and Brazil (Amazonas). Forest on terra firme to 500 or 1200
m alt. Collected in flower Oct-Nov. Fig. 26A.
Specimens examined. COLOMBIA. Amazonas: Rio Loreto-yacu, Black & Schultes 46-259 fl (F).
PERU. Loreto: Florida, Rio Putumayo, Klug 1990 fl (K, S); Boquer6n Padre Abad, Schunke V.
5818 fr (F 2x). San Martin: Nr. Moyobamba, Zepelacio, Klug 3275 fl (A, F, GH, K, MO, S, US),
3380 fl (A, F, GH, K, MO, S, US). Huanuco: Palo de Acero, Woytkowski 7646 fl (F, K, UC, US).

Local name. Peru, Loreto: barbasco caspi (Schunke V. 5818).

This species is near to Esenbeckiapilocarpoides. Differences are the broader
leaflets, the inflorescencetwice as long and wide, the flowers largerin all respects
and differentin color, and the only slightly tuberculatefruits.
One of two collections cited by AlbuquerqueunderE. cornuta is an isotype of
E. amazonica. The other 1 (or 2?) collection(s) I did not study, but the photo-
graphs and the description clearly indicate that they belong to E. amazonica.
This is shown by the largeleaves and the purplishbase of the petals. Albuquerque
apparentlydid not notice the articulationof the leaves.
19. Esenbeckiaalata (Karsten & Trianain Triana)Triana& Planchon, Ann. Sci.
Nat. Bot. s6r. 5. 14: 306. 1872;Engler in Martius,Fl. bras. 12(2): 145.
1874. (See Addenda note 2). Fig. 29.
Kuala alata Karsten & Triana in Triana, Nuev. jen. 22. 1855; Karsten and Triana in Karsten,
Linnaea 28: 429. 1857 (" 1856"); Muller in Walpers, Ann. 7: 529. 1869.
Kuala laevis Karsten & Triana in Triana, Nuev. jen. 22. 1855; Karsten and Triana in Karsten,
Linnaea 28: 429. 1857 ("1856"); Miiller in Walpers, Ann. 7: 529. 1869, syn. nov. TYPE.
Triana s.n. ("3742; Tocaima, prov. de Bogotd, 600 m"), Colombia. Cundinamarca: Tocaima,
fr (lectotype, G, photo 26407 made by F, F, MO, NY).
Esenbeckia alata (Karsten & Triana in Triana) Triana & Planchon var. laevis (Karsten & Triana
in Triana) Triana & Planchon, Ann. Sci. Nat. Bot. ser. 5. 14: 306. 1872; Engler in Martius,
Fl. bras. 12(2): 145. 1874.

Shrub or small tree to 5 m tall, with hairs 0.2-0.3(-0.4) mm long; branchlets

minutely appressed-pubescent,becoming glabrous, light grayish-brown.Leaves
alternate, 1-3-foliolate, with (sub)sessile leaflets; petiole terete, slightly winged
or not, slightly thickened at base, (1-)2-10 cm long, with a small tubercle, ap-
pressed-pubescent,becoming glabrous, the wings, if any, distally to 0.5-1.0 mm
broad and terminatingin an ear or graduallynarrowingtowards the tip; leaflet
blades (ob)ovate, elliptic, or oblong, (8-)11-23 x (3-)4.5-9.5 cm, (long-)attenuate
at base, especially the smaller lateral ones unequal at base, 6-14 x 3-6 cm,
(slightly) acuminate at apex with the very tip obtuse or emarginateto 10 mm,
margin revolute, the blade subcoriaceous, glabrous on both sides or with few
appressedhairsnearbase, venation camptodromousto slightlybrochidodromous,
midvein and main veins prominentparticularlybeneath. Inflorescences terminal,
branchedat base into 3-5 widely or narrowlypaniculatefew-branched,minutely
pubescent partial inflorescences 8-13 x 2-11 cm and shorter than the leaves;
branches alternate, dropping off when sterile; bracts ovate, to 1.8 x 0.6 mm,
subglabrousor pubescent like the branchletsof the inflorescence but less dense,
with or without a false midrib0.1 mm long on the back; pedicels 2-8 mm long;
bractlets 2, alternate or subopposite. Flowers 8-9.5 mm in diam.; calyx lobes
quincuncial, depressedly ovate to roundish, 1.5-1.7 x 1.5-2.5 mm, rounded to
subobtuseat apex, coriaceous, marginpapery, subglabrous,nerves parallel,often
provided ca. V3 from tip with a false midrib ca. 0.3 mm, petals deciduous or
persistent, cochlear, very widely spreading, elliptic, 3.3-4.5 x 2.3-3.7 mm, ob-
tuse to roundedat tip, papery, greenish-yellowor faintly violet, papillose above,
92 Flora Neotropica

29. Esenbeckia
FIG. .A, alata. habit (Triana s.n., paratype of Kuala laevis, COL- 17077). B,
postfloral flower, with B 1, insertion of filament between the lobes of the disc (Karsten s.n., W). C,
fruit (Killip et al. 38300, US).
Esenbeckia 93

(sub)glabrousbelow, venation actinodromous;filaments inserted on the lower

part of the disc, charged at base with an abaxial appendagegreatly swollen im-
mediately after anthesis, the upper part of the filamentseemingly inserted in an
apical pit of the disc, this appendage0.2 mm long adnate to the disc, postflorally
becoming as high as it, the distal free part of the filamentssubulate, 0.8-0.9 mm
long, glabrous; anthers ovoid, 0.5 x 0.4 mm and 0.3 mm thick, the connective
not protruding;disc cup-shaped,postflorallyconstrictedinto 2 superposedparts,
0.5-0.9 mm high (as high as the ovary), 2.5-2.8 mm in diam., fleshy, the upper
part formingbetween the filaments5 ? flat, slightly pustulate lobes 0.3-0.6 mm
broadand providedwith few tuberclesto 0.1 mm long; carpels adnateto the disc,
0.7-1.0 mm high, postflorallydeveloping a solid apical apophysis ca. 1.2-2 x 1
mm; apophyses and rest of carpels tuberculate, with glands some of which be-
come enlargedinto small tubercles, glabrousor sparsely pilose with hyalinehairs
to 0.1 mm long; style inserted near base of the carpels, 1.6 mm long, free from
the ovary in the upper half, and 0.2 mm thick but thickened towards the tip,
glabrous;stigma capitate, 5-lobed, 0.2 x 0.4-0.5 mm, each lobe provided with a
large yellow gland. Fruits depressed, stellately 5-lobed, 1.5-2.3 x 2.4-3.5 mm
when dehisced, loculi trigonous, dorsally 13-/2 from base provided with a re-
curved horn-likeapophysis 5-8 mm, muricate, wrinkled, glabrous or with some
appressed hairs 0.1-0.2 mm; seed 1 per loculus, obliquely tear-shaped, 9.5-
10.5 x 5.8-8 x 5-6 mm, flattenednearbase, the apex with an (obsoletely) curved
beak to 1 mm, testa dull brown, reticulate-colliculatewith shininginterspaces0.1
mm long; chalazalarea ovate, ca. 2-3.7 mm long, acuminate,somewhatelevated,
black; hilum 0.2-0.6 mm broad; embryo 1 per seed, cotyledons punctate, with
ears 0.5-0.7 mm, the radicle enclosed.
Type. Triana s.n. ("Tocaima, prov. de Bogota, 600 m"), Colombia, Cundi-
namarca,Tocaima, fr (lectotype, W; isolectotype, W).
Distribution.Colombia,in Cundinamarca,Tolima, and Valle del Cauca:valleys
of Rios Magdalenaand Cauca. Collected from 350-800 m alt.; floweringMay and
Dec. Fig. 26A.
Specimens examined. COLOMBIA.Cundinamarca:Tocaima, Archer 3355 st (US), Goudot s.n.
(1844, fl May, fr Aug) fl (P), Karsten s.n. ("Kuala alata") fl/fr (W, paratype?), Killip et al. 38300 fl
& fr (A, COL 2x, US), Triana s.n. (alt. 600 m, no. 3841?), fl/fr & fr (G, paratype), s.n. (alt. 700 m,
no. 3741?), fl (P, paratype?),s.n. (alt. 600 m, no. 3741?), fl/fr(B, destroyed, photo 12511madeby F,
F, MO, NY; E, K, US ex BM, paratypes?),s.n. (withoutdata), fr (COL-Herb.Triana,paratype?),
s.n. (alt. 800 m), fr (K, photo NS 2867 made by NY, NY, paratype?), s.n. (alt. 700 m), fl/fr & fr (BM,
paratype?)(all collections cited so far identified"Kuala alata"; the next collections of Trianawere
named 'Kuala laevis,") s.n. (alt. 600 m, no. 3762?), fr (K-Herb. Hooker, paratype of Kuala laevis),
s.n. (alt. 500 m), fr (K-Herb. Hooker, photo NS 2868 made by NY, NY, paratype of Kuala laevis?),
s.n. (alt. 700 m), fr (MPU, fragment of paratype of Kuala laevis?), s.n. (alt. 700 m, Feb 1854), fl/fr
(COL-Herb. Triana 2x, paratypes of Kuala laevis); Honda, Holton 66 ("815") fl (K, NY), Karsten
s.n. (Oct 1872)fr (GOET,LE, NY, W). Tolima:Goudot 1 (Feb 1844)fr (P). Valle del Cauca:Garcia-
Barriga4312 st (COL, US). Locality unknown:Mutis4033 fr (US).
Local names and uses. Cualaand cuala-cuala(several mentions);coya (Goudot
s.n., 1844). The bark is used as insecticide (Garcia-Barriga 4312).
This species shows many similaritieswith Esenbeckiapilocarpoides. However,
there are good differentialcharacters, e.g. the 3-foliolate leaves, the very large
filament-appendages,and the fruit with apophyses inserted lower than in other
species.
Most of the originalmaterialof Kuala laevis has the petioles narrowlywinged.
There are also collections, e.g. Killip et al. 38300, which have some petioles
winged and others not. No differences other than the wings could be found, so
94 Flora Neotropica

there is no reason to maintainvar. laevis. Trianaand Planchon(1872:306) already

had their doubts on this question.
I disagree with Sprague (1934: 394) as to the date of effective publication of
the protologue. The cover of the book bears the imprint: "Imprentadel Neo-
Granadino-1855"; the effective date of publicationis 1855, not 1854as given on
the title page. The assignation of the date 1854 by Karsten (1857: 276) and by
Trianaand Planchon (1872) is incorrect.
Since both Karsten (1857) and Triana and Planchon (1872) give Karsten and
Trianaas authors, this may be assumed to be correct, as Sprague(1934)already
supposed. The sheet in Triana's own herbarium(now COL) with the inscription
"Kuala laevis Karst. et Tr." is in accordance with these facts.
In the originalpublicationno types were designated,only the locality indication
as "provincia Tequendama [=western part of Cundinamarca] crescent alt. 600
metri." Selection of the types is difficult because several specimens were col-
lected at the type locality. Some of these came to the herbariavia Karsten, others
via Triana (Karsten accompanied Triana in Feb 1854 to La Mesa, Anapoima,
Tocaima, Ibague and other places in Tequendama(see Dugand, 1944).Karsten's
labels do not always mention a collecting date. Karsten apparentlyvisited the
same area at least in Oct 1872(accordingto a herbariumsheet in GOET;cf. also
Chaudhri et al. in Index Herbariorum II, Collectors, I-L, 1972). It cannot be
concluded with certainty, therefore, whether some of the Karsten specimens
belong to the originalmaterialor not.
Concerningthe Triana specimens, most of these bear a label "Voyage de J.
Triana 1851-1857, Tocaima (some also with "Le Magdalena") prov. de Bogota."
Trianahowever visited Tocaima in Jan 1853, Feb 1854 (together with Karsten),
Jul 1854, and Sep 1855. If he gatheredE. alata in 1855, those specimens do not
belong to the originalmaterial.Withouta precise date there is no way of telling
which specimens are original.
In selecting the lectotype I chose from among Trianaspecimens of which the
label gives as altitude "600 metr." Several of these bear the names "Colythrum
alatum. Kuala alata Tr. et Karst.," but as Colythrum was not mentioned in the
protologue I suppose these labels were written later on. Therefore I chose as
lectotype one of the two specimens in W labeled "Kuala alata Tr. et Karst.
Tocaima prov. de Bogota, hauteur 600 metr.," a fruiting specimen.

20. Esenbeckia warscewiczii Engler in Martius, Fl. bras. 12(2): 148. 1874; Mac--
bride, Field Mus. Nat. Hist., Bot. Ser. 13: 674. 1949. Fig. 30.
Esenbeckia venulosa Macbride, Candollea 5: 376. 1934; Macbride, Field Mus. Nat. Hist., Bot.
Ser. 13: 674. 1949, syn. nov. TYPE. Weberbauer 6584, Peru. Junin: Rio Mantaro, Huancayo,
rd. to Huachicua, 16 Apr 1913, fl (holotype, F-629065, photo 49591 made by F, F; fragment
of holotype, G; isotypes, MOL (n.v.), S, US).
Esenbeckia dielsiana G. M. Schulze in Diels, Biblioth. Bot. 29 (Heft 116): 100. 1937; Macbride,
Field Mus. Nat. Hist., Bot. Ser. 13: 675. 1949, syn. nov. TYPE. Diels 1164, Ecuador.
Chimborazo: Cafion of Rio Chanchan, above Huigra, ca. 1400 m, Sep 1933, fl & fr (B.
destroyed, n.v.).

Shrub or tree-like shrub 3-4 m tall; branchlets 2-6 mm in diam., densely se-
riceous-pilose with creamy hairs to 0.8 mm long, becoming less densely pilose,
grayish-brown,young tips cream to light ochreous; leaf scars on young shoots
surroundedby centrifugal,straight,creamy hairs but becoming glabrous. Leaves
alternate, 3-foliolate, petiolulate; petiole semiterete, the upper side flattened or
(widely) canaliculate, (1.5-)3-8 cm long, densely pilose like the branchlets or
sparsely so, tubercle or triangularappendageoften present; petiolule of terminal
Esenbeckia 95

I.

FIG. 30. Esenbeckia warscewiczii (Camp E 2976, NY). A, habit. B, indument of lower side of
leaf. C, flower with Cl, filament. D, fruit. E, seed.
96 Flora Neotropica

leaflet (0-)3-7 mm, that of lateral leaflets to 2 mm long, not separatedfrom the
decurrentbase of the leaflet; leaflet blades (narrowly)obovate to elliptic, that of
the terminal leaflet 5.5-15 x 2-6.7 cm, attenuate and slightly unequal at base,
those of lateralleaflets 3.5-12 x 1.7-6 cm, shortly attenuateand stronglyunequal
at base, apex subacute to (sub)obtuse, or occasionally emarginate,rarelyround-
ed, and often slightly acuminate, the margin (sub)revolute, the blade subcoria-
ceous when mature, dull, sometimes darker above than beneath, glabrous to
sparsely appressed-pubescentabove with hairs to 0.6 mm long, spreading-pubes-.
cent towards the base or only in the proximal part of the midvein, subglabrous
to densely pubescent below with spreading hairs to 0.6 mm long, sometimes
minutely strigillose, the midvein and principal veins bearded or pilose with
spreadinghairs, venation camptodromous,midvein plane or slightly impressed
above, lateral veins prominentbeneath. Inflorescences terminal, erect, racemi-
form-paniculate,not longer than the leaves, to 13 x 4(-6) cm, few-flowered.,
densely or sparsely pilose with spreadingcreamy-tawnyhairs to 0.4 mm long;
side-branchletsalternate;bracts broadly triangular,concave, to 1.8(-2.5) x 1.3
mm, beset with few appressed hairs to 0.2 mm long above and with more hairs
below; pedicels ca. 1-2.5 mm long; bractlets up to 2, frequently subtendingsec--
ondary pedicels, occasionally obsolete. Flowers (9-)10-13.5 mm in diam.; calyx
lobes quincuncial,heart-shaped,1.1-1.5(-1.7) x (1.3-) 1.6-2 mm, acuteto obtuse, or
rounded at apex, thick with membranousmargin,glabrous above, minutely pu-
bescent to glabrous beneath with appressed, whitish hairs to 0.2 mm long, the
margin glabrous; nerves parallel and branchingtowards tip; petals deciduous,
cochlear, widely spreading,adnate at base to the disc, elliptic to subovate, 4.5--
6.5 x 2.5-3.6 mm, obtuse, subcoriaceous, rarely thick-papery,dark violet or ni--
grescent-maroonwhen fresh, when boiled tinged with some violet or rose but the
semi-transparentmargin creamy- to rose-tinged, minutely velvety above with
hairs 0.05-0.1 mm long, minutely strigillose below with whitish hairs to 0.3 mm
or (sub)glabrous,venation actinodromouswith median nerve a little thicker;fil-
aments deciduous, insertedjust below the carpels on the level of the calyx lobes,,
adnate at base to the disc, 1.7-2.7 mm long and 0.2-0.4 mm thick, glabrous, the
basal part consisting of a ? oblong swelling 0.8-1.3 x 0.4-1 mm; anthers heart-
shaped, 0.9-1(-1.3) x 0.9-1 mm and 0.4 mm thick includingthe protrudingtip of
the connective 0.1-0.2 mm, papillose; disc cup-shaped, the lower part adnate to
the ovary, with 5 incurvationsup to half its thickness, the lobes plicate or not.,
the free part ca. 0.5-1 mm high and 0.2-0.6 mm thick, 2-2.9 mm in diam., fleshy.,
pustulate, sparsely tuberculate;carpels almost completely adnate to the disc ex-
cept for the 5 dorso-apical apophyses, immersed around the base of the style.,
0.3-1 mm high excluding apophyses 0.2-0.4 mm high and at anthesis not yel
present or barely visible, the apophyses provided with glandular,(sub)glabrous
protuberancesto 0.1-0.3 mm; style inserted at the same level as the free parts of
disc and petals, just above the immersed ovary, thickened towards the tip, 0.7-.
1.4 mm long and 0.1-0.4 mm thick, extended to 1.9 mm after flowering,glabrous:
stigma clavate to capitate, +5-lobed, 0.2-0.3 x 0.2-0.6 mm. Fruits depressed.
1.5-2 x 2.5-4 cm, stellately 5-lobed with large dorsal horn-likeapophyses ca. 6
mm long inserted just above middle of the loculi, these rounded at base, the
nerves of the exocarp observable externally, ? wrinkled, slightly muricate,with
numerous dark glands, glabrous, septicidally dehiscent, but not up to the axis.
and to 2/9 from base; seeds I or 2 per loculus, obliquely tear-shaped, ventrally
flattened near the base, 10.5-13.5 x 6-8 mm, testa dark chestnut-brown,retic-
ulate with flattened,linearinterspacesca. 0.25 mm; chalazalarea broadlyspathu-
late, ca. 3.5-4.5 x 3-3.5 mm, black;hilum0.6-1 mm broad;embryo 1, cotyledons
Esenbeckia 97

unequal, with ears to 2 mm, radicle projecting ? 1 mm beyond the ears, plumule
2-leafed, 0.4 mm long.
Type. Warscewiczs.n., N Peru, nr. village of Sonda, fr (B, destroyed, photo
12519 made by F, F, MO).
Distribution.Ecuadorand Peru. Scrubchaparralwith a few seepages and small
swamps; deciduous woods or evergreen forest; alt. 800-2500 m. Collected in
flower chiefly Feb-May. Fig. 26A.
Specimens examined. ECUADOR. Chimborazo:Nr. Huigra, Cafion of Rio Chanchan, Camp
E-2976fl & fr (F, G 2x, GH, K 2x, LE, M, NY 2x, P, S, UC, US, W); nr. Huigra,Hda. de Licay,
Rose & Rose 22197 fl & fr (GH, NY, US).
PERU. Tumbes:Tumbes, E of Hda. Chicama, Weberbauer7645 fl (F 2x, K, S, US). Cajamarca:
Jaen, SE of San Felipe, Weberbauer7110 fl & fr (F, G, GH, US). Amazonas:Chachapoyas,Matthews
3045 fl & fr (BM 3x, CGE, E, G 2x, G (s.n.), GH, K 2x, K (s.n.), S). Junin:Rio Mantaro,Huancayo,
Weberbauer6584 fl (F, photo, F; G, S, US).
Local name. Peru, Cajamarca: angohuara (Weberbauer 7110).
The hair-cover of Esenbeckia warscewiczii varies from dense to sparse. The
leaves are densely pubescent to nearly glabrousexcept dorsally along all nerves
or the midvein only. The calyx lobes are pubescent to nearly glabrousbelow.
Esenbeckia venulosa differs in the vestigial indument. The leaves in bud are
beset with long hairs but become glabrous when mature except for the midvein
below. Because of the really variableindumentin E. warscewicziiI see no reason
for maintainingE. venulosa as a separate taxon. It should be mentioned that
Macbridehimself alreadydoubtedwhetherhis species mightnot be identicalwith
E. warscewicziibut the flowers of the latter were unknown(Macbride,1949:674).
Esenbeckia dielsiana agrees well with E. warscewiczii, a species not known to
Schulze. Macbride(1949: 675) assumed that the fruits of E. dielsiana should be
ecornute, but this does not appearfrom the description.At the type locality Camp
made another collection in 1945.
The bases of the leaflets of E. warscewicziiare rarelynot separated,the leaflets
seemingly basally connate.

21. EsenbeckiascrotiformisKaastra,Acta Bot. Neerl. 26: 479, t. 5. 1977.

Fig. 3 1.
Shrub or tree? with creamishhairs to 0.2(-0.4) mm long; branchletsca. 3 mm
in diam., light grayish, shining, minutely appressed-pubescent,becoming gla-
brous. Leaves alternate, 3-foliolate, with stalked or subsessile leaflets; petiole
semiterete, adaxially canaliculate, 4-9 cm long, minutely pubescent; petiolules
not sharply separated from base of the leaflets, 0.5-10 mm long; leaflet blades
obovate or elliptic, 6.5-19 x 2.5-9 cm, attenuate, the terminal one slightly un-
equal and the lateral ones strongly unequal at base, shortly acuminate at apex
with the very tip emarginateor retuse, marginrevolute, the blade chartaceous,
dull, darker above than beneath, minutely puberulouson both sides mainly on
the nerves, venation brochidodromous,midveinplane above, lateralveins prom-
inent beneath. Inflorescencesterminal,paniculate,consisting of up to 3 spreading
or erect partialinflorescences shorterthan the leaves up to ca. 9 x 5 cm, branch-
lets few-flowered, obtuse-angled,minutelypubescent with hairs to 0.4 mm long;
side-branchlets alternate; bracts triangular,concave, ca. 0.6-1 x 0.3-0.6 mm,
with hairs shorter than on the branchlets; pedicels 2-5 cm long; bractlets 1-2,
obsolete, sometimes subtendingsecondary pedicels. Flowers ca. 9.5-12 mm in
diam., protandrous;calyx lobes quincuncial, heart-shaped,0.8-1 x 1-1.3 mm,
obtuse at apex, mostly membranous,coriaceous towards base, glabrous above,
98 Flora Neotropica

FIG. 31. Esenbeckia scrotiformis (Ule 9501, type, G). A, habit. B, flower bud with two petals
removed. C, flower.
Esenbeckia 99

subglabrousbelow, nerves paralleland branchingtowardstip; petals valvate with

the inner margininduplicate, ovate, 3.5-5.5 x 2.2-3 mm, of variable breadthin
the same flower, acuminate at very tip, mostly coriaceous but thinner centrally
at base, dark purplish above, greenish below, except centrally at base densely
villous above with shining lilac hairs to I mm long, the hairs unicellular and
consolidated with longitudinal thickenings, glabrous beneath, venation clado-
dromous; filaments adnate to the disc at base, ca. 2 mm long, pale creamish,
glabrous, with a double, abaxial swelling 0.5 mm in diam. at base, gradually
diminishingin width to subulateat apex; anthersdouble-ellipsoidal,ca. 0.7 x 0.6
mm, pale creamish, 1 or 2 smaller than the others, glabrous; disc cup-shaped,
proximallyadnate to the carpels, deeply cleft into five 2-lobed parts, ca. 1.5 mm
high, the free part 0.6-0.7 mm high, ca. 3 mm in diam., fleshy, creamish-yellow,
with glands, papillose; carpels raised on a gynophore 0.1 mm high, completely
immersedin the lower part of the disc and adnate to it, connate, ca. 0.4 mm high
and provided with an apical apophysis 0.2 mm charged with glandulartubercles
of 0.1 mm, glabrous; style inserted on the ovary among the apophyses, ca. 0.7
mm long and 0.1-0.2 mm thick, projecting 0.4 mm beyond the apophyses, the
free part elongatingto ca. 1.2 mm long after anthesis, creamish,glabrous;stigma
capitate, obsoletely lobed, in the male phase ca. 0.2 x 0.2 mm. Fruits unknown.
Type. Ule 9501, Brazil. Acre: SeringalS. Francisco (nr. Rio Acre, above Ina-
pari), Sep 1911, fl (holotype, G; isotypes, K, L).
Distribution.Known only from the type locality. Fig. 26A.
This species is distinct from all others in Esenbeckia by its villous corolla with
valvate aestivation. It takes its name from the resemblance of the filaments to
the genital parts of the human male.

II. Esenbeckiasubgen. OppositifoliaKaastra, Acta Bot. Neerl. 26: 479. 1977.

Leaves (sub)opposite. Inflorescences terminal with side-branchlets
(sub)opposite. Flowers protandrous;calyx lobes provided with a false midribof
glandulartissue; petals imbricateto valvate, nervation camptodromoustending
to become supra-basallyactinodromousor parallel;anthers attached 1/3-2/5(-1/2)
from base. Seeds mostly 2 per loculus; chalazal area dark brown; granularen-
dosperm remainspresent, as far as known.
Type species. Esenbeckiafebrifuga (Saint-Hilaire)Jussieu ex Martius.
Distribution.S Brazil, Paraguay,and Misiones (Argentina).Fig. 4A.

22. Esenbeckiafebrifuga(Saint-Hilaire)Jussieu ex Martius,Nov. Gen. sp. pl. 3:

82, t. 233. Jan-Mar? 1831;T. F. L. Nees, P1. med. Suppl. t. 94? 1833
(the plate copied from Martius);Spach, Hist. nat. veg. 2: 343. 1834;
Walpers, Repert. 1: 501. 1842; Martius, Syst. mat. med. bras. 39.
1843. Fig. 32.
Euodia ("Evodia") febrifuga Saint-Hilaire, [M6m. Mus. Hist. Nat. 10: 146, 155, 159. after 21
Apr, 1823, nom. nud.] Bull. Sci. Soc. Philom. Paris (1823): 129. Sep 1823; Mem. Mus. Hist.
Nat. 10: 363. after 6 Nov, 1823; de Candolle, Prodr. 1: 724. mid Jan 1824; Saint-Hilaire, Fl.
Bras. mer. 1: 79. 26 Mar 1824; PI. us. Bras. 1: t. 4: 1. cum icon. 31 Mar 1824; Hist. pl. rem.
Br6s. Parag. 1: 149. 4 Nov 1825; P1. us. Bras. 1. substitute: t. 4. cum icon. 13 Aug 1827;
Pohl, PI. Bras. 2: 43. 1830, pro syn.; Dietrich, Syn. pl. 1: 498. 1839; Tulasne, Ann. Sci. Nat.
Bot. s6r. 3. 7: 283. 1847.
Esenbeckia febrifuga (Saint-Hilaire) Jussieu, Mem. Mus. Hist. Nat. 12: 486. 1825 = page 103 of
reprint of Dec 1825, comb. invalid. ex Art. 33 ICBN; Engler in Martius. Fl. bras. 12(2): 141.
1874; Chodat and Hassler, Bull. Herb. Boissier. s6r. 2. 4: 1285. 1904; Cowan and Smith in
Reitz, Fl. Ilustr. Cat. I (RUTA) 43, t. 11, fig. a-d. 1973.
100 Flora Neotropica

FIG. 32. Esenbeckia febrifuga. A, habit (Herb. Warming 246311, C). B, indument of lower side
of leaf (Assis 178, MO). C, flower (Herb. Warming 246311, C). D, fruit (Glaziou 18973, C). E, axial
part of endocarpwith two obturators(Curran18, NY). F, seed (Glaziou 18973, C).
Esenbeckia 101

Esenbeckiafebrifuga(Saint-Hilaire)Martiusin Spix and Martius,Reise 2: 789. between 1828and

1831, comb. invalid. ex Art. 33 ICBN; Saint-Hilaireand Tulasne, Ann. Sci. Nat. Bot. ser.
2. 17: 139. 1842.
Esenbeckiafebrifuga (Saint-Hilaire)G. Don, Gen. hist. 1: 795. early Aug 1831.
Colythrumfebrifugum (Saint-Hilaire)Schott, Rutac. 14. 1834.
Metrodoreagracilis Schumann,Bot. Jahrb. Syst. 30. Beibl. 67: 30. 1901, syn. nov.; Glaziou,
Bull. Soc. Bot. France 52. Mem. 3: 85. 1905;Engler in Englerand Prantl,Nat. Pflanzenf.
(ed. 2) 19a: 282-283. 1931. TYPE. Glaziou 20246, Brazil. Minas Gerais: Between Piedade
and Santa Luzia, 21 Nov 1893,fl (holotype, B-Herb. Urban, destroyed, photo 12488made
by F, F, NY; isotypes, BR, C, G, K, LE, LY, P-Herb. Glaziou 2x).
Esenbeckiafebrifuga (Saint-Hilaire)Jussieu var. fruticosa Hassler, Repert. Spec. Nov. Regni
Veg. 10: 345. 1912, syn. nov. TYPE. Hassler 8936a, Paraguay.Caaguazu:Nr. Caaguazu,
Mar 1905, fl & fr (lectotype, G-Herb. Chodat & Hassler, photo 26406 made by F, F, MO,
NY; isolectotypes, BM, K, P).
Shrub or tree, 0.15-11 m tall with trunkto 10 cm in diam.; branchlets2-5 mm
in diam., often reddish-brownwhen young, grayish-brownwhen older, minutely
pubescent with spreading hairs 0.1-0.2 mm long, becoming glabrous. Leaves
(sub)opposite, 3-foliolate, with stalked leaflets; petiole semiterete, strongly can-
aliculate, winged or not, 2-6 cm long, the wings 0.2-0.4 mmbroadand terminating
in a small ear 0.2 mm long, minutely puberulouswith hairs 0.2 mm long, a tri-
gonous tubercle 0.2-1 mm long usually present; petiolules to 20 mm long, not
sharply separatedfrom base of leaflets; leaflet blades elliptic or slightly obovate,
rarely ovate, (4.5-)5.5-16 x (2.0-)2.5-5.5 cm, lateral leaflets somewhat smaller
thanterminalone and unequalat base, long-attenuate,acuminateat apex, the very
tip obtuse or occasionally emarginate,rarelysubcaudate,the marginrevolute, the
blade chartaceousor rarely (sub)coriaceous,dull, rarely sublustrous,much paler
beneath, nearlyglabrousabove but the midveinsparselyminutelypuberulouswith
hairs 0.05-0.3 mm long, sericeous-pubescentbelow near the proximalpart of the
midveinwith mostly straighthairsto 0.8 mmlong, occasionally subglabrousbelow
with the midvein minutely pubescent with appressed hairs 0.05-0.3 mm long,
venation camptodromous,in subcoriaceous leaflets prominentbeneath, midvein
impressed, rarelyplane. Inflorescencesterminal,erect, (occasionally widely) pa-
niculate, nearly always longer than the leaves, (5-)13-25 x (4-)8-25 cm, usually
loosely but many-flowered,the side-branchletsof first order opposite and sub-
tended by a bract, or the lower ones by a 3-foliolate, rarely simple leaf, the side-
branchletsof higher order alternatedistally, pilose with spreadinghairs 0.05-0.2
mm, distally more dense, glabrescent;bracts except the lower ones persistent,
ovate, ca. 0.8-2 x 0.5-1 mm, minutelypuberulouswith hairs 0.05-0.1 mm long;
pedicels to 2 mm long; bractlets subopposite. Flowers usually 4.5-5.5 mm in
diam., protandrous;calyx lobes quincuncial,very broadly to depressedly ovate
with rounded tip, 0.7-1.5 x 1-2 mm, coriaceous with transparentmargin, gla-
brous above, glabrousor minutelypubescent below with hairs 0.05-0.1 mm long,
venation parallellodromous,false midrib present; petals persistent, imbricate,
spreading,subovate, elliptic or oblong, 2-2.5 x 1.2-1.6 mm, roundedat tip, semi-
transparent-papery,whitish when fresh, brown when dried with 0.1-0.3 mm
broad, the ochreous marginsbecoming creamish-whitewhen boiled, glabrousor
minutely puberulousbelow with appressed hairs 0.1 mm, venation camptodro-
mous to somewhat suprabasallyactinodromous, occasionally some showing a
false midrib;filaments persistent, adnate at base to the disc, subulate, slightly
convex abaxially, convex adaxially, 1.4-1.5 mm long and 0.3-0.4 mm thick, gla-
brous; anthers dorsifixed 1/3 to 2/5 from base, heart-shaped,0.6-0.7 x 0.5 mm,
including a tip 0.05 mm; disc cup-shaped, with 5-10 slight incurvations and a
thickened upper margin, 0.5 mm high (as high as the ovary with its protuber-
ances), 0.2-0.3 mm thick at the margin, 1.4-1.6 mm in diam., glabrous;carpels
adnate to the disc and connate in their lower half, 0.5 mm high, provided with
102 Flora Neotropica

numerous finger-like,glandularprotuberancesto 0.2-0.4 mm long and glabrous

or rarely beset with some hairs 0.1 mm long; style inserted near base of the
carpels, 0.8-0.9 mm long and 0.2-0.3 mm thick at maturity,free for 0.6-0.7 mm,
before the sheddingof the pollen 0.4-0.6 mm long and free for 0.3 mm, glabrous;
stigma capitate, 0.1-0.2(-0.5) x 0.3 mm. Fruits ca. 4-10 per infructescence,
subglobose, 10-15 x 10-14 mm (-20 mm when dehisced), glabrous;loculi round-
ed on the back, muricate with numerous + hooked prickles ca. 2(-4) mm long
which are sharp with a broad base or occasionally 2 with a common base, and
beset with glands, dehiscent septicidally to ca. 2 mm above the base and loculi-
cidally to 1/5 or 1/2 from the base; seeds (1-)2 per loculus, superposed,(narrowly)
ovoid, 6.5-7.5 x 2.7-2.8 x 2.8-2.9 mm, when 2 truncate at base or ventrally
near the base, flattenedon the back, beaked at apex, testa darkbrown, colliculate
with interspaces 0.05 mm in diam.; chalazal area irregularin shape, 1-1.2 mm in
diam., often much swollen, darkbrown;hilum0.5 mm broad;granularendosperm
0.1-0.2 mm thick; embryo 1, the cotyledons provided with ears 0.3 mm long,
finely punctate, radicle long-projecting,thick-conical,0.8-1.1 x 0.7-0.8 mm, the
plumule 0.1 mm long.
Type. Saint-Hilaire717, Brazil. Minas Gerais: Nr. Belo Horizonte and Itabira,
Bois vierge dans la Capitale des Mines, Feb 1817, fl (lectotype, P-Herb. St.-Hil.
Fl. Bras. Mer.; isolectotype, P ("Forets du prov. de R.J., M. Geraes et Espirito
Santo"); fragment,P-type Herb. Baillon (s.n.).
Distribution. SE and S Brazil (Ceara and Mato Grosso (rare), Minas Gerais,
Sao Paulo, Rio de Janeiro, Paranai);Paraguay;Argentina(Misiones chiefly near
IguaguFalls). Commonin forests and capoeirao, sometimes on sandy soils. Once
reportedfrom the restinga near Rio de Janeiro;alt. to 1100 m. Flowering round
the year but mainly Oct-Feb. Fig. 26B.
Specimens examined. BRAZIL. Ceara: Allemdo 281 fl (R). Mato Grosso: Manso s.n. (184., Cuiabd)
fl (BR). Minas Gerais: Assis 178 fl (GH, MO, UC); Barreto 163 fr (F), 6103 fr (F, R), 8960 fl (F, R),
10334 fl (R), 10373 fl (R); Gardner 4439 fr (BM, K 2x); Glaziou 18973 fl & fr (C, K, LE, P 3x, R);
Regnell II 53 (14 Aug 1866) fr (S 2x); Riedel 324 fl (LE 2x), 1394 fl (LE), 1439 fl (LE 2x), 1851 fl
(LE); Sellow 1424 fl (Minas?) (K, LE, UC, ZT); Stephan s.n. (1843) fl (BR); Warming s.n. (Lagoa
Santa) fl (K, P 2x, S, W); Herb. Warming 2463/1 fl (C, P), /3 fr (C), /4 fl (C), 15 fr (C), /6 fl (F);
Widgren 1078 fl (BR); L. 0. Williams & Assis 6093 fr (GH). Sao Paulo: Bockermann 79 fl (SP);
Burchell 4935 fr (K), 5045 fr (GH, K, P); Edwall CGGSP 3409 fl (SP); Gehrt SP 19855 fl (SP), SP
37019 fl (SP); Heiner s.n. (Campinas) fl & fr (S); F. C. Hoehne & Gehrt 39536 fr (SP); W. Hoehne
SP 54145 fl (SP); Koscynski 295 fl (SP); M. Kuhlmann 782 fr (SP), 904 fl (SP), 4507 fl (SP); Langsdorff
s.n. (Dec 1825, Itti) fl (LE); Lofgren CGGSP 420 fl (C, SP), CGGSP 496 fl (SP), CGGSP 623 fr (C,
P, SP); Lund s.n. (Feb 1834, Herb. Warming 2462) fr (C); Martius 1821 fl (M), s.n. (Dec, "inter
Portum Felicam et Ypanema") fl/fr (M 2x); Mosen 1184 fr (S); Novaes SP 2032 fl (SP); Vecchi ESP
212 fl (R). Rio de Janeiro: Galvdo s.n. (18 Dec 1877, =Herb. Glaziou 11853) fl (C, K, LE, P); J. G.
Kuhlmann 211 fl (NY); Schott 69v fl (W); Schreiner R 71497 fl (R). Parana: Duarte & Pereira 1632
fl (NY); Dusen 22/33 fl (S), 11198 fl (MICH, S), 11259 st (S), 16818 fl & fr (GH, MO, S); Falcdo 93-
51 fl (NY); Hatschbach 7568 fl (L), 10390 (=Pereira 7774) fl (U), 11215 fl (L), 15740 (=Lindeman
& de Haas 4250) fl (NY, U, W, WIS), 19077 fl & fr (UC), 21113 fr (MO, UC), 26685 fl & fr (NY,
UC); Hatschbach & de Haas 16638 (=Lindeman & de Haas 5518) fr (U); Hatschbach et al. 13238
fl (U 3x); Lindeman & de Haas 916 st (U), 1480 st (U), 1569 fr (U), 1705 fl & fr (K, NY, U, WIS 2x).
4351 fl (K, NY, U, Z), 5377 fr (K, NY, U); Schwarz 7415 fr (NY), 7489 fl (BR, S), 7490 fr (LD).
State unknown: Muller s.n. (S. Brazil) fl (P); Riedel s.n. (C, E, G, GH, GOET, K, L, LE, M, NY.,
P 3x, S, W, ZT); Sellow 2177 fl (S), B 2177. c 2173 fl (ZT), 5310 fl (LE), s.n. st (UC).
PARAGUAY. Amambay: Hassler 11439 fl (A, BM, G 3x, K, MPU, NY, P 2x), 12076 fl (A, G 5x.
MPU). San Pedro: Krapovickas et al. 14261 fl (C, UC); Pedersen 9379 fl (C, K, L, NY, S, UC, ZT):,;
Woolston 256 fl (NY, S), 256-c fl (K). Caaguazi: Balansa 2516 fl (G, GOET, K), 2518 fl (G, GOET,
K, S); Hassler 8936 fl (BM, G, K, P). Alto Parana: Fiebrig 6331 fl (BM, G 4x, GH, K); Hassler 4905
fl (A, G 2x, K, NY, P, S, UC, W). Guaira: Jorgensen 3975 (5 Oct or Nov 1928) fl (DS), 3975 (20 Oct
or Nov 1928) fl (MO), 3975 (19 Oct 1929) fl (C), 3975 (Sep 1931) fl (S), 3975 (Oct 1931) fl (NY); Rojas
12002 fl (W). Department unknown: Hassler 1439 fl (G).
ARGENTINA. Misiones: Curran 18 fr (GH, NY); Devoto BAB 90810 fr (BAB 2x); Herb. LPS
Esenbeckia 103

22278 fr (LP); Lillo 10593 bud (F); Meyer 5350 fl & fr (A, BM, BR, F 2x, S, UC, WIS); Rodriguez
433 fr (F, UC, US), 436 fr (GH); Rothkugel & Devoto BAB 54371 fr (BAB); Sandeman 4729 fl (OXF,
K), 4780 fl & fr (OXF, K).
CULTIVATED. F. C. Hoehne SP 31389 fl & fr (F, M, NY 2x, P, SP); (Mennega) 66-1774 (from
seed Lindeman & de Haas 1705) fl (U).

Local names and uses. Brazil: Tres-folhas (-vermelha)and laranjeira-do-mato

in Ceara, Minas Gerais and Sao Paulo (Allemdo 281, Saint-Hilaire 717, F. C.
Hoehne & Gehrt SP 39536, F. C. Hoehne SP 31389); tres-folhas-do-mato and
mendanhain Minas Gerais (Stephan s.n. 1843)); grumarim(-de-campos)in Rio
de Janeiro (Galvdo s.n.); mamoninhain Sao Paulo and Rio de Janeiro (F. C.
Hoehne & Gehrt SP 39536, J. G. Kuhlmann 211, Vecchi ESP 212). Paraguay:
ivira-fieti-(m)iin San Pedro (Hassler 1439, Woolston256). Very hard, white wood
of first quality for tooth-picks (Duarte & Pereira 1632). The bark resembles An-
gostura bark, and was imported into Europe about 1813 as "Brazilian China
bark." It has a strong and bitter taste (see PHYTOCHEMISTRY).
This species has much in common with Esenbeckia densiflora, and E. pumila,
and resembles vegetatively E. hieronymi. The leaflets are nearly always ? seri-
ceous-pubescent beneath, but occasionally subglabrous.The sericeous hairs are
somewhat thicker than those of E. densiflora, forminga less lanate indumentof
straighterhairs.
There has been much confusion about who first validly made the combination
Esenbeckiafebrifuga. Martiusdid so, but apparentlyonly a few months before
G. Don.
The collection F. C. Hoehne SP 31389 has largerflowers, 6-6.9 mm in diam.,
with petals 3-3.5 x 1.7 mm and minutelypuberulous;calyx minutelypubescent,
filaments 1.6-1.8 mm long, fruits subglobose but slightly stellately-lobedand up
to 18 x 17 mm with fewer prickles. The leaves are subglabrous,and in the inflo-
rescence some small 1-foliolateleaves are present.
The collection J. G. Kuhlmann211 is somewhat more robust than other spec-
imens. The leaflets are up to 19.5 x 6.7 cm and lack sericeous hairs. The inflo-
rescence is not yet fully developed, but the flower organs are 0.1-0.3 mm longer,
the petals 2.7-3 x 1.5 mm. Similarto this collection is Dusen 11259 (st).
Metrodoreagracilis is not different and therefore I reduce it to synonymy of
E. febrifuga. Formerly, Cowan transferred M. gracilis to Esenbeckia without
having seen the type. His transferwas promptedby a collection which appears
to represent E. densiflora.

23. Esenbeckiadensiflora(Chodat& Hassler) Hassler, Repert. Spec. Nov. Regni

Veg. 10: 344. 1912;Engler in Engler and Prantl, Nat. Pflanzenf. (ed.
2) 19a: 282. 1931. Fig. 33.
Esenbeckiafebrifuga (Saint-Hilaire)Jussieu var. densifloraChodat& Hassler, Bull. Herb. Bois-
sier. ser. 2. 4: 1285. 1904.
Esenbeckiagracilis Cowan, Sellowia 12: 89, t. 3, fig. g-j. 1960,syn. nov.; Cowanand Smith in
Reitz, Fl. Illustr. Cat. 1 (RUTA)39, t. 8, fig. g-j. 1973,non Metrodoreagracilis Schumann.
TYPE. Reitz4355, Brazil. SantaCatarina:Ilha de S. C., Florian6polis,Naufragados,14Dec
1951,fl (lectotype, US-2059577;isolectotypes, G, NY, S, U, UC, all ex HBR).
Shrub or small tree 2-12 m tall with trunk 10-40 cm in diam.; branchlets 1-3
(-6) mm in diam., green, becoming reddish-brown,shining, appressedlypuber-
ulous with hairs 0.2 mm long, becoming glabrous. Leaves (sub)opposite, 3-fo-
liolate, with leaflets stalked, some in the inflorescence 1-foliolate or simple;
petiole semiterete, canaliculate,obsoletely winged, 1.5-5.5 cm long, in the inflo-
rescences to 3 mm long, minutely pubescent with hairs 0.1-0.2(-0.3) mm long,
104 Flora Neotropica

FIG. 33. Esenbeckia densiflora. A, habit (Pedersen 4210, C). B, indument of lower side of leaf
(Hassler 12358, NY). C, flower (Pedersen 4210, C). D, fruit (Balansa 2518b, LD).
Esenbeckia 105

provided with a trigonous, bluntish, hairy tubercle to 1 mm long; petiolules in-

distinctly separatedfrom base of the leaflets, canaliculate,6-18 mm long, that of
terminalleaflet longer than the rest; leaflet blades elliptic or slightly ovate, rarely
slightly obovate, 6-12.5 x 1.8-4 cm, very long-attenuateat base, the lateralones
unequal at base and shorter than the terminal one, acuminate or subcaudateat
apex with the very tip obtuse, acutish, or emarginate, the margin thick and
(sub)revolutetowards the base, those in the inflorescencesup to 2.5 cm long, the
blades chartaceous,dull green above, dull and palerbeneath, (sub)glabrousabove
except for some hairs 0.1-0.3 mm near base, (sub)glabrousbeneath but hairy
proximallyalong the midvein or sometimes up to the margin with hairs 0.6-0.9
mm long and sericeous or lanate, spreading,white, and thin, in terminalleaflets
often on either side of the midvein, in lateral leaflets on the admedialside, ve-
nation camptodromousto slightly brochidodromous,the midvein somewhat im-
pressed above. Inflorescences terminal, erect, often longer than the leaves, 12-
17 x 10-17 cm in all, many- and ratherdensely flowered, proximallydispersedly
pilose, distally densely pilose to subtomentellous with hairs 0.1-0.2 mm, com-
posed of 1 terminaland several axillary, paniculately-branchedparts 7-11 x 3-
6 cm, the secondary branchlets(sub)opposite, the branchlets of the third order,
at least the upper ones, alternate;lower bracts representedby 3- or 1-foliolateor
simple leaves; upper ones persistent, triangular-ovate,concave, to 2.5 x 1 mm,
with few appressedhairs 0.1 mm long above, subglabrousor minutelypubescent
below with hairs 0.1 mm; pedicels 1-2 mm long; bractlets opposite or suboppos-
ite. Flowers 6-8 mm in diam., protandrous,fragrant;calyx lobes quincuncial,
subcircular, 1-1.5 x 1-1.6 mm, rounded at tip, thickly coriaceous, minutelypu-
bescent on both sides but very dense at the upper base with hairs 0.05-0.1 mm
long above and 0.1 mm long below, venation parallellodromous,provided with
a glandularfalse midrib;petals persistent, cochlear or quincuncial, spreading,
elliptic, 2.9-3.2 x 1.8-2 mm, obtuse or rounded, papery, ? transparent,yellow-
ish-white when fresh, yellowish when boiled, glabrousabove, minutelypubescent
below with hairs 0.1 mm long, venation camptodromoustending to become su-
prabasally actinodromous, the median nerve thick and strongly branched;fila-
ments persistent, narrowly triangularin outline, abaxially flattened, adaxially
convex, 1.5-1.6 mm long and 0.5 mm thick, glabrous;anthers dorsifixed1/3 from
the base, heart-shapedto subcircular,0.8-0.9 x 0.7-0.8 mm with a mucro 0.05
mm; disc cup-shaped, 10-plicate,0.5-0.8 mm high (as high as the ovary with its
protuberances),0.2 mm thick and 1.6-1.8 mm in diam., coriaceous, glabrous;
carpels adnate to the disc, free distally, 0.5-0.6 mm high, sericeous-pilose at tip
with hairs 0.3 mm long, densely beset with finger-likeglandularprotuberancesof
? equal height (0.3-0.5 mm); style inserted midway on the carpels, ca. 0.5 mm
long but becoming 0.8-0.9 mm after the sheddingof the pollen, and 0.2-0.3 mm
thick; stigma capitate, slightly irregularly5-lobed, 0.2 x 0.4 mm. Fruits 3-4 per
infructescence, subglobose with trigonous apophyses 4-5 mm long, 10-15 x 13-
21 mm when dehisced, with glands, glabrous, muricate(-echinate when young)
with often slightly recurved prickles initially up to 4 mm but finally shrinking
down to 2 mm long, some of them confluentat base, dehiscent septicidallyup to
the base and loculicidallyfromthe tip up to V/9from apex; seed (not seen) probably
6-10 mm long.
Type. Hassler 1695, Paraguay.Nr. Sapucai, Dec, fl & fr (lectotype, G-Chodat
& Hassler; isolectotypes, BM, G 2x, K, NY 2x, P, UC ex G).
Distribution. Brazil (Santa Catarina, Ilha de Santa Catarina)and Paraguay.
Secondaryforests, along marginsand rivers; alt. 20 m (1 record only). Flowering
(Sep-)Oct-Jan. Fig. 26B.
106 Flora Neotropica

Specimensexamined. BRAZIL. Santa Catarina:Klein6926 fl (US), 6927 fl (US); J. G. Kuhlmann

RB 73689 fl (NY).
PARAGUAY. Amambay: Rojas (in Hassler) 10659 fl (K). La Cordillera: Fiebrig 518a fl (A, E, F,
G 3x, K, M, PR);Hassler 1579fl (G), 2107 fl & fr (A, BM, G 5x, K, NY, P, UC, W), 2107a fr (G 6x,
photo, F, MO, NY), 3401 fl (A, BM, G 2x, K, NY, P, UC, W), 12358fl & fr (BM, C, E, F 2x, G 4x,
GH, L, MO 2x, NY 2x, S, UC, Z); Pedersen 4210 fl (C, K, L); Rojas 1579 fl (S). Central: Morin 716
fl (MICH); Pedersen 5137 fl (C); Ziircher 165(52.36) fl (Z). Guaira: Balansa 2518b fl & fr (BM, LD,
S); Jorgensen 3975 (Oct 1930)fl (F), 3975 (Oct 1931)fl (A), 4793 fl (F only). Departmentunknown:
Balansa 2518a fl (G 2x, GOET, K).

Local names. Paraguay:ivira-ovi-guagu(Morin 716); Ilha de Santa Catarina:

cutia; canela-de-veado (J. G. Kuhlmann RB 73689).
This species differsvegetativelyfrom Esenbeckiafebrifugain the leafletswhich
are densely sericeous or lanate along the midvein beneath. In E. febrifuga the
midveinis beset with muchfewer and straighterhairsor is occasionallyglabrous.
Esenbeckia gracilis was published by Cowan as a new combinationbased on
Metrodorea gracilis. It is indeed true, that M. gracilis does not belong to that
genus, as Cowan mentions. However, the specimens studiedby Cowan and cited
under E. gracilis are referable to E. densifora. Metrodorea gracilis is synony-
mous with E. febrifuga.

24. EsenbeckiahieronymiEnglerin Engler and Prantl, Nat. Pflanzenf.3(4): 159.

Mar 1896; Bot. Jahrb. Syst. 21. Beibl. 54: 27. 12 May 1896;Cowan,
Sellowia 12: 89, t. 3, fig. k-n. 1960;Cowan and Smith in Reitz, Fl.
Ilustr. Cat. I (RUTA) 41, t. 8, fig. k-m, t. 10. 1973. Fig. 34.
Treelet or shrub 2-5 m tall; branchlets 2-4 mm in diam., grayish-brown,the
younger ones chestnut, sparinglyminutelypubescent with appressedhairs to 0.2
mm long, becoming glabrous. Leaves (sub)opposite, 3-foliolate, petiolulate or
not; petiole terete, canaliculate, slightly ribbed below the leaflets, 1.4-4.5 cm
long, minutely pubescent with hairs to 0.1 mm long, provided with a short tri-
angularappendageprojectingbeyond the very base of the terminalleaflet; pet-
iolules terete, canaliculate, in terminalleaflets (0-)2-12 mm, in lateral ones (0-)
0.7-8 mm long; leaflet blades (narrowly)elliptic, the terminalone 5-12 x 2-4.3
cm, the lateral ones 4-8 x 1.5-3.1 cm, attenuate or obtuse at base, the lateral
leaflets slightly unequal at base with the exmedial side broadest, (sometimes
slightly) acuminate at apex with a blunt or acute tip up to 1 cm long, margin
revolute, the blade chartaceous, dull green above, paler beneath, subglabrous'
venation camptodromousto slightlybrochidodromous,midveinimpressedabove.
Inflorescences terminal, erect, paniculate, usually longer than the leaves, to
11 x 9 cm, many- but ratherloosely flowered, minutelypubescent with hairs to
0.1 mm long, side-branchlets(sub)opposite, slender;bracts (very broadly)ovate,
+ conduplicate, 1-2 x 0.7-1.8 mm, glabrous or with some short, silky hairs at
base above, minutely pubescent below, with 3 parallelnerves branchingfrom a
common point just below the base, the median nerve distinctly thicker and sur-
roundednearthe tip above with glandulartissue; pedicels ca. 1 mm long; bractlets
2, (sub)opposite, sometimes subtending secondary pedicels. Flowers 2.5-5 mm
in diam., protandrous;calyx lobes quincuncial,very broadlyovate to very broad-
ly triangular,0.7-1.1 x 1-1.5 mm, obtuse, coriaceous, with creamymarginswhen
dried, with short silky hairs 0.05 mm long on both sides, parallel-nervedwith a
false midrib above; petals persistent, valvate to cochlear, erect, ovate, 1.8.-
2.5 x 1-1.4 mm, obtuse, soft-coriaceous with papery margins, white or creamy
when fresh, brown with creamy margins when dried, glabrous above, minutely
puberulousbelow with hyaline hairs 0.05-0.1 mm long, parallel-nerved,the me-
dian nerve very thick (often +1/3 of the total breadth)and somewhat branching
Esenbeckia 107

FIG. 34. Esenbeckiahieronymi. A, habit (Reitz & Klein 826, U). B, flower (Reitz & Klein 826,
U). C, fruit (Reitz & Klein 826, U). D, embryo (Reitz & Klein 826, US).

towards the tip; filamentspersistent, partiallyadnateto the disc at base, subulate

with flattenedbase, 1.2-1.4 mm long, glabrous;anthersdorsifixedV3-/2 from the
base, heart-shaped,ca. 0.6-0.7 x 0.4-0.5 mm, includinga tip 0.05 mm; disc cup-
shaped, with 10 slight incurvations,0.8-0.9 mm high and 0.1 mm thick at tip, as
108 Flora Neotropica

high as the ovary with its protuberances, 1.5-1.7 mm in diam., thinly coriaceous
with thickened tip, glabrous; carpels adnate to the disc near base, free distally,
0.3-0.4 mm high, provided with glandularprotuberancesvarying to 0.3 mm long
and 0.1 mm in diam. and sparselypilose with hairs to 0.1 mm long; style inserted
/3 from base of the carpels, at last 0.7-0.9 mm long and 0.2 mm thick, glabrous;
stigma capitate, 5-lobed, 0.2-0.3 x 0.3-0.4 mm. Fruits globose, 10-14 mm in
diam., to 20 mm when dehisced, punctate with glands, the loculi trigonouswith
a dorsal apophysis 2/3 from base, muricatewith irregularlypyramidalblunt prick-
les to 2 mm long beset with some hairs, dehiscent septicidally from base to tip
but not up to the axis, the nerves of the exocarp visible externally on the sides;
seeds mostly 2 per loculus, obliquely tear-shaped, 4.4-5.4 x 3.1-4 x 3.4-3.6
mm, the upper seed with flattenedbase, beaked at apex, testa darkbrown, gran-
ulate, dull except for the granules, somewhat rugose; chalazal area roundishto
elliptic, thickened, 1.5 x I mm, dark brown; hilum impressed; granularendo-
sperm0.1-0.2 mm thick; embryo 1, cotyledons unequal;radicleprojectingbeyond
ears of cotyledons, recurved, and lying against them, 0.9-1 x 0.4-0.5 mm, plu-
mule with 2 very small leaf primordia.
Type. Ule 500, Brazil. Santa Catarina:Itajai, Jan 1886, fl (holotype, B, de-
stroyed, photo 12515made by F, F, MO, NY; isotype, HBG).
Distribution. Brazil, Mato Grosso (1 collection), Parana (1 collection), and
Santa Catarina.Capoeira,restinga, and also on river banks; alt. 5-300 m. Flow-
ering Jan-Apr, sometimes again in Jul; in Parana:Dec. Fig. 26A.
Specimensexamined.BRAZIL.MatoGrosso:AnonymusR 71068fl (R). Parana:Hatschbach28587
fl & fr (K, NY, S, UC). Santa Catarina:Bresolin 558 fl (US); Klein 1212 fl (NY, UC, US); Klein &
Bresolin 9373 fl (US); Reitz & Klein 826 fl & fr (G, NY, S, U, UC, US); 4104 fl (BR, GH, K, L, NY
2x, U. UC, US, WIS), 8279 fl (BR, F, G, K, L, M, NY, S, UC, US, Z).
Vegetatively this species can be confused with Esenbeckiafebrifuga and E.
densiflora (see under E. febrifuga).
III. Esenbeckiasubgen. LateriflorensKaastra, Acta Bot. Neerl. 26: 479. 1977.
Leaves alternate. Inflorescences lateral, reduced in size. Petals valvate, uni-
costate; anthers dorsifixed at middle. Seed 1 per loculus; chalazal area black-
brown; granularendospermremains absent.
Type species. Esenbeckia almawillia Kaastra.
Distribution.Remote localities in S. America. Fig. 4A.

25. EsenbeckiaalmawilliaKaastra, Acta Bot. Neerl. 26: 481, t. 6. 1977.

Fig. 36,2.
Shrub 1-10 m tall with tawny-ferrugineousindument;branchlets 2-4 mm in
diam., grayish-brownwhen dried, minutely pubescent with hairs 0.1-0.5 mm,
becoming glabrous and losing the longer hairs in age. Leaves alternate, subuni-
foliolate with sessile leaflet; petiole flattened adaxially, slightly canaliculate or
not, 4-10 mm long and 0.6-1 mm thick, minutely pubescent with hairs 0.1-0.2
mm long; leaflet blade (narrowly)elliptic or (narrowly)subobovate, 3-12 x 1.5--
4.5 cm, at base narrowly cuneate, acute, or obtuse, and usually equal, at apex
obtuse to acuminateor caudate-cuspidate,the acu.menemarginate,2-20 mm long
and laterally curved or not, the margin revolute at least towards the base, the
blade (sub)coriaceous,grayish-greenon both sides, + shining above, glands ex-
ternally not visible, subglabrousbut base, midvein, and marginpubescent with
hairs0.1-0.2 mm above and 0.1-0.3(-0.5) mm beneath,the hairsappressedbelow,
Esenbeckia 109

venation brochidodromous,midvein impressed above and longitudinallyribbed

or not, prominentbelow, lateralveins prominulousor plane. Inflorescencessev-
eral, lateral, each of them subtended by a bract or sometimes by a foliage leaf,
usually alternatingwith foliage leaves, consisting of a densely contractedpanicle
?0.5 x 0.5 cm of 3-8 subsessile flowers, minutely pubescent with hairs to 0.1
mm long; primarybracts caducous, narrowly triangularor subulate, 3-15 mm
long and ca. 0.4-1 mm broad, sometimes with a vestigial blade ca. 2.2 x 1.2 mm;
peduncle up to 2.5 mm long, in fruit 4-13 mm long and then occasionally once-
branchedbut all secondary branchletsusually deciduous; pedicels in fruit up to
1.5 mm long; bracts of higher order and bractlets caducous, narrowlytriangular
or provided with a reduced blade, up to 5 mm long. Flowers 5-6 mm in diam.;
calyx ca. 1.2-1.7 mm high, lobes separate, (broadly)triangular,ca. 0.7-1 x 0.5-
1 mm, acutish, coriaceous, glabrous above, strigillose below with hairs 0.2-0.3
mm long; petals persistent, valvate, ovate or elliptic, 2-2.5 x 1-1.2 mm, acute
at apex, coriaceous, yellowish, the internal glands not externally observable,
glabrous or papillose above, strigillose below with hairs 0.2-0.3 mm long, uni-
costate; filaments persistent, subulate, 1.7-1.9 mm long and 0.2-0.4 mm thick,
glabrous or papillose; anthers heart-shaped, 0.8-0.9 x 0.6-0.8 mm including a
mucro 0.2 mm long, pale yellow, brown-papillose;disc basally adnate to the
ovary, cup-shaped, 5-lobed, ca. 0.6-0.9 mm high and 1.5 mm in diam., higher
and darkerin color than the ovary, each lobe with a short tubercle in the middle,
glabrous; carpels adnate to the disc, connate, depressed, ca. 0.2-0.3 mm long
(excluding the apophyses), densely strigillose with hairs 0.2-0.3 mm long, pro-
vided with 5 free apophyses 0.2-0.3 mm long, hairylike the ovary and with many
minute glands; style long-persistent,inserted on the tip of the carpels among the
apophyses, subquinquangular,1 mm long and 0.2 mm thick, half hidden among
the apophyses, glabrous;stigmacapitate, 0.2-0.3 x 0.3 mm. Fruits, 1 per infruc-
tescence, depressed, stellately-lobed, 1-1.1 x 1.5-2 cm when dehisced; each loc-
ulus provided with a dorso-apical blunt apophysis 2-4 mm long and minutely
pubescent or strigillose with hairs 0.1-0.2 mm, dehiscent septicidally from 1-2
mm above base up to the tip; exocarp slightly wrinkledand minutelytuberculate,
prominentlynerved on both sides, the nerves runningparallel from the ventral
axis towardsthe tip of the apophyses; seed 1 per loculus, ovoid, ca. 7 x 3.2 x 3.2
mm, ratherstraighton the back, at apex roundedand shortly mucronateor not,
testa medium to dark brown, shining, finely and flatly reticulate-colliculatewith
interspaces0.05 mm in diam.; chalazalarea distinct, darkbrown-black;hilumca.
I mm broad;embryo 1 per seed, ovoid, cotyledons unequalwith ears ca. 0.4 mm,
punctate, radicle 0.6 mm long projecting0.2 mm beyond the ears, plumule with
2 leaf initials.
Type. Ule 9493, Bolivia. Pando: Rio Acre, nr. Cobija, Dec 1911,fl & fr (ho-
lotype, K).
Distribution.NE and W Brazil (Pernambucoand Acre) and E Bolivia (Pando
and Santa Cruz in the Chiquitos area). Forests; collected in flower Oct-Dec
(Acre, Pando), and Jun (Pernambuco).Fig. 35A.
Specimens examined. BRAZIL. Pernambuco: Serra do Araripe, Araripina, Andrade Lima & Md-
galhaes 52-1070 fl (R). Acre: Alto Xapuri, Seringal Guanabara, Ule 9494 fl (G).
BOLIVIA. Santa Cruz: Chiquitos, nr. San Rafael, Weddell 3448 fr (P 3x); nr Santa Ana (?),
d'Orbigny 764 fr (P).
Uses. The ChiquitoIndiansused and collected these plants for Paraguayyerba
(mate) (Weddell's catalogue, no. 3448).
The collection from Pernambuco has coriaceous, obtuse to subacuminate
110 Flora Neotropica

" '-
v E.amow'il,o
Ia/ '-7
;.1 ]
E.cowan1 |

* '
Mmaraccasanci' , (" ; '
M 7ida
fv- - /
v'~M'IG
. i....r .
3M. ...o
wosspceofEeb
stipularibtno ka(A,fMe a Runa (C).

'
, ' " .
R.mech'inatco l I--J]
Esenbeckia 111

FIG 36 Esenbeckia almawillia. A, habit (Ule 9494, G). B, flower with one petal removed (Ule
9494, G). C, fruit (d'Orbigny 764, P). D, seed (d'Orbigny 764, P).
112 Flora Neotropica

leaves. Although these leaves are rather different from those of the Ule speci-
mens, which are caudate-cuspidateand subcoriaceous, the specimens are iden-
tical in other characters. The leaves of the specimens from Santa Cruz (Bolivia)
are intermediatebetween those from plants from the two other areas.
One of the fruits of Ule 9493 is transformedinto galls that are echinate with
bristles 5-7 mm long.
This species was dedicated to Dr. A. M. W. Mennega who was so kind as to
help me solve some anatomicalproblems about this species, in particularwhen
I had only a single specimen at my disposal.

26. Esenbeckiacowanii Kaastra, Acta Bot. Neerl. 26: 481, t. 7. 1977. Fig. 37.
Tree 5-6 m tall with brown-hyalinepubescence of appressed hairs to 0.1 mm
long; branchlets 3-4 mm in diam., grayish-brown,becoming glabrous. Leaves
alternate, 1-foliolate, the leaflet sessile; petiole semiterete, winged, strongly
ribbed, 7-10 mm long, the wings ca. 0.2 mm broad, ending in an ear 2-4 x 0.9-
2.5 mm with rounded lateral-apicaltip; leaflet blade (narrowly) elliptic, 6.5-
18 x 2.5-7 cm, obtuse at base, the margin slightly revolute towards the base,
acuminateand often somewhat recurved at apex with an emarginatetip 5-7 mm
long, the blade chartaceousto subcoriaceous, shiningabove, dull beneath, glands
inconspicuous,glabrousexcept for midveinand base, venationbrochidodromous,
midvein plane and ribbed above. Infructescences several, lateral, axillary, erect
or spreading,paniculate, shorterthan the leaves, 4-5 x 2-4 cm, pubescent with
hairs to 0.2 mm; with many young fruits but only 1 in each infructescenceusually
maturing,the side branchlets alternate;bracts and bractlets caducous. Flowers
(studiedon young fruits of which 1 or 2 loculi are transformedinto galls) probably
ca. 4 mm in diam.; calyx lobes depressedly triangular,0.6-0.9 x 1-1.3 mm, ob-
tuse, thickly coriaceous, glabrousabove, appressed-pubescentbelow; petals per-
sistent, broadly ovate, ca. 1.4 x 1.1 mm, acute and slightly thickened at apex,
coriaceous, pubescent as the calyx lobes; filaments persistent, glabrous; disc
probably glabrous; carpels provided with an appressed-pubescentapophysis.
Fruits 1-2 per vegetative branch, broadest in the upper half, 1.8-2.3 x ca. 2.5
cm when dehisced, slightly yellowish-hoary with appressed hairs 0.1 mm long;
loculi provided /3 to /? from tip with a blunt apophysis 1-2 mm long, obsoletely
wrinkled,septicidallydehiscent from base to tip, often becoming completelyfree
when old and finally falling off, the exocarp internally smooth, its nerves only
slightly visible on the sides externally; seeds (only 1, empty, seed observed in
Cowan 38833 (F)), probably 1 per loculus, very obliquely tear-shaped ca.
7 x 4 x 4 mm, with apex 0.2 mm long; testa brown, shining, finely irregularly
colliculate with interspaces 0.05 mm in diam., chalazal area irregularlyshaped,
ca. 3 x 2 mm, black, shining;hilum 0.6 mm broad; embryo unknown.
Type. Cowan 38833, French Guiana. Guyane: Montagnede Kaw, alt. 250-270
m, 14 Dec 1954fr (holotype, US-2168712;isotypes, F, NY).
Distribution. Known only from the type locality. Occasional in low forest on
bauxite, and along seasonal pond; alt. 220-270 m. Collected in fruit in Dec. Fig.
35A.
Specimen examined. FRENCH GUIANA. Guyane: Montagne de Kaw: Cowan 38757 fr (K,
NY, P).

Unfortunatelythe flowers of Esenbeckia cowanii are poorly known, but the

position of the species is most likely to be in subgen. Lateriflorens.As to the fruit
E. cowanii is close to E. leiocarpa, but the fruit is less grayish and more yellow-
Esenbeckia 113

..
...
FIG.- 37. Esen b.. ..Cn .'. . '.....: . : B seed.
habit.
* F:IG : . .. , . ,. 37. . . ' .. .. . '
Esenbecki
.i :n ,38833t!' ,. , . ., .
 114 Flora Neotropica

ish-hairy. In vegetative characters however, there are distinct differences, e.g.

the 1-foliolateleaves with winged petioles and obtuse leaflet-bases.
Many young fruits of both known collections are transformedinto galls, echi-
noid in appearance,with numerousthin pricklyprotuberancesca. 5 mm long and
pilose at tip with hairs 0.05 mm long.
This species was dedicated to Dr. Cowan, the collector of the specimens. He
made his first botanicalinvestigationsin the Rutaceae, on which he has published
several articles.

InsufficientlyKnown
Esenbeckia pilocarpoides Kunth var. guianensis Engler in Martius, Fl. bras.
12(2): 144. 1874. TYPE. Schomburgk 569, Guyana (n.v.; disap-
peared?).
According to the description this variety should differ from Esenbeckiapilo-
carpoides in the presence of a dense hair-cover on the lower side of the leaves,
bracts, bractlets, and calyx lobes. All material of E. pilocarpoides studied in-
cluding the collections made in Guyana has glabrousleaves and calyx lobes.

Excluded from Esenbeckia

Esenbeckia altissima Blume, Bijdr. 119. 1825. =Neesia altissima (Blume) Blume,
Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 17: 83, t.
6. 1835.
Esenbeckiaatata Pittier, Arb. arbust. nuev. Venez. decada 1: 7. 1921;Bol. Com.
Industr. (Caracas) 13: 421. 1921 (n.v.); Trab. Mus. Corn. Venez. 7:
339. 1930;Englerin Englerand Prantl,Nat. Pflanzenf.(ed. 2) 19a:282.
1931;Record and Hess, Trop. Woods 64: 12. 1940. =Helietta plaeana
Tulasne, Ann. Sci. Nat. Bot. ser. 3. 7: 281. 1847.
All the type collections of both species appear to be similar, and belong to
Helietta. In the sheet of Pittier 9207 at US (paratypeof E. atata) young fruits
were observed which already show the wings typical of Helietta. The type of E.
atata has pentamerousflowers, the type of H. plaeana tetramerousones, while
the paratypes of E. atata show both of them. Because H. plaeana has priority,
E. atata is reduced to synonymy with it. Types studied:H. plaeana, A. Plee s.n.
(anno 1826) fl (holotype, P, photo 35726 made by F, n.v.; isotypes, P 3x, P-type
Herb. Baillon);E. atata, Jahn 492 fl (type, US-1198273);Pittier 7681 bud (para-
types, GH, US), 9207 fl (paratypes,G, GH, US-1122035).
EsenbeckiaberlandieriBaillon, Adansonia10: 151. 1871,nom. invalid.;non Hem-
sley, Biol. centr.-amer., Bot. 1: 170. 1879. =Helietta parvifolia Asa
Gray ex Bentham, Hooker's Icon. P1. ser. 3. 4: 66, t. 1385. 1882(see
this study, page 38).
Esenbeckia coriacea A. C. Smith in Gleason and Smith, Bull. Torrey Bot. Club
60: 358. 1933. =Metrodorea flavida Krause, Notizbl. K6nigl. Bot.
Gart. Berlin 6: 146. 1914, according to Macbride, Field Mus. Nat.
Hist., Bot. Ser. 13: 671. 1949.
Esenbeckia cuspidata Englerin Englerand Prantl,Nat. Pflanzenf.3(4): 159. Mar
1896,nom. nud. = next following.
Esenbeckia? cuspidata Engler, Bot. Jahrb. Syst. 21. Beibl. 54: 28. 12 May 1896.
Synonyms: Helietta longifoliata Britton in Morong and Britton, Ann.
Esenbeckia 115

New York Acad. Sci. 7: 69. 1892;Cowan and Smithin Reitz, Fl. Ilustr.
Cat. I (RUTA) 57. 1973(nom. illegit. ex Art. 63, ICBN), accordingto
R. E. Fries, Bull. Herb. Boissier. ser. 2. 7: 1002. 1907.Helietta cus-
pidata (Engler)Chodat& Hassler, Bull. Herb. Boissier. ser. 2. 4: 1285.
1904(nom. illegit. ex Art. 67, ICBN). =Helietta apiculata Asa Gray
ex Bentham, Hooker's Icon. PI. ser. 3. 4: 67. 1882.
The type of H. apiculata (Balansa 2515, holotype, K; isotypes, G 2x, GOET,
K, P 2x) constitutes also one of the syntypes of H. longifoliata. Helietta apiculata
was publishedunder H. parvifolia (plate 1385in Hooker's Icon. PI.). Therefore
this name has mostly been overlooked. Benthamundoubtedlyintendedto publish
H. apiculata as a distinct species, as appearsfrom the text. He must have pub-
lished it under the allied species H. parvifolia only for lack of a suitableplate.
Esenbeckia glaziovii Engler in Engler and Prantl, Nat. Pflanzenf. 3(4): 159. Mar
1896;Engler, Bot. Jahrb.Syst. 21. Beibl. 54: 27. 12 May 1896;Engler
in Engler and Prantl, Nat. Pflanzenf.(ed. 2) 19a: 282. 1931. =Helietta
plaeana Tulasne, Ann. Sci. Nat. Bot. ser. 3. 7: 281. 1847.
The isotypes and the photo of the holotype of E. glaziovii agree well with the
type of H. plaeana, as does the description.The isotypes at K and P show young,
winged fruits. According to Glaziou (1905) and some of the labels, the type lo-
cality was in Rio de Janeiro.There are three more collections from Rio de Janeiro
which appearto belong to H. plaeana: Galvdo387 (P-Herb. Glaziou)and Galvdo
565 (R 71216, ex Herb. Saldanha5680), both from Sao Fidelis, and Glaziou 18177
(R 7912) from the type locality (Alto Macahe). Consideringthat the main distri-
bution of H. plaeana is in Venezuela and Colombia,the occurrenceof the species
in Rio de Janeirois strange. In my opinion the four collections of E. glaziovii are
of doubtful origin. Glaziou was concerned with them, and he was sometimes
guilty of piracy (cf. under Pilocarpus microphyllusand P. spicatus subsp. ara-
catensis in the present treatment).However that may be, E. glaziovii has to be
transferredto Helietta and into synonymy with H. plaeana. Type specimens
studied:H. plaeana (see E. atata above); E. glaziovii, Glaziou 18171fl (holotype,
B, destroyed, photo 12514made by F, F, MO, NY; isotypes, C-Herb. Warming
2x, K, photo NS 2866 made by NY, NY; LE, P, P-Herb. Glaziou 2x).
Esenbeckia glaziovii Engler ex Glaziou, Bull. Soc. Bot. France 52, Mem. 3: 85.
1905,nom. nud. Based on Glaziou 18171(P-Herb. Glaziou). =Helietta
plaeana Tulasne, Ann. Sci. Nat. Bot. ser. 3. 7: 281. 1847(see above).
Esenbeckia lucida Rusby, Bull. New York Bot. Gard. 8: 98. 1912. =Galipea
lucida (Rusby) Kaastra, Acta Bot. Neerl. 26: 487. 1977.
Esenbeckia mollis Miquel, Linnaea 22: 796. 1850?(" 1849"); Engler in Martius,
Fl. bras. 12(2): 142. 1874. =Helietta mollis (Miquel) Kaastra, Acta
Bot. Neerl. 26: 487. 1977.
Esenbeckiapittieri Krause, SmithsonianMisc. Collect. 61(16): 1. 1913;Englerin
Engler and Prantl, Nat. Pflanzenf. (ed. 2) 19a: 281. 1931. =Conomor-
pha verisim. verticillata Zahlbruckner,Ann. Naturh. Hofm. Wien 7:
3. 1892(Myrsinaceae).
The leaves have brown peltate hairs (superficiallyresemblingoil glands!). The
flowers are pentamerous,glabrous,but the sepals and petals are glandular-ciliate;
sepals subcontortedin aestivation, ca. 0.6 mm long; petals contorted to the right
in aestivation, connate at base, 1.5 mm long; fertile stamens epipetalous, adnate
to the petals, alternatingwith 5 short, blunt staminodia; anthers + sagittate,
116 Flora Neotropica

latero-introrse;disc absent; ovary 1, unilocular,with basal placenta and probably

3 ovules with the micropyledirected downwards. Type: Pittier 1014 fl (holotype,
US-531213).
Esenbeckia plicata Nees ex Bridel, Bryol. univ. 2: 754. 1827. =Garovaglia pli-
cata (Nees ex Bridel)van den Bosch & van der Sande Lacoste, Bryol.
javan. 2: 79. 1863, n.v.
Esenbeckia riedeliana Engler in Martius, Fl. bras. 12(2): 142. 1874. Synonyms:
Helietta multiflora Engler in Martius, Fl. bras. 12(2): 185. 1874, ac-
cordingto Englerin Englerand Prantl,Nat. Pflanzenf.3(4): 174. 1896.
BalfourodendroneburneumMello ex Oliver, Hooker's Icon. P1. 13(1):
t. 1203-1204. 1877 (nom. illegit. ex Art. 63), according to Engler in
Engler and Prantl, Nat. Pflanzenf. 3(4): 174. 1896. =Balfourodendron
riedelianum (Engler in Martius) Engler in Engler and Prantl, Nat.
Pflanzenf. 3(4): 174. 1896;Chodat and Hassler, Bull. Herb. Boissier.
ser. 2. 4: 1286. 1904;Cowan, Sellowia 12: 93. 1960;Cowan and Smith
in Reitz, Fl. Ilustr. Cat. l(RUTA) 61. 1973.

METRODOREA
2. MetrodoreaSaint-Hilaire, Fl. Bras. mer. 1: 81. 11 Jun 1825; Adr. Jussieu,
Mem. Mus. Hist. Nat. 12:487. after27 Jun, 1825;Meisner,P1.vasc. gen.
1: 63 and 2: 45. 1837;Endlicher, Gen. pl. 1: 1153. 1840;Lemaire, Fl.
Serres Jard. Eur. 4: t. 337. 1848; Engler in Martius, Fl. bras. 12(2):
148. 1874;Englerin Englerand Prantl,Nat. Pflanzenf.3(4): 160. 1897,
and (ed. 2) 19a: 282. 1931;Macbride,Field Mus. Nat. Hist., Bot. Ser.
13: 670. 1949;Albuquerque,Anais Acad. Brasil. Ci. 40: 511. 1968.
Esenbeckiasect. MetrodoreaHooker in Benthamand Hooker, Gen. pl. 1: 300. 1862.
Terminalbuds completely enveloped by the sheaths of the upperpairof leaves.
Leaves opposite, 1-3-foliolate; sheaths of the leaves largely grown out distally,
adnate to the branchletsat base only, the upper part cucullate, round-tipped,the
inner margin densely ciliate with multicellularhairs, later on reflexing and be-
coming glabrous, finally falling off with the leaves; petiole present or seemingly
absent, adnate to the dorsal side of the sheath, semiterete to subterete, ? can-
aliculate, pilose when young, becoming glabrousin age; base of leaflets unequal,
venation brochidodromous.Inflorescences erect, (mostly widely) paniculate,the
lower side-branchletsopposite, the upper ones alternate, the basal ones usually
arising from the axil of the uppermostpair of leaves; bracts with a dorsal pseu-
docosta, the lower bracts caducous and sometimes leaf-like; bractlets usually
subtendingsecondary pedicels or buds. Flowers pentamerous;calyx with a con-
nate part up to half the length of the lobes, the lobes subseparateor subquincun-
cial, depressedly ovate, with a dorsal pseudocosta which originatesat base of the
calyx; petals valvate, the inner margininduplicate,with mostly unguiculatebase
and cladodromousvenation; filamentsaccumbentbetween the lobes of the disc,
reflexing, subulate, flattenedat base, glabrous;anthersdorsifixedat middle, ver-
satile, mucronate at apex, purplish;disc adnate to the ovary in the lower half,
annular,glabrous;ovary pentalocular,carpels connate; ovules 2 per carpel;style
terete or obtuse-angled. Fruits usually pentalocular subglobose capsules, with
glands, glabrous,the exocarpinternallyprominentlynerved, the loculi completely
connate and provided or not with apophyses, septicidally dehiscent along the
dorsal commissures, and loculicidally from the base along the sutures over the
Metrodorea 117

top; seeds with coriaceous testa, the chalazal area usually externally visible;
embryo 1 (as far as known), punctate or not.
Type species. Metrodorea nigra Saint-Hilaire. The name was dedicated by
Saint-Hilaire to "Metrodorus Sabinus, the first man, according to Plinius, who
illustrated plant descriptions." Plinius however, mentioned several Metrodori but
not any Metrodorus Sabinus. Probably Saint-Hilaire had in view ? 135 of book
XXXV of the Naturalis Historia, where Plinius mentions Emilius Paulus, who
looked for an educator for his little son and who was advised to ask Metrodorus
"pictor idemque philosophus, in utraque scientia magnae auctoritatis" (Schou-
ten, Maartensdijk, in verbis).
Distribution. Mainly Brazil, only few collections from Surinam and Bolivia (see
Fig. 35B).

Key to the Species of Metrodorea

1. Petioles completely adnate to the sheaths. 5. M. stipularis.
1. Petioles partlyadnate to the sheaths, free distally.
2. Branchletswhen young densely villous; sheaths 2.5-4.5 mm long; leaflets to 5 cm long;
fruits without apophyses. 3. M. mollis.
2. Branchletswhen young glabrousor pilose; sheaths 4-10 mm long; leaflets (3-)5-25 cm
long; fruits with apophyses.
3. Petioles 1-2(-15) mm long; leaves coriaceous; disc distally lobed; ovary deeply im-
mersed into disc, developinglarge apical apophyses immediatelyafter flowering.
1. M. maracasana.
3. Petioles 4-60 mm long, rarely shorterbut then the leaves chartaceous;disc distally
partite;ovary projectingbeyond disc, apophyses developingat a later stage.
4. Petals claret or purplish;filaments0.8-1.5 mm long; disc rathersmooth, charged
with minutetuberclesto 0.1 mm;fruits with dorsalapophyses3-5 mm long; seeds
mucronate. 4. M. nigra.
4. Petals yellow-white;filaments2.4-3 mm long; disc densely chargedwith tubercles
0.2-0.8 mm; fruits with dorsalapophyses 5-10 mm long; seeds roundedat apex.
2. M. flavida.

1. Metrodorea maracasana Kaastra, Acta Bot. Neerl. 26: 484, t. 8. 1977.

Fig. 38A-C.
Tree 5-12 m tall, with trunk to 25 cm in diam.; branchlets 2-6 mm in diam.,
grayish-brown, somewhat shining, lenticels roundish or spindle-shaped ca. 0.3-
1 mm, glabrous. Leaves 1-foliolate with ? sessile leaflet, glabrous; sheath 4-7
mm long, ciliate with hairs ca. 0.5 mm long; petiole semiterete, widely shallowly
canaliculate, 1-15 mm long and ca. 1-2 mm thick; leaflet blade elliptic or ob-
ovate, 5-14 x 2.9-7 cm, shortly attenuate or narrowly subcuneate at base, obtuse
or emarginate at apex, margin revolute, coriaceous, grayish-green on both sides,
shining, venation prominulous below, midvein impressed. Inflorescences termi-
nal, widely paniculate, about as long as the leaves, 3-9 x 4-10 cm, with stocky
branchlets, densely flowered, pubescent with spreading hairs 0.1-0.2 mm long,
the lowermost pair of side-branchlets subtended by the upper pair of leaves; the
upper bracts ovate or triangular, to 4 x 2 mm, glabrous above, pubescent below
with spreading hairs 0.1 mm long; pedicels to 2 x 0.5 mm; bractlets 2, alternate
or subopposite. Flowers ca. 6-7 mm in diam., perfumed; calyx lobes
(sub)quincuncial, connate and overlapping at base, depressedly ovate, 0.7-
1.2 x 1-1.5 mm, slightly acuminate and obtuse at very tip, or obtuse, glabrous
above, minutely pubescent below with hairs 0.05-0.1 mm long, parallel-nerved,
the nerves branching towards the tip, the pseudocosta indistinct; petals subquin-
cuncial to subvalvate, adnate to the disc at base, reflexed, elliptic, 2.7-3 x 1.2-
118 Flora Neotropica

FIG. 38. A-C, Metrodorea maracasana. D-F, Metrodorea mollis. A, habit (Pereira 9656 & Pabst
8545, type, F). B, flower and C, l.s. of flower (Pereira 9656 & Pabst 8545, type, M). D, habit (Glaziou
14588, type, C). E, immature fruit (Glaziou 15887, C). F, fruit (Gifford & Fonseca G 323, E).
Metrodorea 119

1.4 mm, postflorallyup to 3.6 x 1.5 mm, unguiculatewith claw ?0.5 mm acutish
at tip, coriaceous, in age becoming thinly so or semitransparentwith membra-
naceous margin, yellowish white, minutely pubescent on both sides with hairs
0.1 mm long; filaments 1.3-1.7 mm long and 0.2-0.3 mm thick, with same color
as petals; anthersheart-shapedca. 0.5-0.6 x 0.5-0.6 mm, purplish;disc in upper
half 5-10-lobed, longer than ovary, 0.6-0.8 mm high, ca. 0.4 mm thick at tip, 1.2-
1.5 mm in diam., becomingdark-coloredin age, smooth except for a few tubercles
0.05 mm; carpels deeply immersed into disc, ca. 0.3-0.5 mm high, glabrous,
charged with sometimes ? confluent tubercles to 0.3 mm long, postflorallyde-
veloping apical apophyses ca. 1.5 mm high and 1 mm thick, chargedwith internal
glands along the margin;style ca. 0.8 mm long and 0.2-0.3 mm thick, projecting
ca. 0.3-0.4 mm beyond the tuberclesof the ovary, glabrous;stigmacapitate, 0.1-
0.2 x 0.3 mm, dark. Fruits (nearlymature)depressedlyglobose, stellately-lobed,
18-20 x 20-30 mm, loculi provided with a dorsal apophysis 3-7 mm half-way to
the tip, transverselywrinkledwith prominentparallelnerves on both sides of the
wall, muricate-tuberculate,the tubercles to 2 mm.
Type. Pereira 9656 & Pabst 8545, Brazil. Bahia:Between Itiruguand Maracas,
23 Jan 1965,fl (holotype M; isotypes, F-1629627,LP, all ex HB 35012).
Distribution. Brazil (Bahia, Chapada de Maracaisand environs). Rain and
coastal forest. Flowering Oct-Feb. Fig. 35B (only type mapped).
Specimens examined. BRAZIL. Bahia: Between Itiruguand Maracas,Jesus 400 fr (U); between
Jaguaquaraand Rio Bahia,Pinheiro1865fr (U); Gandu,estradaa Itubera,Santos 1158fl (U); Itacare,
estradado Aeroporto,Santos 2934 fl (U).
This species superficiallyresembles Metrodoreastipularis but the petioles are
partly free and the leaflets are uniformly 1-foliolate.
The lobed (not partite) disc and the deeply immersed ovary which develops
well-sized apophyses are distinct charactersof this species. There is some resem-
blance with M. nigra, but the inflorescenceis more stocky with thickerpeduncles,
the flowers are a different color, and the leaves are more rigid; M. nigra has
usually longer petioles than M. maracasana.
2. Metrodoreaflavida Krause, Notizbl. K6nigl. Bot. Gart. Berlin 6: 146. 1914;
Engler in Engler and Prantl, Nat. Pflanzenf. (ed. 2) 19a: 282. 1931;
Macbride, Field Mus. Nat. Hist., Bot. Ser. 13: 671. 1949. Fig. 39.
Esenbeckiacoriacea A. C. Smith in Gleason and Smith, Bull. Torrey Bot. Club 60: 358. 1933;
Macbride,Field Mus. Nat. Hist., Bot. Ser. 13: 671. 1949, pro syn. TYPE. Krukoff1667,
Brazil. Rond6nia("Mato Grosso") probably:MachadoRiver region (=Rio Jiparana),nr.
source of JatuaranaRiver, 23 Dec 1931, dec. fr (holotype, NY; isotypes, BM, G, K, P,
S, U).
Shrub or usually tree (1.5-)3-18(-27) m tall; trunk 10-20(-40) cm in diam.;
branchlets 3-7(-10) mm in diam., brown, reddish-brownwhen young, shining,
lenticels spindle-shapedca. 0.3-1 mm long, puberulouswith spreadinghairs ca.
0.05 mm, becoming glabrousin age. Leaves 1-3-foliolate with usually sessile or
shortly stalked leaflets; sheath 4-10 mm long, inner marginciliate with hairs to
0.4 mm long, abaxial side pubescent with hairs to 0.4 mm, becoming glabrousin
age; petiole subterete, (shallowly) canaliculate, not or obsoletely winged, 4-40
mm long and 1-2 mm thick, pubescent with spreadinghairs ca. 0.05 mm, becom-
ing glabrous in age; wings to 0.2 mm broad and usually not eared; petiolules 0-
2 mm long; leaflet blades elliptic or slightly obovate, rarely ovate, 7-25 x 2.5-
9.5 cm or sometimes smaller towards apex of the branchlets, (shortly) attenuate
or narrowlycuneate at base, at apex acuminateor occasionally slightlyacuminate
or obtuse, the very tip obtuse, the margin not revolute or only slightly so, the
120 Flora Neotropica

FIG. 39. Metrodorea flavida. A, habit (Ule 9491, type, U). B, flower (Capucho 475, F). C, fruit
(Capucho 475, F). D, seed (Capucho 475, F).
Metrodorea 121

blade (sub)coriaceous, shining, glabrous or occasionally pubescent at base like

the petioles, becoming glabrousin age, venation ? prominent,midvein plane or
prominulous,sometimes slightlyimpressedtowardsbase. Inflorescencesterminal
or axillarynear tips of branches, widely paniculate,shorteror usually longerthan
the leaves, (7-)10-25(-30) x (5-)8-25 cm, numerous-flowered,pubescent with
spreading hairs 0.05-0.1 mm, the lowermost side-branchlets usually subtend-
ed by the upper pair of leaves; bracts (depressedly) ovate or triangular,to
2(-5) x 1.2-1.5(-4.5) mm, rarelythe lowest bractsleaf-like and up to 12 x 1 mm,
minutely pubescent below with hairs 0.05-0.1 mm long, less dense above; ped-
icels 1.5-6 mm long and 0.2-0.5 mm thick; bractlets 2, (sub)opposite. Flowers
6.5-8.3 mm in diam., usually 7.5-8 mm, not fragrant(Capucho475); calyx lobes
subquincuncial or subcochlear to separate, depressedly ovate, (0.4-)0.7-
1 x (0.7-)1-1.4 mm, slightly acuminateor obtuse, glabrous above, minutelypu-
bescent below with spreadinghairs 0.05 mm long; petals deciduous, valvate, very
widely spreading,ovate, unguiculate,3.5-4 x 1.5-2.5 mm, the claw 0.2-0.5 mm,
acutish at tip, thickly coriaceous becoming thinly so, yellow-white when alive,
brownish-yellowwhen dried, densely pubescent above with spreadinghairs ca.
0.05 mm long, or papillose, pubescent or subglabrousbelow with spreadinghairs
0.05-0.1 mm long; filamentspersistent, 2.4-3 mm long and 0.3 mm thick, yellow-
ish tending to become rose, glabrous; anthers broadly ovate to nearly heart-
shaped, 0.5-0.6 x 0.4-0.6 mm, becoming purplish, occasionally papillose; disc
in the upper half 5-partite,not projectingbeyond the ovary, (0.7-)1 mm high in
all, (1.8-)2(-2.5) mm in diam., becoming dark, upper part densely chargedwith
thick tubercles ca. 0.1-0.4 mm long; carpels 0.5-0.6 mm on a raised receptacle,
densely charged with clavate or finger-like, thick protuberances 0.2-0.8 mm,
which are glabrousor pilose with spreadinghairs0.05-0.1 mm long; style inserted
half-way on the carpels, (1-)1.2-1.5 mm long and 0.3-0.5 mm thick, projecting
0.5-1 mm beyond protuberances of ovary, with some basal glands, glabrous;
stigma capitate, 5-lobed, 0.2-0.3 x 0.4-0.5 mm. Fruits ? depressedly globose,
stellately-lobed, (15-)20-28 x (15-)20-30 mm(-48 when dehisced), loculi near
middlebearinga dorsal, bluntapophysis 5-10 mm long, transverse-parallelnerves
hardly visible externally except near the commissures and sometimes at tip,
strongly muricatewith tubercles to 3-5 mm long, dehiscent septicidallyfrom ca.
3 mm above base up to some mm below tip and loculicidallyup to (half-way)the
apophysis; seeds usually 2 per loculus, superposed, ovoid, 5-7 x 3.5-4.5 x 4-
5.5 mm, flattened at or near base, rounded at apex, testa thinly coriaceous, dull
mediumbrown, irregularlypustulateand reticulate-colliculate;chalazalarea tear-
shaped, shining, dark brown, ca. 3.5 x 2.5 mm; hilum ca. 1 mm broad; embryo
punctate, radicle projectingamong the ears.
Type. Ule 9491, Brazil. Acre: Rio Acre, Monte M6, Dec 1911,fl (holotype, B,
destroyed, photo 12487made by F, F, NY; isotypes, G, K, L, U, UC).
Distribution. Surinam, NW Brazil, and Bolivia. High forest on terra firme;
woods on lateritic soils, and capoeira;alt. 35-1500 m. FloweringmainlyOct-Jan.
Fig. 35B.
Specimens examined. SURINAM. Nickerie: Oldenburger et al. ON 508 st (U), ON 534 fl & fr
(K, U).
BRAZIL. Amapa: Egler & Irwin 45962 fr (U). Para: Capucho 475 fl & fr (F); Ducke MG 15116 fl
(BM, G, US), MG 15872 fl (BM, G), RB 13621 fl (K, U), RB 17734 fr (U), RB 17735 fl/fr (U), RB
17736 fr (U), RB 20474 fl (K, S, U, US); Prance & Silva 58694 (=L. S. 64) fr (F, G 2x, K, NY, P,
RB, S, U, UC, US); M. Silva & Souza 2266 fr (U); N. T. Silva 181 fl (U). Maranhao: Prance & Silva
58975 fl (GH, K, NY, S, U, US); N. T. Silva 57778 fl (NY, US 2x). Acre: Forero et al. 6335 fr (U);
Krukoff 5517 fr (A, BM, F 2x, G, K 2x, LE, LP, M, MICH, MO, NY, PR, S, U 2x, UC, US); Ule
9492 fl (G, K, L, UC). Rond6nia: Prance et al. 8339 bud (U), 8344 fl (U); Rodrigues & Wilson 3426
122 Flora Neotropica

(US); Rusby 2617 fl (GH, K, LE, MICH, NY 2x), 2663 fl (LE, NY 2x, MICH). Mato Grosso:Berg
et al. (in Prance) 18595 fl (U), 19842 fr (U); F. C. Hoehne Comm. Rondon 994 fl (R); Prance et al.
18225bud & fr (P, US).
BOLIVIA. La Paz: Cardenas 992 fl & fr (F, NY 3x); R. S. Williams 482 fl (NY 3x).

The flowers differ from those of Metrodorea nigra in the yellow-white color,
longer filaments, style already projectingbefore the shedding of pollen, and in
the more stronglytuberculateand less deeply partitedisc. VegetativelyM.flavida
resembles M. nigra.
Although the leaves are usually 3-foliolate, occasionally specimens show
1-foliolateleaves only.

3. Metrodoreamollis Taubert, Bot. Jahrb. Syst. 15. Beibl. 34: 5. 1892;Engler in

Engler and Prantl, Nat. Pflanzenf. 3(4): 160. 1896. Fig. 38D-F.
Metrodoreamollis Taubertvar. glabrata Taubert,Bot. Jahrb.Syst. 15. Beibl. 34: 6. 1892, syn.
nov. TYPE. Glaziou 11850, Brazil. MinasGerais:Serrado Lenheiro, nr. Sao Joao del Rei,
22 May 1877,fl (holotype, B, destroyed;isotypes, C, K, LE, P-Herb. Glaziou).
Small shrubor tree to 12m tall with a narrowcrown and low branches;branch-
lets 2-5 mm in diam., dull darkreddish-grayish-brown,lenticels roundishor spin-
dle-shaped 0.3-1.5 mm long; young branchlets densely villous with pale tawny
hairs to 1.5 mm long, the brownish indument clearly visible to the naked eye,
becoming glabrousin age. Leaves (2-)3-foliolate with leaflets sessile; sheath sub-
orbicularor ovate, 2.5-4.5 mm long, ciliate with hairs to 0.2 mm long, densely
villous like the young branchlets,or tomentose; petiole semiterete, (deeply) can-
aliculate, 2-10(-12) mm long and 0.5-1 mm thick, the indumentlike that of the
sheath; leaflet blades obovate or sub-elliptic, (0.6-)1.2-5 x (0.4-)0.7-2 cm, base
very shortly attenuateor sometimes minutelyauriculate,or terminalleaflets nar-
rowly cuneate, obtuse at apex, marginrevolute towards base, the blades sub-
coriaceous, shining, the upper side grayish-green,becoming darker, the lower
side brown-green,minutelypubescent on both sides but particularlybelow with
tawny hairs to 0.2 mm long, (up to 0.5 mm towards base), or the upper side
glabrous except for the base, venation prominulous,midvein plane or prominu-
lous towards base above. Inflorescences terminal, paniculate, longer than the
leaves, 4-6 x 3-5 cm, many-flowered, pubescent with hairs to 0.6 mm long,
lowermost side-branchletsinserted at base of inflorescence; lower bracts some-
times leaf-like and up to 2.5 x 1.1 mm, upper bracts ovate, ca. 1.5-2.5 x 0.5-
1.1 mm or smaller, with few appressedhairs ca. 0.1 mm above, pubescent below
with hairs to 0.2 mm long; epipodium of pedicels 1.5-4 mm long and 0.5 mm
thick; bractlets2, (sub)opposite.Flowers 6-6.5 mm in diam.; calyx lobes subquin-
cuncial or subseparate, depressedly ovate, 0.5-0.8 x 0.6-1.2 mm, obtuse or
acute, pubescent below with hairs 0.05-0.1 mm long; petals deciduous, valvate,
very widely spreading,elliptic, 2.8-3.2 x 1.4-1.7 mm, unguiculatewith the claw
0.2-0.3 mm, acutish at the tip, thickly coriaceous, becoming thinly so in age,
yellowish-whitewhen fresh, brownishwhen dried, densely pubescent above with
hairs 0.05-0.1 mm long, or minutely papillose-velvety above, puberulousbelow
with hairs 0.1-0.2 mm long; filamentsusually persistent, ca. 1.6-2.5 mm long and
0.2-0.3 mm thick, rose, glabrous; anthers suborbicularto heart-shaped, 0.6-
0.7 x 0.5-0.6 mm, papillose,dark;disc 5-lobed, shorterthanthe ovaryincludingits
tubercles, 0.7-0.8 mm high and 1.5-2 mm in diam., dark, with tubercles to 0.2
mm; carpels ca. 0.5 mm, on a raised receptacle, chargedwith orbicularor ovoid,
glandulartubercles0.2-0.3 mm which are glabrousor puberulouswith hairs0.05-
0.1 mm; style before the shedding of the pollen 0.9 mm long and 0.4 mm thick,
Metrodorea 123

projecting0.5 mm beyond the tubercles, after the shedding of the pollen ca. 1.3
mm long, 0.2 mm thick and projecting ca. 0.9-1 mm, rose, glabrous; stigma
capitate, 0.2 x 0.2 mm, dark. Fruits depressedly globose, 15-20 x 18-25 mm
when closed, their loculi dorsally rounded, without apophyses, glabrous, tuber-
culate or tuberculate-muricatewith tubercles to 1 or 3 mm long, septicidally
dehiscent from some mm above the base up to the top, and loculicidallyto /3or
/? below the top; seeds unknown.
Type. Glaziou 14588, Brazil. Minas Gerais: Serra da Caraqa,nr. Ouro Preto,
Apr 1884,fl (lectotype, C; isolectotypes, B-Herb. Eichler, destroyed, photo 12489
made by F, F, NY; BR, F, G, K, LE, LY, P, P-Herb. Glaziou 2x).
Distribution.Brazil (Ceara, Bahia, and known from Glaziou collections said to
be collected in Minas Gerais and in Rio de Janeiro near Espirito Santo). The
Glaziou collections are possibly collected by Allemao in Ceara, and pirated by
Glaziou. Forests and thickets. Flowering, accordingto Glaziou (1905), Feb-Jun.
Fig. 35B.
Specimens examined. BRAZIL. Ceara: Allemdo 277 fl (P, R 6x), 277 = Herb. Saldanha 7583 fl
(R); Gifford & Fonseca G 323 fr (E). Pernambuco: Andrade Lima & Magalhdes 52-1092 fr (R). Bahia:
Gifford & Fonseca G 275 fr (E). Minas Gerais: Glaziou 15887 fr (C, K, LE, LY, P 2x). Rio de Janeiro:
Glaziou 10452 fl (C, K, P).

This species is easily recognized by its small and puberulous, sessile leaflets.
It may be confused with Helietta mollis (Miquel) Kaastra but it differs in the
presence of sheaths, and the inflorescences are coarser.
The leaves become subglabrousabove in age, as in the type, or some are
subglabrousalready when adult. Therefore, there is no reason to maintainvar.
glabrata.
Taubertreportedfruits only 15mm in diam. but he studiedthe young, immature
fruits of Glaziou 15887. I came across mature fruits (without seeds) in Gifford
& Fonseca G 323.
Gifford & Fonseca G 275 is labeled: "leaves in 2's, 3's + 4's; stems spiny";
the specimen does not show this.

4. Metrodorea nigra Saint-Hilaire, Fl. Bras. mer. 1: 81, t. 16. 1825; Walpers,
Repert. 1: 501. 1842; Henfrey, Gard. Mag. Bot. 3: 49, cum icon. 1851;
Lemaire, Jard. fleur. 2: t. 130-131, cum icon. 1852; Engler in Martius,
Fl. bras. 12(2): 150. 1874; Urban, Jahrb. K6nigl. Bot. Gart. Berlin 2:
397. 1883. Fig. 40.
Metrodorea pubescens Saint-Hilaire & Tulasne, Ann. Sci. Nat. Bot. ser. 2. 17: 139. Mar 1842;
Walpers, Repert. 1: 501. 6-8 Nov 1842; Engler in Martius, Fl. bras. 12(2): 149, t. 33. 1874,
syn. nov. TYPE. Gaudichaud 795, Brazil. Rio de Janeiro: Nr. Rio de Janeiro, 1831-1833,
bud (lectotype, P; isolectotypes, F ex Herb. Drake/Franqueville, P ex eodem (not signed by
Saint-Hilaire, with label of Richard; Engler vidit2), P (s.n., signed by Saint-Hilaire, "Cueille
pres R. J. par Gaudichaud")).
Metrodorea atropurpurea Fischer ex Lemaire, Fl. Serres Jard. Eur. 4: t. 337, cum icon. 1848;
Walpers, Ann. 2: 247. Jan 1852; Lemaire, Jard. fleur. 2: t. 130-131, cum icon. before 15 Jul
1852; Payer, Trait6 d'Organogenie t. 22. 1857; Koch in Koch and Fintelman, Wochenschr.
Gartnerei Pflanzenk. 2: 151. 1859; Engler in Martius, Fl. bras. 12(2): 150, pro syn. TYPE.
Lemaire, Fl. Serres Jard. Eur. 4: t. 337 (lectotype).

2
Erroneously reported as no. 799 by Engler in Martius, Fl. bras. 12(2), who misread the old French
"5." The real Gaudichaud 799 was identified by Richard too. This specimen (at P) is accompanied
with a separate number label. It represents a different taxon, however.
124 Flora Neotropica

FIG. 40. Metrodorea nigra. A, habit (Glaziou 4780, C). B, habit (F. C. Hoehne SP 3456, GH).
C, flower (Glaziou 4780, C). D, fruit (Lindeman & de Haas 1699, U).
Metrodorea 125

Pilocarpusatropurpureus(Fischerex Lemaire)Koch in Koch and Fintelman,Wochenschr.Gart-

nerei Pflanzenk.2: 152, 419. 1859.
Metrodoreanigra Saint-Hilairevar. brevifoliaEnglerin Martius,Fl. bras. 12(2): 150. 1874,syn.
nov. TYPE. Sellow (B) 2172, Brazil. MinasGerais:Fda da Galena, 1818,fl (lectotype, K ex
B, photo NS 2869 madeby NY, NY; isolectotypes, B-Herb. Kunth,destroyed,photo 12486
madeby F, F, NY; K, LE ex B, PR, S (collected underno. 563), US ex Manilla,ZT, all ex
B; Sellow B 2173 (probably wrongly indicated for B 2172, see photo 12486 in F, NY),
probablyisolectotype, GH ex B).
Metrodoreaselloana Englerin Englerand Prantl,Nat. Pflanzenf.3(4): 160.Mar 1896;Bot. Jahrb.
Syst. 21. Beibl. 54: 29. 12 May 1896, syn. nov. TYPE. Sellow 144, Brazil. Rio de Janeiro:
Nr. Rio de Janeiro,fl (lectotype, B, destroyed, photo 12490made by F, F, NY).
Metrodoreabrevifolia(Englerin Martius)Engler[in Englerand Prantl,Nat. Pflanzenf.3(4): 160.
Mar 1896,nom. nud.], Bot. Jahrb.Syst. 21. Beibl. 54: 29. 12 May 1896;Glaziou,Bull. Soc.
Bot. France 52. Mem. 3: 85. 1905.

Shrub or small tree to 6 or 8 m tall, trunk 10.5-30 cm in diam.; branchlets2-

5(-10) mm in diam., brown, shining, lenticels spindle-shapedor fissured, up to
more than 5 mm long, (sub)glabrous.Leaves 1-3-foliolatewith leaflets subsessile;
sheath to 7 mm long, ciliate with hairs to 0.5 mm, densely pubescent, becoming
glabrousin age; petiole subterete, 10-60 mm long and 1-3 mm thick, rarelyshort-
er, usually glabrous, or pubescent with spreadinghairs to 0.05 mm long; wings
obsolete or up to 0.1-0.2 mm broad;leaflet blades (narrowly)elliptic or obovate,
(3-)5-20(-28) x (1.2-)2-7.5(-13) cm, mostly up to 13 x 5 cm, (long-) attenuate
to narrowlycuneate at base, acuminateto, occasionally, obtuse or emarginateat
apex, the very tip obtuse or occasionally emarginate, margin revolute toward
base, the blades chartaceous or somewhat subcoriaceous, somewhat shining,
mostly glabrous on both sides, or pubescent below with spreadinghairs to 0.3
mm long and soft to the touch, venation ? prominent,particularlybeneath, mid-
vein plane or prominulentabove. Inflorescences terminalnear tips of branchlets,
widely paniculate, in general longer than the leaves, (10-)20-35 x 2-30 cm, with
numerousflowers, pubescent with spreadinghairs0.05-0.2 mm long, lower bracts
sometimes leaf-like and up to 10 x 2 mm, upperones usually narrowlytriangular,
0.8-2(-5) x 0.5-1.2 mm, usually ? densely pubescent on both sides with hairs
to 0.1 mm long, or subglabrousbelow; pedicels 1.5-7 mm long and 0.4-0.7 mm
thick, pubescent with spreadinghairs 0.05 mm; bractlets 2, subopposite. Flowers
(6-)6.5-10(-12) mm in diam., especially the buds becoming very dark purplish
when dried; calyx lobes separateto subvalvate, depressedly ovate or -triangular,
0.5-1 x 0.7-1.3 mm, obtuse, upper side glabrous or sometimes near marginpu-
bescent, lower side subglabrousor pubescent with spreading hairs to 0.1 mm
long; petals deciduous, valvate, very widely spreading, ovate or elliptic, 3-5
(-5.5) x 1.5-2.5(-3) mm, unguiculatewith claw 0.2-0.5 mm, acute at apex, thick-
ly coriaceous becoming thinly so, claret or dark purplishwhen living, brownish-
or red-purplishwhen boiled, the upper side densely pubescent with spreading
hairs to 0.1 mm long, the lower side glabrous, or towards the tip pubescent like
the sepals; filaments deciduous or persistent, 0.8-1.5 mm long and 0.1-0.2 mm
thick, the tip acuminatebut later on subobtuse due to shrinking,purplishin the
upper half, glabrous;anthers very broadly heart-shaped,0.5-0.8 x 0.5-0.8 mm,
finely purplish-pigmentedand papillose; disc in upper half 5-partite, each part
slightlyor obsoletely lobed, no longerthanthe ovary, ca. 1-1.2 mmhighand 1.7-2.1
(-2.5) mm in diam., purplish, chargedwith tubercles up to 0.1 mm; carpels 0.6-
0.8 mm, usually on a raised receptacle, provided with orbicularor ovoid, glan-
dulartubercles 0.1-0.2 mm, which are usually glabrous,or pilose with hairs0.05-
0.1 mm long; style inserted half-wayon the carpels, not or only slightlyprojecting
beyond them before the shedding of the pollen, but after that slightly elongating
126 Flora Neotropica

to become 0.5-0.8 mm long, 0.2 mm thick, and projectingup to 0.5 mm, becoming
purplish, with glands, glabrous; stigma capitate, slightly lobed or not, 0.2-
0.3 x 0.3-0.4 mm. Fruits somewhat depressed, stellately-lobed, 14-20 x 22-30
mm, up to 35 mm wide when dehisced, loculi provided with a dorsal, blunt
apophysis 3-5(-7) mm long 1/2 or 2/5 below the tip, transversely wrinkled with
prominentparallelnerves on both sides of the wall, + muricate-tuberculate,the
tubercles to 2 mm long, septicidally dehiscent from some mm above base to 2
mm below top, and loculicidallyup to half-way the apophysis; seeds 1 or 2 per
loculus, (depressedly) ovoid, or roundish, 5-8.2 x 4-6 x 4-6 mm with a mucro
0.2 mm, flat at or near base; testa coriaceous, dull dark brown, often provided
with slight longitudinal pustulate ridges; chalazal area roundish, ca. 3-4 mm
broad, dark brown-black,shining;hilum ca. 0.6 mm broad; cotyledons strongly
unequal, not punctate, with ears 0.5 mm, radicle thick, 0.4-0.7 mm long, obtuse,
projectingbut not exceeding the cotyledons, plumule0.2-0.3 mm long.
Type. Saint-Hilaire669, Brazil. Rio de Janeiro:Nr. Rio de Janeiro, "dans le
lit du ruisseau qui tournet ses eaux a l'aqueduc," Nov 1816, fl (lectotype, P;
isolectotypes, F ex P ex Herb. de Bunge, P-Herb. Cosson ex Herb. de Bunge
(sine coll. et num.), P-Herb. Saint-HilaireFl. Bras. mer. (det. Tulasne), P (det.
Tulasne)).
Distribution. Brazil, S Piaui to Parana. Dry places in woods, capoeirao, and
gallery and rain forests; on red sand (1 record); alt. up to ca. 500 m. Flowering:
in the northernstates Nov-Feb(-Mar), in the southernstates Aug-Feb. Fig. 35B.
Specimens examined. BRAZIL. S Piauf: Liiutzelburg13058 fl (M). Bahia: Blanchet 2378 fl (G, P);
Campos Porto RB 20481 fl (BM, IAN); Fr6es 33 (in Krukoff 12667) fl (A, DS, NY), 19971 fl (NY),
19983 fl (K 2x); Pinheiro 1251, fl (U). Minas Gerais: Burchell 3582 fl (K, L, OXF, P); Glaziou 13651
fl (BR, C, F 2x, G 2x, K, LE, LY, P 3x, US), 14589 fl (BR, C, G, K, LE, LY, P 3x); Regnell I 273'
fr (LD, S 2x); Widgren s.n. (1845) fr (S). Espirito Santo: Wied-Neuwied s.n. (Brasilia) fl (LD), s.n.
(Capitania) fl (BR). Sao Paulo: Amaral SP 35609 fl (SP); Andrade EFSP 24 fl (SP); Burchell 5051 fr
(K); Comm. Geogr. Geol. S. P. 140 fl (C); Edwall Exp. R. F. 60 fr (P, SP); Helmreichen 17 fl (NY,
W); Hemmendorf 43 fl (S); W. Hoehne SP 54147 fl (SP); Hunger Filho s.n. (Navarro de Andrade, Jul
1928) st (P); M. Kuhlmann 728 fr (SP); Lofgren CGGSP 34 fr (C, P, SP), CGGSP 946 fl (C, P, SP),
CGGSP 1390 fl (P, SP); Lofgren & Edwall CGGSP 2729 fl (SP), CGGSP 2740 fl (SP); Lund s.n. (Feb
1834, Herb. Warming 2462) (pro parte) fr (C); Herb. Martius 79 fl (BR, E, G, K, L, W); Mosen 2444
fr (S), 2804 fl & fr (BR, C 2x, K, LD, LE, M 2x, P, S 2x), 3165 fl (S), 4061 fl (BR, C, K, LD, LE,
M 2x, P, S 5x); Novaes 521 fl (US); Pacifico CEFSP 280 fl (SP); Pickel 5450 fl (US); Sellow 563
(2172.c 2169) fl (ZT 2x); Viegas IAC 2376 fl (IAN); Viegas et al. IAC 3000 fl (SP). Rio de Janeiro:
Anonymus s.n. (liheos, 1855) fl (S); Comm. Th. Bernhardi s.n. (1886) fl (L); Casaretto 541 fl (G, TO);
Constantino RB 2549 fl (K, S, U, US); Duarte 5026 fl (LP); Ducke & J. G. Kuhlmann RB 16511 fl
(S, U); Dusen s.n. (Herb. d'Aleizette, "Feb 1904") fl (L); Glaziou 1391 fr (BR 3x, C, P 2x), 2526 fl
(BR 2x, C 2x, K, P 2x), 4780 fl (C 2x, K, LY, P 2x), 11851 fl (C, K, P), 11852 fl (C, K, LE, P);
Guillemin 1033 fl (G 2x, P 2x); F. C. Hoehne 318 (SP 24859) fl (SP); Luschnath s.n. (Rio, "fruct. nr
1") fl (OXF), s.n. (Rio, 1833, "fruct nr 36") fl (LE); McLean R 79 fl (BM); Miers 3370 fl (BM, K 2x),
4485 fl (K 2x, P), s.n. (Organ Mts, Dec 1837) fl (BM), s.n. (Rio de Janeiro) fl (F, HBG, MO, P 2x,
S, US), s.n. (Corcovado) fl (BM); Mikan s.n. fr (W); Nadeaud s.n. (7 Nov 1862) fl & fr (A, P 7x,
US); Netto 117 fl (S); Pereira 4217 fl (F, M); Pohl Herb. Bras. 27 fl (W); Riedel 54 fl (L), 469 fl (C,
LE 2x, M), s.n. (Monte Tijuca, 1836) fl (P, U), s.n. fl (E 2x, G, GH, GOET, K, L, M, NY, P 3x, S,
US, W, ZT); Schwacke 1535 fl (GOET); L. B. Smith 1238 fl & fr (GH, S, US); Ule 3556 fl (HBG);
Widgren 483 fl (S 2x), 1077 fl (BR). Parana: Braga & Lange 78 fl (US); Hatschbach 12937 fl & fr (F,
K 2x, US), 16935 fl & fr (K), 16948 fl (MO, NY 2x, S, UC), 17006 fl (C), 17055 fl & fr (F), 21517 fr
(MICH, NY); Lindeman & de Haas 577 st (U), 1525 st (U), 1699 fr (K, NY, U), 5524 fr (U, WIS).
State unknown: Anonymus 150 fl (W); Bowie & Cunningham s.n. fl (BM); Comm. Martius I (Sebas-
tianopolis, 1841) fl (GOET); Herb. Richard 138 fl (P); Schiicht s.n. fl (W); Sellow 676 fl (BM), 1406
fl (K), s.n. fr (F, NY (on same photo as Sellow 144)); Herb. Swartz s.n. fl (S); Tube 5998? fl (W), s.n.
(Apr 1871) fl (W); Wied-Neuwied s.n. (comm. 1828) fl (BR).
CULTIVATED. F. C. Hoehne SP 3456 (Hort. Butantan) fl (GH, NY, SP, US); Hort. BO 15 fr
(MPU); VII E 6 fl (L); Hort. C s.n. (Jan 1876) fl (C); Hort. K s.n. (1854) fl (LD), s.n. (1856) fl (K),
s.n. (Apr 1862) fl (K); Hort. W s.n. (1861) fl (W), s.n. (1862) fl (W), s.n. (1864) fl (W 2x), s.n. (22 May
Metrodorea 127

1875 or 1878?) fl (W); Koscynsky 184 (Horto Fl. Sao Paulo) fl (SP); Herb. Magnus s.n. (Hort. B, 20
Mar 1867) fl (HBG); Th. Moore s.n. (Hort. Chelsea, Dec 1852) fl (K); Riedel s.n. (Hort. LE) fl (US);
Vogel s.n. (30 Jan 1897) fl (W).

Local names and uses. Bahia: vira sarere (Pinheiro 1251); Sao Paulo: chupa-
ferro (several records in SP); Parana: carrapateiro (several records). Wood hard
(Lifgren CGSP 946).
The presence of a (short) petiole is a good character for distinguishing this
species from Metrodorea stipularis. Some specimens have some or all petioles
ca. 1-2 mm long. In other respects however, they are clearly referable to M.
nigra.
The color of the flower, the disc, the ovary, the slender inflorescence, etc., are
quite distinct from those of M. maracasana, the one species of Metrodorea with
short petioles.
Metrodorea selloana is not different, according to the protologue and the photo
of the type. The shortly stalked, large leaves and the inflorescences come within
the range of M. nigra. The type collection was originally identified by Engler as
M. nigra. In 1895 he changed the name to M. nigra var. brevifolia.
The inflorescence varies in size. Very large ones with side-branchlets e.g. to
20 cm long occur in Rio de Janeiro (Guanabara) and Sio Paulo. These specimens
belong to M. nigra var. nigra. Smaller inflorescences with side-branchlets to 9
cm (usually ca. 2 cm) were collected in Sao Paulo and in the other states of Brazil
but not in the former Guanabara; these plants represent M. nigra var. brevifolia.
Because there are no other differences, and intermediate inflorescences exist, I
reduce M. brevifolia to synonymy with M. nigra.
Hort. C s.n. (1876) (C) has inflorescences with hairs to 0.5 mm long. Two
specimens from a tree cultivated in Hortus BO have coriaceous leaves with their
midvein slightly impressed.
Ripe seeds are rarely found in herbaria.

5. Metrodorea stipularis Martius, Flora 20(2). Beibl. 124. 1837; Herb. fl. bras.
124. 1837; Walpers, Repert. 1: 501. 1842; Engler in Martius, Fl. bras.
12(2): 149. 1874, pro syn. Fig. 41.
Metrodorea pubescens auct non Saint-Hilaire & Tulasne.

Tree 4-15 m tall with trunk ca. 10-50 cm in diam.; wood white without heart-
wood (according to a label); subverticillately branched, glands bulging and often
conspicuous; branchlets 3-6 mm in diam., dark brown, shining, lenticels roundish
or spindle-shaped 5-7 mm long or fissured, glabrous or distally pubescent with
spreading hairs to 0.6 mm long, becoming glabrous in age. Leaves (2-)3-foliolate,
with leaflets stalked; sheath ca. 10-15 mm long, ciliate and pubescent with brown
hairs to 0.5 or 0.7 mm, becoming glabrous in age; petiole seemingly absent;
petiolules inserted half-way on the sheath, 0-5 mm long and 2-3 mm thick,
mostly glabrous; leaflet blades (narrowly) obovate, occasionally elliptic, (4-)8-
25(-31) x 2-10(-12) cm, gradually narrowing towards the narrowly cuneate or
long-attenuate base, obtuse or subacuminate at apex, the margin slightly undulate
and revolute towards base, the blades subcoriaceous, somewhat shining, glabrous
or pubescent at base, the midvein plane or impressed above, with hairs 0.05-0.2
mm long, venation prominulous below. Inflorescences terminal, occasionally also
axillary near tips of branchlets, widely paniculate, no shorter than the leaves, 15-
30 x 11-20 cm, with numerous flowers, pubescent with spreading hairs to 0.4
mm, lowermost side-branchlets subtended by the uppermost leaves; lower bracts
128 Flora Neotropica

olo
o

FIG. 41. Metrodorea stipularis. A, habit (Regnell II 54, Mar 1875, S). B, insertion of leaflets
(Regnell II 54, 19 May 1870, S). C, fruit (Irwin et al. 15603, U). D, seed (Irwin et al. 15603, U).
Metrodorea 129

sometimes leaf-like and up to 18 x 3 mm, upper ones triangular,to 3-5 x 1.5-2

mm, the upper side glabrous or minutely pubescent towards tip, the lower side
minutely pubescent with hairs 0.1 mm long; pedicels to 2 mm long and 0.2 mm
thick; bractlets 2, alternate. Flowers 5.5-7.5 mm in diam., very fragrant(Warm-
ing 2467/11);calyx lobes separateor, less frequently,subquincuncial,very broadly
to depressedly ovate, 0.7-1.3 x 0.9-1.2 mm, obtuse or acuminate, acute at very
tip, glabrous above, usually minutely pubescent below with hairs 0.05-0.1 mm;
petals persistent, valvate, adnateto the disc at base, very widely spreading,ovate
or elliptic, (2.5-)3-3.8 x (1-)1.2-1.7 mm, unguiculatewith claw to 0.3 or 0.4 mm,
recurvedand acute at apex with minutelyapiculatetip, coriaceous,becomingthinly
so and semitransparentin age, yellowish-whitewhen fresh, yellow-brownishwhen
dried, minutelypubescenton both sides with hairs0.05-0.1 mm long or glabrousin
lower half above, occasionally glabrousbeneath;filamentspersistent, 1.3-1.7 mm
long and 0.2-0.4 mm thick, glabrous; anthers broadly heart-shaped, 0.4-
0.6 x 0.5-0.6 mm, when fresh minutelypunctate with pink; disc 5-lobed, as high
as the ovary includingits tubercles, ca. 0.5-0.8 mm high and 1.5-2 mm in diam.,
thick at margin,rose-violet, beset with few tubercles 0.05-0.1 mm; carpels 0.5-
0.6 mm high, charged with obconical tubercles 0.3 mm or some up to 0.6 mm
after the shedding of the pollen; tubercles at tip filled with glands, pilose with
hairs 0.05-0.1 mm, rarelyglabrous;style inserted near tips of carpels, ca. 0.5 mm
long before the shedding of the pollen, afterwards(still at anthesis) ca. 0.9 mm
long and 0.2-0.3 mm thick, projectingca. 0.5 mm beyond the tubercles, glabrous;
stigma capitate, 5-lobed or entire, 0.1-0.2 x 0.3-0.5 mm. Fruits depressedlyglo-
bose, stellately-sublobed,25-30 x 35-45 mm (-55 when dehisced), loculi provid-
ed or not with a short apophysis, very thick-walled,densely chargedwith coarse,
blunt, warty tubercles up to 12 mm long, dehiscent septicidally with a fissure
only, and loculicidally to 13 or /? below the top; seeds (in 1 collection seen only)
2 per loculus, ovoid or roundish,4.5-7 x 3-3.5 x 3.5-5 mm, with roundedapex,
at base not flattenedor slightly so; testa coriaceous, dull darkbrown, flatly collic-
ulate-reticulate;chalazal area roundish, slightly impressed, 0.7-1.3 mm broad;
caruncle black, up to 0.4 mm in diam.; cotyledons unequal with ears to 0.6 mm
long, not punctate; radicle projectingbeyond the ears, 0.5-0.7 mm long, thick,
plumule 2-leafed, ca. 0.3 mm long.
Type. Martius Herb. Fl. Bras. 181 (A. L. P. da Silva Manso 245), Brazil. Mato
Grosso: Nr. Serra de Cuiabai, fl (lectotype, W; isolectotypes, BR (Manso
245), G).
Distribution.Brazil, Mato Grosso, Distrito Federal, Minas Gerais, Sao Paulo,
and Rio de Janeiro.Locally common, forests, probablydry places; alt. up to 1000
m. Flowering Oct-Jan(-Mar). Fig. 35B.
Specimens examined. BRAZIL. Mato Grosso: Manso s.n. (Cuiabd, 1835) fl (BR); Goias: Burchell
6240 fr (K); Pohl s.n. fl (BR). Distrito Federal: Irwin et al. 15603 fr (U), 18048 fr (U); Pires et al.
9496 fr (US). Minas Gerais: Barreto 6104 fr (F); Glaziou 18173 fl (BR, G, K, LE, LY, NY, P 2x),
18179 fl (C); Helmreichen 64 fl (F, M, NY, US, W), s.n. fl (BR); Regnell 11 54 (5 Dec 1845) fl (K, LE,
P, U, US), (6 Dec 1868) fl (BR, M, P 2x), (17 Feb 1869) fr (C, LD), (May 1869) fr (S), (9 Mar 1870)
fl/fr (S), (19 May 1870) fr (P, S 2x), (Mar 1875) fl (S); Sellow 2170 fl (K, S); Warming 510 (herb. 2467/
2) st (C, P), 549 (herb. 2467/3) fr (C), 869(?) (herb. 2467/1) fl (C), s.n. (2 Jan 1865) fl (LE), s.n. st (C
2x); Widgren s.n. (1845) st (BR, S). Sao Paulo: Anonymus 29 (SP 19838) fr (SP); Comm. Geogr. Geol.
S. P. 92 fr (C); Frazdo RB 15258 fl (S); Glaziou 12526 st (C, P, US); Heiner 334 fl (S); F. C. Hoehne
SP 1934 fr (SP), SP 29729 fr (F, NY 2x, SP); Hunger Filho s.n. (Navarro de Andrade, Jul 1928) st
(P); M. Kuhlmann 1160 fl (SP); Lofgren CGGSP 1342 fr (SP); Mosen 1185 fr (LD, S 2x), 4060 fl (P,
S 3x); C. Novaes CGGSP 3799 fl (SP); Pickel 5551 fl (US); Riedel 3 fl (LE), 126 fl (LE, M); Vecchi
CPEF 162 fl (SP). Rio de Janeiro: Campos Porto RB 15255 fl (U). State unknown: Helmreich(en?)
s.n. fl (BR, G 3x, LE, NY 2x, US 2x); Riedel s.n. fl (E, GH, K, P), s.n. fr (K); Sellow s.n. fl (K).
130 Flora Neotropica

Local names and uses. Minas Gerais: limoeira-do-mato,laranjeira(-do-mato),

bananeira-do-mato(Sic!) (several records of Glaziou and of Warming);arco-de-
pipa (Barreto 6104). Siio Paulo: catagua (Comm. Geogr. Geol. S. P. 92); caputuna
(F. C. Hoehne 29729); chupa-ferro (F. C. Hoehne SP 1934, confused with Me-
trodorea nigra?). For building,as filling-and firewood;possibly also for barrels,
cf. "arco-de-pipa."For medicinaluses see PHYTOCHEMISTRY.
The species differs from all others in the lack of petioles.
As a result of a misinterpretationby Engler (1874a)nearly all collections were
identifiedas Metrodoreapubescens Saint-Hilaire.Martius'name of 1837must at
least have priorityupon M. pubescens (1842) but it has now turned out that the
type of M. pubescens belongs to M. nigra!

Insufficientlyknown
Metrodoreaexcelsa Allemao ex Th. Peckolt, Ber. Deutsch. Pharm.Ges. 9: 339.
1899.
This species, called amare or amari, occurs in Ceara and Alagoas. From the
descriptionit is not clear whetherit belongs to Metrodoreaor not, but it is surely
not referable to any of the known species. For practical purposes I give here
the complete description, which occurred in the Heil- und NutzpflanzenBrasi-
liens: "Baum bis 30 m hoch, Stamm 30 bis 45 cm Durchmesser, mit grossen
dreibldttrichen, lederartigen, verkehrt einformig-lanzettlichen, oberseits kahlen,
unterseits rostfarben, haarigen Blatter, grosse zusammengesetzte Rispe mit weis-
sen befilzten Bliiten. Kapsel holzig, hockerig. Bitterschmeckende Rinde zur
Wechselfieber. "

Excluded from Metrodorea

Metrodoreagracilis Schumann,Bot. Jahrb.Syst. 30. Beibl. 67: 30. 1901. =Esen-
beckiafebrifuga (Saint-Hilaire)Jussieu ex Martius(see page 103).

RAULINOA
3. RaulinoaCowan, Sellowia 12: 90. 1960;Cowan and Smith in Reitz, Fl. Ilustr.
Cat. I (RUTA) 47. 1973.
Branchlets with spindle-shaped lenticels, frequently armed with opposite
branch-spines.Leaves opposite, simple, stalked; venation of the blades brochi-
dodromous, its loops formingan intramarginalvein by joining the superadjacent
secondary one at an obtuse angle. Inflorescences lateral, axillary on branchlets
and spines, greatly reduced in size, cymose, raceme-likeor subfasciculate;flow-
ers tetramerous,slightly zygomorphic;filamentssubulate;anthers mucronateby
the protrudingconnective; disc slightly tetragonalwith antefilamentalincisions;
carpels 4, connate, with a free apophysis.
Type species. Raulinoa echinata Cowan. The name was given to honorRaulino
Reitz, the editor of the Flora IlustradaCatarinense.A photographof PadreReitz,
made at the type locality, is given in Cowan and Smith (1973).
Distribution.Brazil, endemic in Santa Catarina(one species known).

1. Raulinoa echinata Cowan, Sellowia 12: 90, t. 4, fig. a-d. 1960; Cowan and
Smith in Reitz, Fl. Ilustr. Cat. 1 (RUTA) 48, t. 13, fig. a-d, tab. 1-2.
1973. Fig. 42.
Shrub 2-3 m tall, glabrous except for young petioles; branchlets 2-5 mm in
diam., grayish-brown,especially fast-grownbranchletsarmedwith straight,slen-
Raulinoa 131

FIG. 42. Raulinoa echinata. A, habit (Reitz & Klein 7359, U). B, flower (Reitz & Klein 7359,
US).

der spines 0.7-2.5 cm long, which bear flowers and leaves near the base. Petiole
semiterete, 2-3 mm long and ca. 0.7 mm thick, not sharply separated from base
of blade, minutely pubescent when young with hairs 0.05 mm long; blade narrowly
obovate, 2.2-6.5 x 0.6-1.7 cm, attenuate-cuneate at base, rounded at apex, the
very tip emarginate or entire, margin subrevolute, the blade subcoriaceous, shin-
ing, grayish-green, venation prominulous, costa plane above and prominent be-
low. Inflorescences erect or spreading, to 8 mm long and wide, 1-5-flowered;
peduncle to 1 mm long, side-branchlets (sub)opposite; bracts and bractlets ovate
to triangular, ca. 0.5-1 mm long; pedicels to 2.5 mm long. Flowers 8-9.5 mm in
diam., glabrous; sepals imbricate, overlapping at anthesis, nearly free or connate
at base, very broadly ovate, 1-1.7 x 1.2-1.7 mm, the innermost two smaller than
the others, rounded to obtuse at apex, coriaceous with papery margins, venation
parallel, the nerves branching distally, with a dorsal false midrib; petals imbricate,
reflexed, (broadly) elliptic, 3.5-4.7 x 2.5-3 mm, the innermost ones smaller than
the others, obtuse to rounded at apex, subcoriaceous with papery margin, dark
red, papillose above, venation parallel to actinodromous, the nerves somewhat
branching distally; filaments persistent, seemingly inserted on the disc but ac-
tually completely surrounded at base by the disc, reflexed, I mm long and 0.2-
0.3 mm thick, fleshy; anthers dorsifixed just below middle, versatile, very broadly
heart-shaped, 0.7 x 0.7 mm including a mucro 0.05 mm; disc annular to cup-
132 Flora Neotropica

shaped, equallingthe carpels or lower, becoming0.8 mm high and 2 mm in diam.,

fleshy towards tip, rose to dark purplish, smooth or with few protuberancesca.
0.3 mm long; carpels immersed in the receptacle with their lower half, 0.8 mm
high includinga trigonous apophysis 0.4 mm, the apophyses beset with ca. 3-4
protuberances0.05-0.3 mm; ovules 2 per loculus; style persistent, inserted on
the connate part of the carpels among the apophyses, ca. 0.9 mm long and 0.3
mm thick, projecting 0.3-0.6 mm beyond the apophyses at anthesis; stigma
4-lobed, 0.2-0.3 x 0.4 mm. Fruits 1 per infructescence, (3-)4-locular capsules,
stellately-lobed,0.8-1 x 1-1.2 cm, with glands, glabrous,the exocarp transverse-
ly nerved on both sides, the loculi ? obovate in outline, rounded at apex, with
a dorsal apophysis 0.5-2 mm long, projectingdownwardjust above the middle,
dehiscent septicidally from base up to tip and loculicidally from apex up to the
tips of the apophyses; seed 1 per loculus, ovoid, ca. 6 x 3.5 x 3.5 mm, flattened
at base, curved at apex, irregularlywrinkled, finely reticulate-colliculate;hilum
ca. 0.7 mm broad; embryo 1, cotyledons unequal with ears 0.2-0.3 mm, radicle
conical, projectingbeyond the ears.
Type. Reitz & Klein 3828, Brazil. Santa Catarina:"Indaial" (=Apiuina),Su-
bida, 11 Oct 1956,fl & fr (holotype, US-2278676;isotypes, BR, G 2x, HBR (n.v.),
K, L, M, NY, UC, Z).
Distribution. Known only from the type locality. Frequently inundatedriver
banks; alt. 100 m; collected in flower and in fruit in Oct. Fig. 35C.
Specimens examined. BRAZIL. Santa Catarina: Apiiina, Subida, Reitz & Klein 7359 fl & fr (BR,
GH, K, L, U, UC, US 2x, WIS); Smith & Reitz 12410 fl & fr (C, GH, MO, NY, P, UC, US).

PILOCARPUS
4. PilocarpusVahl, Eclog. 1: 29. after Mar, 1797("1796") Willdenow,Linn. Sp.
pl. 1: 1133. 1798;Jaume Saint-Hilaire,Expos. fam. nat. 2: 356. 1805;
Roemer and Schultes in Syst. veg. 5: 472. 1819/1820;Nees and Mar-
tius, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 11:
176. after 26 Jul, 1823; Saint-Hilaire,Bull. Sci. Soc. Philom. Paris
(1823): 130. Sept 1823;de Candolle, Prodr. 1: 728. 1824;Saint-Hilaire,
Fl. Bras. mer. 1: 82. 11 Jun 1825;Adr. Jussieu, Mem. Mus. Hist. Nat.
12: 488. after27 Jun, 1825;Spach, Hist. nat. veg. 2: 344. 1834;Meisner,
P1.gen. 1: 63 and 2: 45. 1837;Endlicher,Gen. pl. 1153. 1840;van Hall,
Ann. Hort. Bot. 3: 113. 1860;Hooker in Bentham and Hooker, Gen.
pl. 1: 299. 1862;Engler in Martius, Fl. bras. 12(2): 131. 1874;Engler
in Engler and Prantl,Nat. Pflanzenf.3(4): 157. 1897;Wilson, in North
Amer. Fl. 25: 199. 1911;Engler in Engler and Prantl,Nat. Pflanzenf.
(ed. 2) 19a: 278. 1931;Macbride,Field Mus. Nat. Hist., Bot. Ser. 13:
675. 1949; Cowan, Sellowia 12: 85. 1960;Albuquerque,Anais Acad.
Brasil. Ci. 40: 506. 1968; Cowan and Smith in Reitz, Fl. Ilustr. Cat. 1
(RUTA) 24. 1973.
Pre-Linnean synonyms:
Euonymus lati-folius, racemosus, fructu pentagono, atropurpureo Plumier, Cat. pl. amer. 18.
1703 (as "Evonymus"); Burman, PI. amer. 119. 1757 (as "Euonymus latifolius," etc.), non
Tournefort.
Prunus fioribus racemosis Burman, PI. amer. t. 127. 1757, non Linnaeus.

Branchlets with roundish or spindle-shapedlenticels usually to 1 mm long;

terminalbuds covered with large concave perules. Leaves alternateto subverti-
cillate by crowdingat the tips of the branchlets,simple or divided;apex of leaflets
or leaves mostly emarginate.Peduncle of racemes rudimentaryor lacking; ped-
Pilocarpus 133

icels alternate, adaxially incurved when in bud. Flowers pentamerousto tetra-

merous in P. spicatus; calyx toothed or lobed, appressed but remote from the
petals when dried;petals deciduous, coheringwith their hooked tips when in bud,
reflexed when flowering, ovate, with the inner margin thickened and (slightly)
induplicate, the apex acute and uncinately inflexed, carinate above and often
impressed next to the keel; filaments deciduous, accumbent in shallow grooves
of the disc, inflexed over the ovary but finally usually spreading;anthers dorsi-
fixed mostly near middle, versatile, or immobile and recurved on the back after
the shedding of the pollen, yellow; disc annularto somewhat cup-shaped, com-
pletely adnate to the ovary, with internalglands dischargingon the outside and
often some of them confluentto a circularcanal;carpels not immersed,frequently
borne on a globose gynophore, connate at base, free distally; ovules 1 or 2 per
carpel; style inserted with 5 strands, pentangular,often rudimentarybefore the
shedding of the pollen, stigma 5-lobed with a gland in each lobe. Fruits divided
into mericarps, 1-4 of which usually rudimentary,the mericarpsunited at base
only, conchate with flattened sides, rounded dorso-basicallyand mostly dorso-
apically too (except for P. pennatifolius), frequently mucronate at apex by the
persistent bases of the strands of the styles, prominentlytransversely arcuate-
parallel-nervedinternally,and externallymostly too (except for P. trachylophus),
dehiscent loculicidally from the base along the ventral sutures over the tip and
dorsallydown to 13 or 14 above base; seeds usually kidney-shaped,slightlykeeled
dorsally or sometimes double keeled; testa thinly coriaceous, shining; embryo
usually 1 (except for P. pauciflorus and P. riedelianus), densely punctate.
Type species. Pilocarpus racemosus Vahl.
Distribution.Mexico to Argentina,rare in the Amazon basin, see Fig. 4B.

Key to the Species of Pilocarpus

1. Leaves compound.
2. Leaves exclusively 3- or 1-foliolateor imparipinnate1-jugate.
3. Leaves (densely) pilose; flowers 9-11 mm in diam. 8. P. goudotianus.
3. Leaves (sub)glabrous;flowers 7-9(-9.2) mm in diam.
4. Midribof leaflets impressed;petals reddish-brown;filamentssubulate.
7. P. racemosus.
4. Midribof leaflets plane; petals yellowish-green;filamentstruncate.
13. P. pauciflorus.
2. Leaves imparipinnateor mostly so.
5. Leaflets sessile.
6. Wingsof petioles to 0.3 mm broad, not eared;lateralleaflets narrowlyoblong
or narrowlyelliptic; midveinplane above. 5. P. trachylophus.
6. Wings of petioles 0.5-1 mm broad, eared; lateral leaflets ovate to elliptic;
midvein prominentabove. 6. P. microphyllus.
5. Leaflets stalked.
7. Apex of leaflets acuminate,the very tip emarginateor retuse; mericarps15-
17 mm long. 2. P. demerarae.
7. Apex of leaflets roundedor obtuse, the very tip emarginateor retuse; meri-
carps 9-13 mm long (of P. grandiflorusunknown).
8. Base of lateral leaflets rounded, obtuse, or cuneate, rarely some of a
specimen shortlyattenuate,stronglyunequal.
9. Leaflets sessile or shortly stalked, with curved hairs;racemes erect,
firm; pedicels 1-1.5 mm long; fruits tuberculate,externally without
arched nerves. 5. P. trachylophus.
9. Leaflets stalked, glabrousor with + straighthairs; racemes pendu-
lous, slender;pedicels 7-16 mm long; fruits without tubercles, with
arched nerves. 4. P. jaborandi.
8. Base of lateralleaflets attenuate, + unequal.
10. Leaves 5-6-jugate;flowers ca. 13.5 mm in diam., hairy.
3. P. grandiflorus.
134 Flora Neotropica

10. Leaves 1-3-jugate,or some 1-3-foliolateor simple;flowersto 11 mm

in diam., (sub)glabrous.
11. Midvein of leaflets impressed above; leaves usually 3-jugate,
sometimes 1-, 2-, or 4-jugateor some 1-foliolate;flowers 8.5-11
mm in diam.; mericarpsdorso-apicallyblunt, usually with an an-
gle of ca. 90?,apex truncate,the very tip mucronate. 1. P. pennatifolius.
11. Midvein of leaflets plane above; leaves 1-3-jugate, sometimes
some simple or 1-3-foliolate;flowers 7.5-9(-9.2) mm in diam.;
mericarpsdorso-apicallyrounded, apex obtuse or rounded, the
very tip slightly mucronate. 7. P. racemosus.
1. Leaves simple.
12. Leaves taperinginto narrowlycuneatebase, mostlylongerthan20 cm; racemeslateral.
9. P. giganteus.
12. Leaves not tapering,attenuateto narrowlycuneate at base, mostly shorterthan 20
cm; racemes (sub)terminal.
13. Leaves bullatebetween secondarynerves; costa ? prominentabove.
10. P. peruvianus.
13. Leaves plane between secondarynerves; costa plane or longitudinallywrinkled
above.
14. Racemes more than 1.5 cm wide; flowers more than 7 mm in diam.
15. Racemes 5-40 x 1.5-2.5(-3) cm; petals yellowish-green;carpels mostly
strigose. 13. P. pauciflorus.
15. Racemes 5-50 x 3.5-5 cm; petals purplishor dark reddish;carpels gla-
brous. 7. P. racemosus.
14. Racemes less than 1.5 cm wide; flowers to 7 mm in diam.
16. Petals purple;mericarps11-15 x 8-10 mm in outline. 11. P. riedelianus.
16. Petals greenish-yellow;mericarps6-9 x 4.5-6 mm in outline. 12. P. spicatus.

1. Pilocarpus pennatifolius Lemaire, Jard. Fleur. 3: t. 263. 15 Jul 1852; Jard.

Fleur. 3: 52. 1 Jun 1852, nom. invalid. ex Art. 34 (2) ICBN; Muller in
Walpers, Ann. 4: 411. 1857; van Hall, Ann. Hort. Bot. 3: 113, cum
icon. 1860; Carriere, Revue Hort. 40: 60. 1868 (as "P. pinnatifidus");
Engler in Martius, Fl. bras. 12(2): 137. 1874, pro parte (as "P. pin-
natifolius"); Baillon, Adansonia 11: 276-278. 28 Jan 1875; Bentley and
Trimen, Med. pl. 1 (pars 32), t. 48. 25 May 1878, pro parte; Poehl,
Pharm. Z. 19: 142, 1880; Baillon, Traite bot. m6d. phan. 858, fig. 2550.
1884; J. D. Hooker, Bot. Mag. 118, t. 7235. 1892; Holmes, Pharm. J.
Trans. ser. 3. 24: 1066, fig. 2c. 1894; Pharm. J. Trans. ser. 4. 1: 522,
539, cum icon. 1895; Engler in Engler and Prantl, Nat. Pflanzenf. 3(4):
158, fig. 93E-J. 1896 (as "P. pinnatifolius"); Geiger, Ber. Deutsch.
Pharm. Ges. 7: 386, 405, fig. 32a-b. 1897; Duval, Bull. Sci. Pharmacol.
7: 49, t. 1, fig. 1. 1903, Recherches Jaborandis 31. 1905; Hassler, Re-
pert. Spec. Nov. Regni Veg. 10: 345. 1912; Cowan, Sellowia 12: 86,
t. 2, fig. a-c. 1960; Cowan and Smith in Reitz, Fl. Ilustr. Cat. I (RUTA)
25, t. 5-6. 1973.
Usually a shrub or small tree (1-)2-12 m tall with the trunk to 10 cm in diam.
(two records; to 20 cm according to Hassler 416); branchlets 4-7(-9) mm thick,
green- or brown-reddish-grayish, shining when young, perules of terminal buds
densely pubescent with tawny hairs. Leaves alternate, imparipinnate, occasion-
ally the terminal leaflet lacking, 3-jugate, occasionally some 1-, 2-, or 4-jugate,
sometimes 1-foliolate just below inflorescence, with stalked sometimes shifted
leaflets, often (broadly) elliptic or subcircular, 11-34(-39) x 10-25 cm; petiole
(semi-)terete, slightly canaliculate, strongly ribbed and sometimes subwinged, (2-)
3-10(-13) cm long and 0.1-0.3 cm thick; rachis 1.5-14 cm long, the interspaces
1.5-6 cm, strongly ribbed like the petiole, continuing beyond the upper pair of
Pilocarpus 135

leaflets withoutjoint, but occasionallythe uppermostinterspacedoubly articulate;

terminalpetiolule (0-)1-7 mm long, sometimes to 20 mm long in upper leaves;
lateralpetiolules inserted at 45-90?, 2-10 mm long; leaflet blades often (narrowly)
elliptic or oblong, sometimes slightly (ob)ovate, 4-18 x 2-5.4(-6.5) cm, occa-
sionally some smaller, attenuateand unequalat base, obtuse or roundedat apex,
the very tip emarginate, marginrevolute and often slightly undulate-crenulate,
the blade dull green, much paler beneath, venation slightly brochidodromousto
subcamptodromous,principalveins prominulentabove, prominentbeneath, mid-
vein (occasionally slightly) impressed above. Racemes nearly always solitary,
terminal, erect, flexuose, 30-50 x 2-4 cm, with numerousflowers developing in
basipetalfashion soon floweringall together;pedunclemostly absent, or obsolete;
rachis 2-4 mm thick at base; bracts depressedly triangular,ca. 0.2-0.9 mm long
(basal bracts rarely to 1.5 mm) and ca. 1 mm broad, subglabrous;pedicels at
anthesis inserted at 90?, (5-)6-17 (in fruit to 18) mm long and 0.7-1.1 mm thick,
(sub)glabrous;bractlets 2, transversely or occasionally obliquely inserted 13 to
mostly /2 from base of the pedicel, occasionally higher, alternateor rarely sub-
opposite, rarely beset with glandularhairs at the margin. Flowers 8.5-11 mm in
diam.; calyx 5-toothed with apert or valvate to subquincuncialaestivation, 0.5-
1(-1.7) mm long; teeth at anthesis occasionally not overlapping,depressedly tri-
angular,0.2-0.5(-1.4) x 1.1-1.5 mm, obtuse, very thickly coriaceous, beset with
few hairs 0.05 mm especially at margin, otherwise glabrous; petals subvalvate,
adnate to the gynophore, 3.8-5 x 1.9-2.7 mm, inflexed at tip through 0.3-0.5
mm, thickly coriaceous, bordeaux,the upperside carinatetowardstip and slightly
or obsoletely impressed, glabrous, venation parallel to slightly actinodromous,
the nerves branched towards tip, the median nerve thicker; filaments subulate,
slightly flattenedtowards base, 3-4.8 mm long and 0.5-0.8 mm thick, thick, pur-
plish, glabrous;anthers ovoid, 1.5-2.8 x 0.7-1.5 mm, frequently 1.5 x 1.2 mm,
dark ochrous or yellow, with internal gland usually not observable externally;
disc 0.5-1.1 mm high and 1.8-3.5 mm in diam., irregularlyplicate, glabrous;
carpels inserted on a gynophore (0.4-0.5 mm in diam.), 1.1-1.6 mm high, pro-
jecting 0.5-0.8 mm beyond the disc, brown-violet, with discharging internal
glands, glabrous;ovules 2, superposed;style inserted below tips of carpels, cla-
vate or terete, 0.4-0.8 mm long and 0.5-0.8 mm thick, the free part 0.3-0.7 mm,
glabrous; stigma clavate, 0.2-0.4 mm long, green. Mericarps 8-13 x 6-8 mm,
dorso-apicallyblunt usually with an angle of ca. 90?, truncate or slightly convex
at apex, shortly mucronate at the very tip, punctate with glands, glabrous, ex-
ternally wrinkledlongitudinallyat 90?to the nerves, loculicidallydehiscent up to
the dorso-apicalangle; mericarpsreceding from the axis when ripe; axial part of
endocarppersistently connected with the immatureseed; seeds 1 or rarely 2 per
mericarp, kidney-shaped, 7-10 x 4-6 x 3-5 mm, with flattened base and with
curved apex ca. 0.2-0.5 mm, dorsally slightly keeled or doubly keeled, testa
thinly coriaceous, very dark brown, shining, externally finely colliculate, with
flat, irregularroundish collicles ca. 0.03 mm long, hilumjust below apex, 2.5-
3.5 x 1-1.5 mm; embryo ochrous, cotyledons usually ? unequal, ears 0.8-1.2
mm long, radicle conical, 0.8-1.3 x 0.5-0.8 mm and often totally enclosed, plu-
mule obsolete, to 0.2 mm long.

Key to the Varieties of Pilocarpus pennatifolius

1. Leaflets glabrous or beset with spreading hairs 0.05 mm long at base. la. var. pennatifolius.
1. Leaflets rather densely pubescent below with spreading hairs 0.2-0.5 mm long on the whole
surface. lb. var. pilosus.
136 Flora Neotropica

la. PilocarpuspennatifoliusLemaire var. pennatifolius. Fig. 44A-D.

Pilocarpus trijugatusLemaire, Jar. fleur. 3: t. 263. 15 Jul 1852, pro syn. Based on the type of
Pilocarpus pennatifolius Lemaire.
Pilocarpuspinnatus Martiusex Englerin Martius,Fl. bras. 12(2): 138. 1874,pro syn.
Pilocarpusselloanus Englerin Martius,Fl. bras. 12(2):136,t. 30. 1874;Holmes, Pharm.J. Trans.
ser. 3. 5: 641. 1875;Baillon,Bull. Mens. Soc. Linn. Paris 1: 149. 1878;Englerin Englerand
Prantl,Nat. Pflanzenf.3(4): 158, fig. 93A-B. 1896;Cowan, Sellowia 12: 86. 1960,pro syn.
TYPE. Sellow 4021, Brazil. (Rio Grandedo Sul, 1826,accordingto Herter (1949)),fl, (ho-
lotype, B, destroyed, photo 12526made by F, F, NY; isotypes, K ex B, photo NS 2857
made by NY, NY; P, fragmentex B).
Pilocarpussimplex hort. ex Baillon, Adansonia11: 276. 28 Jan 1875,pro syn.; Bull. Mens. Soc.
Linn. Paris 1: 39. after 3 Feb, 1875, pro syn. Based on Hort. P s.n., Dec 1861,fl (P-Type
Herb. Baillon).
Pilocarpusselloanus Englerin Martiusvar. gracilis Chodat& Hassler, Bull. Herb. Boissier. s6r.
2. 4: 1284. 1904;Hassler, Repert. Spec. Nov. Regni Veg. 10: 346. 1912, pro syn. TYPE.
Hassler 4222, Paraguay.(San Pedro), nr. San Estanislao, Aug 1898, fl & fr (lectotype, G
(fl); isolectotypes, G 2x (fr)).
Pilocarpus selloanus Engler in Martius f. brevipedicellata Chodat & Hassler, Bull. Herb. Bois-
sier. ser. 2. 4: 1284. 1904, syn. nov. TYPE. Hassler 3046, Paraguay.(La Cordillera),San
Bernardino,Jun 1898-1899,fl (lectotype, G-Delessert;isolectotypes, G-Chodat& Hassler
2x, MO, MPU, S, W).
PilocarpuspennatifoliusLemairevar. genuinusHassler, Repert. Spec. Nov. RegniVeg. 10: 346.
1912,nom. invalid. ex Art. 24, ICBN.
Pilocarpus pennatifolius Lemaire var. genuinus Hassler f. typicus Hassler, Repert. Spec. Nov.
Regni Veg. 10: 346. 1912,nom. invalid. ex Art. 24, ICBN.
Pilocarpus pennatifolius Lemaire var. genuinus Hassler f. gracilis (Chodat & Hassler) Hassler,
Repert. Spec. Nov. Regni Veg. 10: 346. 1912.
Pilocarpus pennatifolius Lemaire var. genuinus Hassler f. gracilis (Chodat & Hassler) Hassler
subf. deorsum-bracteolatusHassler, Repert. Spec. Nov. RegniVeg. 10:346. 1912(as "deor-
sum bracteolatus"), syn. nov. TYPE. Hassler 9415, Paraguay.Nr. Caaguazu,Sep 1905,fl
& fr (lectotype, G (fl); isolectotypes, G 4x, K (fl & fr)).
Pilocarpus pennatifolius Lemaire var. genuinus Hassler f. gracilis (Chodat & Hassler) Hassler
subf. sursum-bracteolatusHassler,Repert.Spec. Nov. RegniVeg. 10:346. 1912(as "sursum
bracteolatus"), syn. nov. TYPE. Hassler 4222, Paraguay.(San Pedro), nr. San Estanislao,
Aug 1898,fl & fr (lectotype, G (fl); isolectotypes, G 2x (fr)).
Pilocarpus pennatifolius Lemaire var. genuinus Hassler f. latifoliolatus Hassler, Repert. Spec.
Nov. Regni Veg. 10: 346. 1912, syn. nov. TYPE. Hassler 892, Paraguay.Concepci6n:Nr.
Rio Apa, Jul 1885-1895,fl (holotype, G).
PilocarpuspennatifoliusLemairevar. selloanus (Englerin Martius)Hassler, Repert. Spec. Nov.
Regni Veg. 10: 346. 1912;Englerin Englerand Prantl,Nat. Pflanzenf.(ed. 2). 19a: 280, fig.
129A-B. 1931.
Pilocarpus pennatifolius Lemaire var. selloanus (Engler in Martius) Hassler f. brasiliensis
Hassler, Repert. Spec. Nov. RegniVeg. 10: 347. 1912,nom. illegit. ex Art. 26 and 63, ICBN.
Based on Sellow 4021.
Pilocarpus pennatifolius Lemaire var. selloanus (Engler in Martius) Hassler f. intermedius
Hassler,Repert.Spec. Nov. RegniVeg. 10:346. 1912,syn. nov. TYPE.Hassler416, Paraguay.
(La Cordillera),Cordillerade Altos, Jul 1885-1895(G-Chodat& Hassler 2x, K, NY, P).
Pilocarpus pennatifolius Lemaire var. selloanus (Engler in Martius) Hassler f. paraguariensis
Hassler, Repert. Spec. Nov. Regni Veg. 10: 346. 1912, syn. nov. TYPE. Hassler 3046,
Paraguay.(La Cordillera),San Bernardino,Jun 1898-1899,fl (BM, G-Chodat& Hassler 2x,
G-Delessert, K, MO, MPU, NY ex G, S, W).

Branchletsminutelypubescent when young with appressed hairs 0.05-0.1 mm

long, mostly very soon becoming glabrous; perules of terminal buds minutely
pubescent with hairs to 0.2(-0.6) mm long. Petioles puberulous or sometimes
densely pubescent with hairs 0.05-0.2 mm long. Leaflets (thinly) coriaceous,
usually glabrous, but rarely beset with few appressed hairs 0.2 mm long on the
midveinbelow, andfrequentlybeset with spreadinghairs0.05 mm at base. Raches
of racemes glabrous or pilose with hairs 0.05 mm long and sometimes with ap-
pressed hairs 0.1 mm at base.
Type. Originallycollected in 1847as a living plant by Libon near Villa Franca
Pilocarpus 137

(Paraquay,see below). Transportedby de Jonghe from the garden of the Duke

of Croy, Diilmen, BRD, May 1852to hort. BR. Probablyno specimen was pre-
served at that time, but a drawingwas made (lectotype, Lemaire, Jard. Fleur. 3:
t. 263. 15 Jul 1852).
Distribution. Brazil (Goias and Minas Gerais (rare), Paranai,Santa Catarina,
and Rio Grande do Sul), Paraguay, and Argentina (departmentsbordering on
Paraguay). In Brazil in primaryforests and in mata de vairzea;in Paraguayin
woods and thickets in undergrowth;often common or abundant;alt. 100-800 m.
Flowering Apr-Sep, sometimes Dec-Jan(-Mar). Fig. 43B.
Specimens examined. BRAZIL. Goias?: Glaziou 20798a fr (R). Minas Gerais: Brade 17886 fl (RB
2x). Parana: Duarte 1665 bud (U); Dusen 11201 st (S); Falcdo 70 fl (RB); Gomes & Mattos Filho
1049 fr (RB 3x); Hatschbach 9400 fl & fr (US 2x), 11335 fl (F, LP, U), 12946 fr (UC, US), 19303 fl
(K, MO, NY, S, UC), 21514 bud (MO), 26916 fl (S, UC); Hatschbach & Guimardes 19152 fl (C, F);
Hatschbach & de Haas 16583 fl (L 2x); Homrich et al. ICN 5164 bud (U); J. G. Kuhlmann RB 52281
fl & fr (RB 2x); Lindeman & de Haas 650 st (U), 650a bud (U), 710 st (U), 799 st (U), 1482 st (U),
1526 fr (U), 2841 fr (K, NY, U), 5454 fl (K, NY, U); da Rocha Prata 27 fl (U). Santa Catarina: Dusen
11817 bud (F, GH, NY, S); Klein 675 fr (US), 2019 bud (G 2x, L, NY, UC, US, Z 2x), 7445 fl (US);
Klein & Bresolin 7378 fl (US); Plaumann 117 fl (RB); Reitz 1727 dec. fr (US); Reitz & Klein 7401 fr
(F, US), 7578 fl (NY, UC), 8818 fl & fr (BR, F, G 2x, K, L, M, NY, S, UC, US, Z); Smith & Klein
13110 fr (C, R, US); Smith & Reitz 12416 fr (R, US, WIS), 12696 fr (R, US), 12747 fr (LP, MO, NY,
PR, UC, US), 12940 fr (R, US); L. B. Smith et al. 11784 dec. fr (R, US); Ule 1297 fl (HBG, P). Rio
Grande do Sul: Bornmuller 725 fl (A, GH, M); Ceroni et al. ICN 5908 fl (U); Leite 2173 fr (A);
Lindeman et al. ICN 8925 fr (U); Rambo 42102 fl (C, GB, W), 42376 bud (C, US), 42891 fl & fr (BR,
K, MO), 43449 fl (BR, MO, P, W), 47204 fl (BR). State unknown: Anonymus 1 (R 70805) fl (R);
Sehnem 1537 fl (F).
PARAGUAY. Concepci6n: Hassler 8334 fl (BM, G 3x, NY, S). Amambay: Hassler 11282 fl (A,
BM, G 4x, K, NY, P, W, Z); Rojas (in Hassler) 10539a fl (G). San Pedro: Alboff LP 38517 fr (LP);
Woolston 724 fl (C, K, NY, P, S, U, UC). Caaguazu: Burkart 18896 fr (US); Hassler 9337 fl (BM,
K); Krapovickas & Cristobal 13318 bud (MO, UC). Alto Parana: Bertoni 1403 bud (A, L), 1651 fl
(E. G, W); Fiebrig 5383 fl (G 5x, GH, US); Rojas 8162 bud (W). La Cordillera: Fiebrig 19 fl (F, G,
K, L, M); Hassler 11798 fl (A, BM, C, E, F, G 5x, GH, K, L, MO, NY, S, UC, US, Z), 11798a fr
(A, BM, C, E, G 4x, GH, L), 12240 fl (A, BM, C, E, G 4x, GH, K, L, MO, NY, S, UC, US, Z),
12951 fl (A, BM, C, E, G 3x, GH, L); Osten 8815 fl (S); Pedersen 4213 fl & fr (BR, C 2x, G, GH,
NY, US). Central: Archer 4905 dec. fr (US); Bonpland 1226 bud (P 2x); Gibert 55 fl (K); Malme It.
Regn. I 850 fl (R, S 3x), It. Regn. I 850b fl (S 2x); Morin 510 fl (MICH); Morong 635 fl & fr (BM, E,
F, GH, K, MICH, MO, NY 2x, US, WIS), s.n. (Nr. Asunci6n) fl (NY 2x, PHA); Osten 8120 fl (SP);
Pedersen 6039 fl & fl/fr (C 2x, K, L); Schinini 5095 fl/fr (ZT); Zurcher 69 fl (Z). Paraguari: Lindman
It. Regn. I A-3589 (Jan 1894) fl (S 2x), It. Regn. I A-3589 (Apr 1894) fr (S). Guaira: Jorgensen 3977
(5 May 1919) fl (MO), (5 Jun 1929) bud (GH), (4 Jul 1929) fl (C), (17 Jul 1929) fl (DS), (May 1931) fl
(F, US), (Jul 1932) fl & fr (S), (Sep 1932) fl & fr (NY), (Jul) fl (LP); Lourteig 1940 fr (C 2x, P); Montes
15752 bud (MO). Itapua: Anisits 2609 st (S); Lindman It. Regn. I 3589 1/2 (7 Aug 1893) st (S).
Department unknown: Balansa 2514 fl & fr (BM, G 3x, GOET, K 2x, LD 2x, P 2x, S); Egerton s.n.
(1881) bud (K 2x); Kuntze s.n. (Sep 1892) fr (NY 2x).
ARGENTINA. Formosa: Jorgensen 2588 fl (F, GH, MO, US); Morel 3486 fl bud (MO), 3532 bud
(E), 4262 fr (W); Kermes 505 fl & fr (U). Chaco: (?) Meyer 1093 fl (GH, K); Schulz 3948 fl (LP 2x).
Corrientes: Harrola 1264 fr (S). Misiones: Anonymus 62 (LPS 22286) fl (LP); Anonymus 885 (anno
1890) st (HBG); Bertoni 2233 fl (BR, MO), 2869 fl (L, MO), 3657 fl (G), 3698 fl & fr (E, W), 3721 fl
(MO), 3739 fl (W); Cabrera et al. 262 fl (F, LP); Crovetto 5940 fr (BAB, U); Curran 33 fl (A, NY),
682 fl & fr (NY, US); Devoto Herb. Direcc. Forestal 1876 fl (LP); Ekman 1965 fl (LD, S), 1966 fl
& fr (S 2x); Grondona & Piccinini 3228 fr (BAB, U); Jorgensen 163 fl (BAB, U); Krapovickas et al.
15068 bud (C, UC); Lillieskold s.n. (Colonia Bonpland) fl (S); Llamas 769 fl (BAB, U); Maruhak 380
fl & fr (P, R 2x, U, ZT 2x); Meyer 5306 bud (A, F, NY, U), 5478 st (U); Montes 72 bud (L, U, US),
429 fl & fr (BAB, U), 780 fl (BAB, U), 940 fl (A, BM, UC), 2115 fl (A, BM, UC), 2196 fl & fr (F, GH,
K, LP), 2339 fl (S), 2486 fr (LP), 4136 fl & fr (LP), 14773 fr (F, GH, K, M, MO, NY, UC); Muniez
BAB 90811 fl (BAB, U); Rodriguez 293 fl (BAB, U); Scala 138 (?) (Jul 1926) fl (LP 2x, NY, SP, US
2x), 275 (?) (LP 31753, Jul 1927) fl (LP); Schinini 4881 st (WIS); Schwarz 1208 fl (NY), 2593 bud (DS,
GB), 2634 fl & fr (C, GB), 2792 bud (S), 2835 fl (S), 2915 bud (MO), 3020 fl (S), 3083 fl (P, W), 3105
fl (S), 3132 fl (G, W), 3254 fl (S), 4725 fl (G, W), 4766 fl (DS, MO), 7884 fl (G), 7909 fl (BR), 10491
fl (G, W), 10647 fl (S), 10785 fr (NY, S); Schwindt 734 fl (BR), 4887 fl (LD).
CULTIVATED. Badini 3182 (Ouro Preto) fr (R); Bauer (?) s.n. ("Zaopolitan" = Hort. Lwov?,
1861) st (W); Daveau s.n. (Lisboa, 27 Nov 1894) fr (MPU), s.n. (Montpellier, 1908) fl (MPU); Devoto
138 Flora Neotropica

* P.gi nteugrndiforus
v varpilosus
P.pennotifolius

,? 1.)^' t, P. I
<>jaborandi
'C r S / ) . . * P trachylophus
/F~~~~~~~~~~~
~~~~~~~/^---+-<^^l'-^ I
) '-. .
* P.microphyllus

* P.spicatusvar.spicatus '- --------------o

* P.spicatus i.
var.lealii
* P.spicatussubsp.longeracemosus \
-er o sI )
* P.spicatussubsp.arcatensis ) /
v P.pauciflorus 'I * / /

-< - XX
./

IVG.

Ur--*B
**-"--'. -c .... ....

FIG. 43. Distribution of some species of Pilocarpus.

Pilocarpus 139

FIG. 44. Pilocarpus pennatifolius. A-D, var. pennatifolius. E, var. pilosus. A, habit (Osten 8815,
S). B, flower(Osten 8815, S). C, fruit (cultivatedat BR). D, seed (cultivatedat BR). E, indumentof
lower side of leaf (Mosen 3954, type, S).
140 Flora Neotropica

Herb. Direcc. Forestal 1495 (Montevideo) fl (LP); Geier s.n. (Halle, 19 Dec 1960) fl (HAL 4x); Goeze
s.n. (Lisboa, Apr 1876) fl (BM); Greenway 2576 (Amani) fr (K); Hort. BO 149 (Tjibodas) fl & fr (L),
s.n. st (LE); Hort. Cork s.n. (Nov 1893) fl (K); Hort GE s.n. (La Mortola, 1892) st (PHA), s.n. (La
Mortola, 13 May 1893) bud (GE, copy seen), s.n. (Palazzo Orengo, La Mortola, 1894) fr (PHA), s.n.
(La Mortola, Jun 1898) fl (PHA 2x), s.n. (La Mortola 20 Oct 1920) bud (K 2x, US); Hort. K s.n. (Jun
1888) fl (K), s.n. (Nov 1891) fl (K); Hort. LE s.n. fl (LE); Hort. LP LPS 22287 fr (LP 2x); Hort. M
s.n. (Oct 1903) fl (M), s.n. st (M); Hort. Orot (?) s.n. (Jan 1882) fl (Z); Hort. P s.n. (P. simplex) st
(P); Hort. Schonbrunn s.n. (17 Dec 1907) st (W); Hort. STR s.n. (Jun 1893) leaf (PHA); Hort. TO
s.n. (1889) fl (TO); Hort. U s.n. st (U); Hort. W s.n. (1860) fl (W), s.n. (1861) fl (W), s.n. (Nov 1863)
fl (W), s.n. (23 Nov 1868) fl (W); Khek s.n. (Sch6nbrunn, 1896) fl (W); Lavalree 16993 (Meise) fl (BR);
Magnus s.n. (Berlin, 20 Feb 1870) fl (HBG), s.n. (La Mortola, 10 Nov 1892) fl/fr (HBG 2x); Herb.
Neyraut s.n. (Talence, 5 May 1930) st (MPU); Pulle 3053 (Tjibodas) fl & fr (U); Herb. Raphelis 1075
(Alger) fl (MPU); Rn 124 (Hort. Arg. 1892) st (W); Rohrer s.n. (Tanga, 16 Jul 1909) bud (Z 2x);
Rosendahl s.n. (La Mortola, 29 May 1900) fl (S), s.n. (Genova, May 1903) fl (S); Ross s.n. (Miinchen,
Oct 1902) fl (M); Saleman G 6170 (Amani) fl (K); Sampaio 3909 (Rio de Janeiro) fl (R); Veith-Rohrer
s.n. (Sigi, 9 Mar 1910) st (Z); Woloszczak 4047? (cult. v. Hooibrenk, 1884) st (W).

Local names and uses. Goyais:jaborandi(Glaziou20798a); Parana:canela-de-

cutia and guatambu(Lindeman & de Haas 2841); Santa Catarina:cutia-branca
(reports of Klein); Paraguayand Misiones: ivfra-taii,ibira-tai, (several records),
jaborandi (Montes 2196), jaguarandi (Hassler 416); Misiones: yerba-de-cutia
(Montes 780); Chaco: cambai canilla (Meyer 1093).
Infusions against rheums (Schinini 4881 anno 1972);decortations of the bark
against fever as a sudorific, against asthma, and hydropesia (Woolston 724). In
pharmaceuticalliteraturefrequentlymentionedas Paraguayjaborandi,see PHY-
TOCHEMISTRY.
The protologuewas based on livingplants or possibly theirvegetative offspring.
The original plants were gathered, according to Lemaire: "non loin de Villa
Franca (province de St.-Paul, au Bresil) . . . pres d'un endroit nomme Posa
allegre'). . . .') Nous ne saurions garantir l'exactitude orthographique ou geo-
graphique des lieux que nous citons . ..." I have no knowledge of a locality
called Villa Franca with an adjacentPosa allegre in S Brazil. In Paraguayhow-
ever, in Neembucui, opposite to Formosa, at Rio Paraguay, lies a place Villa
Franca, and very close to it PuertoAlegre. Possibly the locality of collection was
wrongly reported as lying in Sao Paulo. Another argumentagainst Sao Paulo is
that from this state only hairy specimens were collected (var. pilosus). Because
the originalplants had glabrousleaves (accordingto the description),it is unlikely
that they were collected in Sao Paulo. Therefore it is plausible that the original
materialwas collected in Paraguay.
Pilocarpus selloanus fully agrees with P. pennatifolius var. pennatifolius. Syn-
onymy was already suggested by Baillon (1878) and Geiger (1897).
Most leaves of Malme 850 are 4-jugatebut one leaf is 5-jugate. Dusen 11817
shows sericeous hairs 0.6 mm long on some petioles. In Montes 940 pedicels ca.
15 mm long are found with 2 alternatebracteoles subtendinga secondarypedicel
ca. 5 mm without bracteoles. The bracteole numbervaries from 2 to 4 in very
rare 6- or 7-merous flowers; some of such flowers were found in the racemes of
Hatschbach 19303and Schwarz2593. The floweringrachis of the lattercollection
is ratherthick: 4-4.5 mm in diam.
This variety has been cultivatedin botanicalgardens. In greenhouses the leaf-
lets grow to 19.8 x 6.2 cm, the leaf-rachisto 17.5 cm, with interspaces up to 7
cm long.

lb. PilocarpuspennatifoliusLemaire var. pilosus Kaastra, Acta Bot. Neerl. 26:

486. 1977. Fig. 44E.
Small tree covered with spreadinghairs 0.2-0.4 mm long on young branchlets,
petiol(ul)es, and raches, and to 1 mm long on the perules of the terminalbuds.
Pilocarpus 141

Leaflets (sub)coriaceous,beset with hairs0.1-0.2 mm long at base above, densely

pubescent below with hairs 0.2-0.5 mm long. Lower part of flowering rachis
pubescent like young branchlets,upper part glabrous, like the pedicels.
Type. Mosen 3954, Brazil. Sao Paulo: Campinas,Jul 1875, fl & fr (holotype,
S (fl); isotypes, C 2x, LD, P (all ex S), S 4x (fl & fr & st, some dated 23 or 25
Jul)).
Distribution. Brazil (Mato Grosso and Goiais(rare), Sao Paulo). Woods and
capoeirao. Flowering Mar-Aug. Fig. 43B.
Specimens examined. BRAZIL. Mato Grosso: Manso 246 fl (BR). Goias: Glaziou 20799a fl & fr
(G 3x, P). Sao Paulo: Dedecca & Teixeira IAC 8165 fl (SP); Duarte 5562 fl (U); Edwall Exp. R. F.
4 fl & fr (SP), IB 19816 fl/fr (NY); Glaziou 3959 fr (P); Kiehl SP 181 fr (SP); Lofgren CGGSP 668 fr
(C, SP); Pickel 5451 fr (US).
Local name and uses. Sao Paulo:jaborandi(Mosen 3954). The tree supplies a
sudorificum according to Mosen 3954. Not used in Goiais (Glaziou 20799a).
This variety has thicker leaves and differs in the indument.It is allopatricand
is the sole variety of Pilocarpuspennatifolius in Sao Paulo.
Engler, in Fl. bras. 12(2)underP. pennatifolius, cited the cultivatedspecimens
which originatedfrom the gatheringby Libon, as well as Manso 246. His de-
scriptionhowever, was based on the latter collection only; this collection belongs
to var. pilosus.
The collection Kiehl SP 181 shows a ratherrobust specimen. The branchletis
12 mm thick, the leaflets are 15-18 x 4.5-5 cm. The rachis of the raceme is very
thick:6 mm at base and still 5 mmin the middle.The hair-coverdoes not differfrom
that in other collections. The mericarpsare roundedat their very apex.

2. PilocarpusdemeraraeSandwith, Kew Bull. (1948): 308. 1948. Fig. 45.

Tree ca. 9 m tall with trunk7.5-10 cm in diam.; branchlets 5-8 mm in diam.,
grayish-brownand somewhat shining when young, puberulous with spreading
hairs 0.05 mm, soon becoming glabrous; perules of terminal buds acuminate,
minutely pubescent towards tip with appressed hairs 0.05-0.1 mm long. Leaves
alternate, imparipinnatebut occasionally without terminalleaflet, (l-)2-4-jugate
(frequently one of a pair shifted) with stalked leaflets, variable in outline, 14-
70 x 18-27 cm; petiole semiterete, slightly canaliculate,ribbed or subwingedto-
wardstip, 5.5-18 mm long and 0.2-0.3 mm thick, indumentlike that of branchlets;
rachis 1.3-25 cm long with interspacesto 9 cm, continuingbeyond the upperpair
of leaflets, upper interspaces doubly articulatewith wings to 0.5 mm broad;pet-
iolules 2-5 mm long, lateral ones inserted at 60-80?; leaflet blades (narrowly)
elliptic to (narrowly)oblong, or ovate, 8-24 x 3.8-7.5 cm, narrowlycuneate to
(very long-) attenuateand unequalat base (lateralleaflets stronglyunequal),acu-
minate or rarely roundedat apex, the acumen emarginateor retuse ca. 0.5-1 cm
long, marginoften slightly undulate-crenulateand revolute, leaflet blades thinly
coriaceous, somewhat shining,paler beneath than above, glabrousor the midvein
pubescent above with spreadinghairs 0.05 mm long, venation brochidodromous,
principalveins and midveinprominulousabove, prominentbeneath. Racemes 1-
2 per vegetative branch, terminalor lateral, erect, 35-85 cm long, with numerous
flowers, minutely pubescent with hairs 0.05 mm long; rachis 1 mm in diam.,
rapidly increasingafter anthesis to 4 mm in diam.; bracts depressedly ovate, to
0.4 x 0.5-1 mm, glabrous;pedicels 1-2 mm long in bud, 3-4 x 0.5 mm postflor-
ally, and 10-15 x 1 mm in fruit; bractlets 2, opposite, transversely or obliquely
inserted just below the flowers. Flowers ca. 8 mm in diam., entirely glabrous;
calyx 5-toothed, 0.8-1.0 mm long; teeth depressedly triangular,0.2-0.5 x 0.7-
1.2 mm, rounded at apex, coriaceous; petals subvalvate, 3.3 x 2.5 mm, apex
142 Flora Neotropica

FIG 45 Pocarpus demerarae (Fanshawe 2811, NY). A, habit, B, seed.

Pilocarpus 143

inflexed through0.5 mm, coriaceous, yellowish-greenwith purplishbase, upper

side carinate towards tip with distinct impressions 3 mm long, venation parallel
to subactinodromous,the nerves branched towards tip; filaments subulate; an-
thers ovoid, 1-1.2 x 0.7 mm, yellow, finely brown-punctate,distallywith a dorsal
gland 0.3 mm; disc 0.9 mm high and 2.5 mm in diam., irregularlyplicate, light-
colored; carpels on a gynophore, 0.7 mm high after anthesis, protruding0.3 mm
beyond the disc, dark brown; ovules 2, superposed or collateral; style obscure
at anthesis, inserted below tips of carpels, 0.3 mm long and 0.2 mm thick after
anthesis; stigma subsessile at anthesis, later on elevated by the developing style,
clavate, 0.6 x 0.6 mm. Mericarpsobovate-ellipticin profile, 15-17 x 13-14 mm,
dorso-apicallyrounded, the very apex obtuse but occasionally subretuse, with
darkglands to 0.8 mm in diam., glabrous, dehiscent down to % below tip; seeds
I per mericarp, ovoid to kidney-shaped, ca. 12 x 9.5 x 7.5 mm, flat at base,
rounded dorsally and apically, slightly keeled dorsally, testa very dark brown,
externallyfinely colliculate with flat collicles ca. 0.04 mm; hilum 5-6 x 2-3 mm;
cotyledons slightly unequal, with ears 2 mm long, enclosing both the radicle 1.2
mm long and the plumule 0.2 mm.
Type. Fanshawe 2065 (Forest Dept. 4801), Guyana. Essequibo: Mazaruni Riv-
er, TakutuCreek to PuruniRiver, 31 Oct 1944,bud & fr (holotype, K, photo NS
2858 made by NY, NY; isotypes, F-1448508n.v., photo 49577 made by F, F; K
3x, NY, U ex K, U ex NY, US ex NY).
Distribution.Known only from Guyana. Ropy mixed forest on lateritic soils.
Flowering Oct-Nov. Fig. 43B.
Specimens examined. GUYANA. Essequibo: Essequibo River, Omai LB, Fanshawe 2811 (FD
5610) fr (K 2x, NY, US).

This species differs from Pilocarpuspennatifolius in the acuminateleaflets, in

the shorter pedicels, in the thinner petals, in the presence of a gland on the
anthers, and in shape and size of fruits and seeds.
Differences from P. grandiflorusare the leaves with fewer, acuminateleaflets,
and the smaller flowers which are differentin color and entirely glabrous.
The small size of the petals as reportedby Sandwithis due to the studying of
flower buds only. My description could be based on the only one petal I found
in a postfloralflower of an isotype in U. The fruit stalks are 2-3 times longerthan
as has given in the originaldescription.

3. Pilocarpus grandiflorus Engler in Martius, Fl. bras. 12(2): 137, t. 31, fig. 1.
1874; Engler in Engler and Prantl, Nat. Pflanzenf. (ed. 2) 19a: 280.
1931. Fig. 46.
Tree, reportedas 10 m tall with trunk 15 cm in diam.; branchlets7-10 mm in
diam., grayish-brown,minutely pubescent with hairs 0.05-0.1 mm. Leaves im-
paripinnate,5-6-jugate, ca. 50 x 25-30 cm; petiole subterete, ca. 12 cm long and
2-3 mm thick; rachis 15 cm long with interspaces 3-5 cm; lateral petiolules in-
serted at 60-70?, ca. 3 mm long (the terminalpetiolule longer), not sharply sep-
aratedfrom the decurrentbase of the leaflet;leaflet blades elliptic or subobovate,
to 16 x 8 cm, the lowermost leaflets smaller, attenuate and unequal at base,
obtuse or rounded at apex, the very tip emarginate, margin strongly revolute,
leaflet blades coriaceous, shining above, dull beneath, green, subglabrous,mi-
nutely appressed-puberulouson the midvein and the base below with hairs 0.05-
0.1 mm, venation brochidodromous,prominent on both sides, the midvein im-
pressed towards base above, very prominentbeneath. Racemes curved, much
144 Flora Neotropica

FIG. 46. Pilocarpus grandiflorus.A, leaf (Engler in Martius, Fl. bras. 12(2): t. 31, fig. 1). B,
inflorescence (Santos 1522, WAG). C, flower bud (Santos 1522, WAG). D, flower bud with petals
removed (Santos 1522, WAG).
Pilocarpus 145

longer than 80 cm, 2-2.5 cm wide, with numerousflowers, puberulouswith fer-

rugineoushairs in all parts; rachis 4 mm thick proximally,becoming transversely
cracked; pedicels at anthesis inserted at 90?, 4-5 mm long and 1.5-2 mm thick;
bractlets 2, alternate, triangular,0.6-0.7 x 0.6-0.9 mm. Flowers ca. 13.5 mm in
diam.; calyx 5-lobed, ca. 2 mm long; lobes separate, depressed-triangular,ca.
0.5-1 x 1.5-2.5 mm, obtuse, thickly coriaceous, minutelypubescent with tawny
hairs to 0.1 mm long; petals valvate, ca. 5 x 3.5-3.7 mm, inflexed at tip through
1 mm, very thickly coriaceous, 0.5 mm thick distally, dark brown-purplish,car-
inate and slightly impressed above, minutely pubescent below with tawny hairs
to 0.1 mm long, venation + parallel;filamentsnarrowlyovate in outline, truncate
at apex, flattened toward base, ca. 3.5 mm long and 0.8-0.9 mm thick, longitu-
dinally ribbed, dark purplish, glabrous or strigillose with hairs 0.1-0.2 mm; an-
thers ovoid, stronglyrecurved, ca. 2-2.5 x 1.3-1.6 mm, with a smalldorsalgland;
disc 1 mm high and 4 mm in diam., irregularlyplicate, brown-purplish,strigillose
with tawny hairs 0.2 mm long; carpels inserted on a gynophoreca. 1 mm, adnate
to the disc, dorsally strongly obtuse-angled,ca. 1.4 mm high, protruding0.7 mm
beyond the disc, dark brown, with internal glands 0.2 mm, densely strigillose
with tawny hairs 0.2 mm; ovules 2, one of them developing;style not protruding
beyond the ovary; stigmasubsessile, capitate, ca. 0.7 x 0.7 mm. Fruitsunknown.
Type. Sellow s.n., Brazil, "inter Victoria et Bahia," see below, fl (B, de-
stroyed; fragment,P-type Herb. Baillon ex B).
Distribution. Brazil (known from Espirito Santo, possibly from Bahia too).
Forests; collected in flower: Mar-Apr. Fig. 43B.
Specimens examined. BRAZIL. Espirito Santo: Linhares, street to Barro novo, Santos 1522 fl
(U, WAG ex CEPEC-Itabuna).
No materialof this species was known after the destroying of the type in B,
except for a good drawing. In P I traced some fragments of the type, a single
leaflet and some flowers. Fortunatelythe species still exists, as demonstratedby
Santos 1522.
Vegetatively Pilocarpus grandiflorus differs from P. pennatifolius in the 5-6-
jugate and rather large leaves. The flowers are quite distinct, e.g. in size and
indument.
There are some vegetative similaritieswith P. demerarae, which see.
There are two possible type localities: 1. between Bahia (=Salvador) and Vic-
toria (=Vit6ria, the capital of Espirito Santo), therefore in the state of Espirito
Santo (1816) or Bahia (1816 or 1817); cf. Herter (1949), or 2. between Bahia
(=Salvador) and Victoria (ca. 100 km N of Bahia), in the state of Bahia (1817 or
1818?).
4. PilocarpusjaborandiHolmes, Pharm.J. Trans. ser. 3. 22: 875. 1892, ser. 3. 5:
581, cum icon. 23 Jan 1875, sine nom., ser. 3. 5: 641. 13 Feb 1875,sine
nom., ser. 3. 23: 1008. 1893, ser. 4. 1: 520. 1895;J. D. Hooker, Bot.
Mag. 122: t. 7483. 1896; Geiger, Ber. Deutsch. Pharm. Ges. 7: 380,
404, fig. 33. 1897;Duval, Bull. Sci. Pharmacol.7: 46, t. 1, fig. 2 and
t. 2, fig. 1. 1903;Recherches Jaborandis:25. 1905. Fig. 47.
Pilocarpus officinalis Poehl, Pharm. Z. 19: 129, 132. 1880, nom. invalid. ex Art. 34(2) ICBN.
Based on: Comm. Poehl s.n., from druggist-shipment from Brazil, st (LE).
Pilocarpus cearensis Rizzini, Leandra6: 34, t. 1, fig. b. 1975, syn. nov. TYPE. Araujo RB
168527, Brazil. Ceara: Vicosa, nr. Brejo Grandeabove Mount Ibiapaba,24 Nov 1974, fr
(holotype, RB; isotypes, RB 4x).
Bush or small tree with a burn smell, quite glabrous except for the buds, or
hairy with + straight somewhat dull tawny hairs; branchlets 3-10 mm thick,
146 Flora Neotropica

FIG. 47. Pilocarpus jaborandi. A, habit (Lynch s.n., CGE). B, leaf with B 1, base of lateral leaflets
(Hort. CGE s.n., 30 Jan 1894, K). C, flower (Lynch s.n., CGE). D, fruit with Dl, mericarp (Araujo
RB 168527, RB). E, seed (Araujo RB 168527, RB).
Pilocarpus 147

grayish, reddish-brown when young, shining, longitudinally wrinkled and

cracked, densely pilose with light tawny hairs to 0.6 mm long becomingglabrous,
perules of terminalbuds densely hirtellous. Leaves alternate, imparipinnate,2-
5-jugate,with the (stalked)leaflets frequentlyalternate,to 35 x 25 cm, gradually
passing through3-foliolate into simple leaves in the proximalpart of the raceme,
graduallymerginginto bracts or all leaves 2-5-jugate and sharply distinct from
the bracts; petiole subterete, canaliculate,winged or not, 3-11 cm long and 1.5-
3 mm thick, the wings 0.1-0.2 mm broad, densely hirtellous with hairs 0.3-0.6
mm long, or glabrous; rachis shaped like the petiole, 4-14 cm long, projecting
beyond the upper pair of leaflets, interspaces 1.5-5.5 cm; petiolules 0.5-7.0 mm
long, the lateral ones inserted at 30-80?, indumentlike that of the petiole; leaflet
blades (narrowly) elliptic or (narrowly) ovate, (4.5-)6-13 x 1.5-5.5(-6.5) cm,
rounded at base (the terminal leaflet too!), occasionally some leaflets shortly
attenuate at base, lateral leaflets strongly unequal at base and terminalleaflet +
equal, obtuse or rounded at apex, the very tip emarginate,marginsubundulate
and revolute, leaflet blades chartaceouswhen young but later on (sub)coriaceous,
dull green somewhat more yellowish below, glabrous on both sides, or sparsely
pilose above with hairs 0.2-0.6 mm long on the midveinand near the marginand
+ densely hirtellous below with hairs 0.4-0.6 mm, venation brochidodromous-
camptodromous,prominenton both sides, midvein plane above and wrinkledor
not. Racemes apical, pendulous, curved, slender, 24-40 cm long and 3 cm wide,
with numerousflowers developing in basipetal fashion from some cm below tip;
peduncle 0-2.0 cm long; rachis ca. 2-2.5 mm thick at base, glabrousor (densely)
hirtellous with hairs to 0.4-0.5 mm long at the basal 10 cm; bracts triangular,to
0.7 mm long, glabrous, the proximalones to 2.5 cm long and 0.7 mm broad and
hirtellouslike the rachis; pedicels inserted at 45-70?, (7-)10-16 mm long and ca.
0.4 mm thick, in fruit a good 1 mm thick, glabrous;bractlets 2, alternate, some-
times subtendingbuds or secondary pedicels (in 1 of the flowering specimens I
studied, all bractletsdoing so, in fruitingspecimens I did not observe it). Flowers
8.5-9 mm in diam., quite glabrous;calyx 5-toothed; teeth separate, depressedly
triangular,0.2-0.5 x 1-1.7 mm, roundedor obtuse, very thickly coriaceous, with
a subapical gland; petals subvalvate, adnate to the gynophore, 3.5-3.7 x 2-2.5
mm, apicallyinflexedthroughca. 0.5 mm, coriaceous,pinkwith yellow marginsbut
the pink hardly observable after drying, carinate towards tip above, without
impressions; venation parallel to actinodromouswith the nerves branched to-
wards tip and the median nerve distinctly thicker; filaments subulate, flattened
towards base, 2.4-2.7 mm long and 0.5-0.7 mm thick, fleshy, rose; anthersovoid
with broadly rounded tip, 1.4-1.5 x 0.8-0.9 mm, pale yellow, with a distinct
dorsal gland 0.3-0.5 mm long; disc 0.5-0.7 mm high and 2.4-3 mm in diam.,
irregularlyplicate, greenish-yellow;carpels on a gynophore0.5 mm, ca. 1.0 mm
high, projecting0.2-0.3 mm beyond the disc, with small glands, rose, glabrous;
ovules 2, collateralgrowing superposed;style clavate, inserted below the tips of
the carpels, 0.6 mm long and 0.3 mm thick, projecting0.3 mm beyond the ovary;
stigma sessile, 0.2 x 0.4-0.5 mm. Mericarps subellipsoidal, ca. 10-15 x 9-13
mm, dorso-apicallyrounded, obtuse at apex, with a dotted appearanceby nu-
merous glands up to 0.5 mm in diam., glabrous, very prominentlynerved, irreg-
ularly wrinkled at base, dehiscent to 1/3 above base, receding from the ventral
axis when ripe; seed 1 per mericarp, kidney-shapedca. 10.5-11.5 x 7.5 x 6.5
mm, flattened at base and rounded at apex, dorsally carinate; testa very dark
black-brown, externally finely reticulate-colliculatewith interspaces 0.05 mm;
hilum ovoid, ca. 4 x 1.3-2 mm; cotyledons subequalwith ears to 1 mm long, the
radicle not projecting.
Type. Comm. E. M. Holmes s.n., Brazil. Pernambuco:Out of commerce, Feb
148 Flora Neotropica

1875, st (lectotype, K, signed in pencil by Holmes: "hairy variety," photo NS

2859 made by NY, NY; photo 462 made by IPA, n.v.; isolectotypes, BM, hairy,
BM, smooth, 12 Jan 1875, K, hairy).
Distribution:Brazil; until recently only known out of commerce from Pernam-
buco, and from botanicalgardensoriginallycollected by J. L. Patersonin Bahia.
Recent collections from Pernambucoand Ceara;alt. 850 m (1 record). Collected
in flower in Nov; fruitingSep and Nov. Fig. 43B.
Specimens examined. BRAZIL. Ceara: Allemdo 274 st (R 2x). Pernambuco: J. G. Kuhlmann RB
46768 st (RB). Out of commerce: Anonymus s.n. (Pharmacie centrale, 1880) st (P), s.n. leaf (P), s.n.
st (ZT); Comm. Craig s.n. (13 May 1875) st (E); Herb. Engelmann 12084 st (MO); Comm. Gubler
s.n. (Apr 1876) st (MPU); Comm. Holmes s.n. (1894) leaves & fruits (BM), s.n. (Nov 1895) fruits (K,
photo 463 made by IPA, n.v.), s.n. (Apr 1896, Feb 1897) fruits & seed (K, on same sheet as the
lectotype), s.n. (Mar 1897) fruits (K), s.n. (Engler vidit) leaf (B, destroyed, photo 12520 made by F,
F, NY); Herb. Holmes s.n. (1898) st (PHA), s.n. ("out of the drug of commerce") st (PHA); Comm.
Kreuzpointner s.n. (1878 from druggist Kleiber) st (M 4x, intermingled with P. pennatifolius).
CULTIVATED. Hort. CGE s.n. (30 Jan 1894) fl (K); Hort. E s.n. (Jun 1888) leaves (K); Hort. K
s.n. (9 Jan 1896) leaves & flowers (K, drawn in Hooker); Lynch s.n. (Cambridge, Oct 1894) fl (PHA),
s.n. (1904) leaf (PHA), s.n. fl (CGE).

Local name and uses. Jaborandi(Allemdo274).

This species was introducedinto Europe as the drug Pernambucojaborandi.
Due to its high percentagepilocarpineit was in great demandby the drugdealers,
but the species is very rare now, see PHYTOCHEMISTRY.
Sterile specimens superficiallyresemble Pilocarpus trachylophus.They differ
in several characters, however, see under P. trachylophus.
Vegetatively this species differs from P. pennatifolius in the bases of the leaf-
lets, and in the more deeply canaliculatepetiole. There have been quarrelsfor a
long time about the hairy variety and the smooth one of Holmes (1875:581). The
latter has been referred to P. pennatifolius ("P. selloanus") but both belong in
fact to P. jaborandi.
Pilocarpus cearensis is identical with this species. It is a pity that Rizzini does
not mention P. jaborandi at all; this could be due to its rarity, however.

5. PilocarpustrachylophusHolmes, Pharm.J. Trans. ser. 3. 24: 1066,fig. 1. 1894;

Pharm. J. Trans. ser. 4. 1: 540. 1895;Geiger, Ber. Deutsch. Pharm.
Ges. 7: 410, figs. 34a-h. 1897;Duval, Bull. Sci. Pharmacol.7: 98, t.
1, fig. 4. 1903;Recherches Jaborandis36. 1905;Engler in Engler and
Prantl, Nat. Pflanzenf.(ed. 2) 19a: 280. 1931. Fig. 48.
Shrub or tree 0.8-8 m tall; indument of cream-tawnyhairs to 0.3(-0.5) mm;
branchlets ca. 3-7 mm in diam., grayish-brown, minutely puberulous when
young; perules of terminalbuds densely pubescent. Leaves alternate,imparipin-
nate, 1-3-jugatewith sessile or shortly stalkedleaflets, subcircular,ca. 3-20 x 6-
18 cm; petiole semiterete, canaliculate, slightly or scarcely winged, 1-5 cm long
and 0. l cm thick, minutelypubescent, the wings to 0.3 mm broad and not eared;
rachis 1.8-8 cm long, the interspaces 1-3 cm; petiolules 0-3 mm; leaflet blades
narrowly oblong or narrowly elliptic, or obovate, rarely some ovate, 3-
11.5 x 1.5-3.7 cm, rounded, obtuse, or narrowlycuneate at base, lateralleaflets
strongly unequal at base, rounded, obtuse, or occasionally truncateat apex, but
the very tip always emarginate,margin recurved, leaflet blades subcoriaceous,
brown-greenabove, distinctly paler and often yellow-green beneath, sparsely or
densely pubescent on both sides with hairs 0.1-0.3 mm above and 0.2-0.5 mm
beneath, venation brochidodromousto camptodromous,prominulous, midvein
plane above and very prominentbeneath. Racemes 1-2 per branch, apical, erect,
Pilocarpus 149

FIG. 48. Pilocarpus trachylophus. A, habit (Macedo 384, SP 52586b). B, base of lateral leaflets
(Macedo 384, SP 52586b). C, flower with Cl, upper side of petal (Macedo 384, SP 52586b). D, fruit
(Anonymus s.n., K).
150 Flora Neotropica

25-40 cm or longer and 0.8-0.9 cm wide with numerous flowers; rachis 2 mm

thick, densely pubescent;bractsandbractlets(depressedly)ovate, to ca. 1 x 0.8-
1 mm, glabrous above, minutelypubescent beneath; pedicels 1-1.5 mm long, in
fruitto 4 mm, densely minutelypubescent;bractlets2, close to the calyx. Flowers
ca. 6.2-7 mm in diam.; calyx 5-lobed;lobes separate, (depressedly)ovate, ca. 1-
2.5 x 1.5-2 mm, acute, coriaceous, minutely pubescent; petals valvate, ca. 2.7-
3 x 1.8-2.5 mm, the tip inflexed through to 0.8 mm, thickly coriaceous, not
transparent, red, purplish when boiled, the keel distinctly winged, the wings
extendingover the ovate, acuminate,distinct impressions,glabrous,or strigillose
beneath, venation parallelto acrodromous;filamentssubulate, flattenedtowards
base, 2.2-2.5 x 0.4 mm, red, glabrous; anthers heart-shaped, 0.7-1.1 x 0.7-1
mm, with dorsal gland 0.2 mm long; disc rosette-like, 5-lobed, shallowlygrooved
at apex, ca. 0.5 mm high and 2.3 mm in diam., purplish, fleshy, wrinkled, gla-
brous; carpels almost completely immersedinto the disc and adnate to it, ca. 0.7
mm high, with small tubercles, densely pilose at tip or glabrous; ovules 2, col-
lateral; style obsolete, inserted midway among the carpels, 0.2 x 0.2 mm, indu-
ment like that of the ovary; stigma capitate-clavate, 0.5 x 0.5 mm, purplish.
Mericarpsobovoid to subellipsoid,8-10 x 7-9 mm, dorso-apicallyrounded, ob-
tuse at apex, not truncate, irregularlywrinkledand muricatewith tubercles to 1
mm high, the parallel-archedribs hardly visible externally and only slightly so
internally, with glands, hoary when young, becoming glabrous except for the
sides; seed I per mericarp,at least 9 x 4.5 mm; cotyledons unequal, eared, ob-
scurely punctate.
Type. Comm. W.J. Bragg & Co. of Liverpools.n., shippedfrom Brazil, Cearfa,
received Feb 1894, st with separateinfructescences (holotype, PHA).
Distribution.Brazil, Cearai,Bahia, and Minas Gerais. Cerradoand marginof
forests, moist stony places, and sandy soils, locally common; alt. 510-620 m, as
far as known; floweringMar-Apr. Fig. 43B.
Specimens examined. BRAZIL. Ceara: Anonymus s.n. fr (K); Comm. Duval s.n. fr (photo 12528
made by F at B, F, NY). Bahia: Espigao Mestre, 6 km S of Cocos, Anderson et al. 37017 fl & fl/fr
(U ex NY); Chique-chique,Serra da Tiririca,Lutzelburg75 (28 Apr 1912)fl & fr (RB 2x); idem,
Joazeirio, Zehntner75 bud & fl/fr (R), 75=975 fl & fr (R). Minas Gerais: 22 km W of Januariaon
road to Serradas Araras,Andersonet al. 9199 fl & fl/fr(U ex NY); 52 km W of Januariaon road to
Serradas Araras,Rio Pandeiros,Andersonet al. 9300 fl & fl/fr(U ex NY); mun. Ituiutaba,chacara
"Vovo Pecho," Macedo 384 fl (BM, MO, SP 2x). State unknown: Herb. Holmes s.n. (leaves in
commerce, 1898) st (PHA).

Local names and uses. Minas Gerais: catiguai(misnamed?Usually a name for

species in Meliaceae)(Macedo 384). Bahia:jaborandi(Liitzelburg75). According
to Duval (1903:98) also known as arruda-do-mato.Dried and fresh as a sudatory
tea (Zehntner75). For its use as a drug (Cearajaborandi), see PHYTOCHEM-
ISTRY.
This species is close to Pilocarpus microphyllus.Yet there are several differ-
ences: the petioles have narrowerwings, the leaflets are narrowerandpale below,
the midvein is plane and not prominentabove. The racemes are pubescent, not
glabrous;the flowers are largerand usually hairy instead of always glabrous.
Sterile specimens in particularare rather similarto P. jaborandi. Differences
are the leaves, which are darkergreen above; the indument(best to be seen at
terminalbuds) is more hyaline-whitishrather than tawny, and more shining, of
stronglycurved hairs. The leaves are shorter, only 1-3-jugate,with usually sessile
leaflets; sessile leaflets were not in P. jaborandi. The racemes, flowers (especially
the petals), and fruits, finally, are quite different.
Except for a record by Engler, this species has only been referredto in phar-
Pilocarpus 151

maceutical literature. J. F. de Toledo, who came first across a noncommercial

collection, was unaware thereby of the existence of P. trachylophus. He gave the
collection a manuscript name but died (in 1952) before he could have published
it. The recent collections of Anderson et al., the specimen seen by Toledo, and
the material of commercial origin, are identical.
According to Geiger (1897), the dried plant has a disgusting scent, but recent
collections do not have it, nor do their labels mention it.

6. Pilocarpus microphyllus Stapf ex Wardleworth, Pharm. J. Trans. ser. 3. 24:

506. 23 Dec 1893. Fig. 49.
PilocarpusmicrophyllusStapf ex Holmes, Pharm.J. Trans. ser. 3. 24: 419. 18 Nov 1893,nom.
invalid. ex Art. 34(3), ICBN.
PilocarpusmicrophyllusStapf, Bull. Misc. Inform.(1894):4 Jan 1894;Oliver,Hooker'sIcon. P1.
24: t. 2331. 1894;Holmes, Pharm. J. Trans. ser. 4. 1: 540, cum icon. 1895;Geiger, Ber.
Deutsch. Pharm.Ges. 7: 394, 412, fig. 31a-c. 1897;Duval, Bull. Sci. Pharmacol.7: 99, t. 1,
fig. 6-7. 1903;RecherchesJaborandis38. 1905;Standley, Contr. U.S. Natl. Herb. 23: 535.
1923,syn. nov. TYPE. Glaziou 13417, Brazil. "Rio de Janeiro:near Rio de Janeiro"(not
correct;piratedby Glaziou, cf. Wurdack, 1970: 1911),Feb 1882(holotype, K (same sheet
as type of correct name), photo NS 2860 made by NY, NY; drawn in Oliver; isotypes,
B-Herb. Eichler, destroyed, photo 12523made by F, F, NY; C, P).

Shrub or treelet 3-7.5 m tall, trunk 3-7.5 cm in diam., bark bitter; branchlets
2-4 mm in diam., grayish-green-brown, shining when young, puberulous with
hairs 0.05-0.1 mm long, glabrous in age, young branchlets often crowded. Leaves
alternate to opposite, imparipinnate, 1-5-jugate but mostly 3-4-jugate with sessile
leaflets, elliptic, 5-15 x 4-6 cm; petiole semiterete, canaliculate by incurved
wings, 0.5-3.5 cm long and 0.2 cm thick, (sub)glabrous, the wings 0.5-1.0 mm
broad with revolute margin and ending in a small, round ear; rachis like the
petiole, 2-6 cm long, interspaces 1-3 cm, the wings broader distally; leaflet blades
ovate to elliptic, the terminal one often obovate, 2-5.5 x 1-3.5 cm, or those of
the upper leaves smaller, shortly attenuate-cuneate and strongly unequal at base,
the terminal leaflet also long-attenuate and usually equal at base, obtuse or round-
ed at apex, the very tip always emarginate, margin revolute towards base; leaflet
blades chartaceous, green and shining on both sides, glabrous, venation brochi-
dodromous-camptodromous, somewhat prominent on both sides, midvein prom-
inent above, plane to prominent below. Racemes solitary, terminal to axillary, to
30 cm long, ca. 0.5 cm wide, with numerous flowers, (sub)glabrous; peduncle
absent or obsolete; rachis 0.5-1(-2) mm thick; bracts broadly triangular to ovate,
0.2-0.3 mm long; pedicels 0.1-1.5 mm long and 0.3 mm thick, in fruit 1.5-2.5 mm
long and 1 mm thick; bractlets 2, alternate, obliquely inserted in upper part of
pedicel. Flowers 4-5 mm in diam., quite glabrous; calyx 5-toothed; teeth separate,
very broadly to depressedly triangular, or subcircular, ca. 0.2-0.4 x 0.4-0.6 mm,
obtuse or rounded at apex, coriaceous; petals subvalvate, 1.8-2.3 x 1.1-1.2 mm,
apex inflexed through 0.3 mm, thinly coriaceous, semitransparent, brown when
dried, the upper side carinate distally and impressed, venation imperfectly ac-
rodromous; filaments subulate, flattened towards base, ca. 1-2 mm long and 0.2-
0.4 mm thick; anthers ca. 0.5-0.7 x 0.4-0.6 mm, with an oblong dorsal gland;
disc ca. 0.5 mm high and 1.2-1.5 mm in diam., slightly 10-plicate, light-colored,
foveolate; carpels ca. 0.5 mm high, projecting 0.2 mm beyond the disc; style
inserted below tips of carpels, not projecting beyond the ovary, 0.2-0.3 mm long
and 0.1 mm thick; stigma capitate, 0.2 x 0.4 mm. Mericarps subovoid, 7-10 x 5-
8 mm, dorso-apically rounded, obtuse or rounded, minutely notched at the very
apex, punctate with glands 0.2-0.3 mm, glabrous, wrinkled towards base, dehis-
152 Flora Neotropica

FIG. 49. Pilocarpus microphyllus. A, habit (Sucre et al. 9448, RB). B, fruit (Lindeman et al.,
180, U) C, seed (Lindeman et a. 180 U).
Pilocarpus 153

cent up to 3/ below tip; seed 1 per mericarp,ca. 5.2-7 x 4.2-5 x 3.8-5.1 mm,
flattened at base, rounded at apex, slightly carinate dorsally; testa very dark
black-brown,externallyflatlyreticulatewith interspaces0.05-0.1 mm long; hilum
ca. 1.4-2.5 x 0.5-1 mm; cotyledons subequalwith ears ca. 0.4-1 mmlong, radicle
conical, 0.5 mm long, plumule0.05 mm long.
Type. Comm. T. H. Wardleworth (Evans, Sons, and Co., from Liverpool) s.n.
(31 Oct 1893) Brazil. Maranhao,st (lectotype, K, photo NS 2860 made by NY,
NY; paratype, PHA, lost).
Distribution.Surinam(1 collection); more frequently collected in Brazil (Ma-
ranhao and Piaui). Ferro-bauxiticsoil in Surinam,primaryforest on terra firme
on rocky, reddish soil in Brazil; alt. 110-710 m. FloweringFeb-Mar(-Jun);fruit-
ing Sep. Fig. 43B.
Specimensexamined. SURINAM. Marowijne:Lely Gebergte,Lindemanet al. 180 fr (U 4x).
BRAZIL. Maranhao: Ilha de Sao Luis, Froes (in Krukoff) 9568 st (U), (in Krukoff) 9573 (U), (in
Krukoff)11668fr (A, F, MO, NY, S), (in Krukoff)11762bud & fl/fr(A, GH, K, LP, MICH,MO, NY,
S, U); locality unknown: Comm. Bragg & Co. s.n. (1895-6) st (PHA); Comm. Holmes s.n. (8 Nov
1893)fr (K, on same sheet as type, photo, NY), s.n. (Mar 1897)fr (K, on same sheet as type, photo,
NY); Comm. Wardleworth s.n. (8 Nov 1893) leaves & fr (K). Piaui: Parnaiba, Clissold s.n. (rec. 10
Nov 1916) seeds (K); Lfizilandia, Mucambo, Sucre (& J. F. da Silva) 9446 fl & fr (U ex RB), 9448
fl & fr (RB 3x). NorthernBrazil:Jobert s.n. (1877-1878)fr (P). Locality unknown:Herb. Holmes
s.n. (out of commerce, 1905) st (PHA); Comm. Wright et al. s.n. (out of commerce, Jan 1905) fr
(PHA).
Local names and uses. Maranhao:as "jaborandi?"importedto Liverpool, 1893
(the type); jaborandi (Froes (in Krukoff) 11668); arruda-brava (Froes (in Krukoff)
11762; Clissold s.n.); arruda-do-mato(Duval, 1903:99). Piaui:jaborandiand ar-
ruda (Sucre 9448).
Known as the drug Maranhamjaborandi. It has the highest contents of pilo-
carpine and the species is still in great demand by drug trading-companies.In
spite of the relative few herbariumspecimens known, the dealers are able to
collect huge quantities, see PHYTOCHEMISTRY.
Vegetatively this species is distinguishedby its smallleaflets and by the winged
petioles and raches. In commerce it has been confused with Swartzia decipiens
Holmes ex Geiger (Leguminosae),now referableto Bocoa (Cowan (Fl. Neotrop-
ica 1: 212. 1967);see Holmes (1896)and Geiger (1897:418)).
The first valid publicationof the name was by Wardleworthin Dec 1893. Al-
though Wardleworthknew that Stapf would publish the name in the next issue
of Kew Bull., he nevertheless published it a few weeks earlier. The description
by Wardleworthis slightly differentfrom that by Stapf. Wardleworthdid not see
the type of Stapf s name, but as he had sent sterile parts to Stapf and to Holmes
(Wardleworth,1893a),those specimens must be regardedas the types.
7. PilocarpusracemosusVahl, Eclog. 1: 29, t. 10. 1797(" 1796");Jaume Saint-
Hilaire, Expos. fam. nat. 2: 356. Feb-Apr 1805;Persoon, Synopsis pl.
1: 243. Apr-Dec 1805; Nees and Martius, Nova Acta Phys.-Med.
Acad. Caes. Leop.-Carol. Nat. Cur. 11: 177, t. 19, fig. I. after 26 Jul,
1823;de Candolle, Prodr. 1: 728. 1824;Sprengel, in Syst. veg. ed. 16.
1: 954. 1824/1825;Jussieu, Mem. Mus. Hist. Nat. 12: t. 22, fig. 29.
after 27 Jun, 1825;Saint-Hilaire,Hist. rem. Bres. Par. 1: 145, t. 16. 4
Nov 1825;Dietrich, Syn. pl. 2: 1017. 1840;Grisebach, Fl. Brit. W. I.
135. 1859; Urban, Bot. Jahrb. Syst. 21: 553. 1896; Duval, Bull. Sci.
Pharmacol.7: 103, fig. 3. 1903;Recherches Jaborandis46. 1905;Wil-
son, in North Amer. Fl. 25: 200. 1911;Standley, Field Mus. Nat. Hist.,
Bot. Ser. 3: 309. 1930.
154 Flora Neotropica

- ' I '\ ' ''= ' ' i' I

* Procemosus
varracemosus _ . -- - '
ov Procemosus
voryucatanus - '" . . , !
Prcemosussubspviridulus ,II . . y ' ,-
* Pgoudotionusvorgoudotionus , /-> --. \.' - J
o Pgoudotianusvor mculis '
* P goudotionussubsp heterochromus - - --- -- - , - - -.-'- _ _ , ',
"

FIG. 50. Distribution of Pilocarpus racemosus and P. goudotianus.

Shrub or small tree 2-8 m tall, (8-20 m tall according to reports from Vene-
zuela), trunk narrow in diam., bark smooth and dark grayish; branchlets 4-8 mm
in diam., dull grayish-brown, shining when young; perules of terminal buds am-
pulliform, rather abruptly terminating in a long, blunt-pointed tip to I mm long.
Leaves alternate, often crowded and subopposite at the ends of the branches,
imparipinnate or rarely paripinnate l-2(-3)-jugate, or 1-3-foliolate, with usually
stalked leaflets, occasionally simple, 8-23(-32) cm long; petiole (semi-)terete, often
slightly winged, varying from 0.5-6(-11) cm, 1-2(-3) mm in diam.; the wings to
0.4 mm broad; rachis 3-5(-7.5) cm long, in apical leaves 1-2.5 cm, wings usually
broader than those of the petiole; petiolules 0-6(-1 1) mm, those of terminal leaf-
lets to 35 mm; leaflet blades (narrowly) elliptic to slightly obovate, rarely some
ovate, 4-16(-30) x 1.5-9.5(-11.5) cm, shortly attenuate at base, (terminal leaflets
also to long-attenuate), base of lateral leaflets + unequal, obtuse or rounded at
apex, sometimes slightly acuminate, the very tip emarginate or less frequently
retuse, margin + revolute; leaflet blades (sub)coriaceous or chartaceous, dull
green, paler beneath, venation brochidodromous to camptodromous, prominent
or plane on both sides, midvein plane above. Racemes usually solitary, subter-
minal, erect, 5-50 x 3.5-4(-5) cm, usually 10-25 cm long, rather loosely flowered
with numerous flowers, mostly glabrous, or minutely pubescent with spreading
hairs 0.05-0. 1(-0.2) mm long; rachis 1.2-3 mm thick at base; bracts broadly or
depressedly triangular, 0.5-0.8 x ca. 0.5 mm; pedicels alternate, rarely some
subopposite, inserted at 45-90?, 8-21 mm long and 0.4-0.7 mm thick, in fruit up
to 20(-26) mm long and I mm thick; bractlets 2, alternate, inserted in the upper
part of the pedicel, close to the calyx or not, 0.3-0.6 x 0.6-1 mm, tufted or not
with hairs to 0.2 mm long. Flowers 7.5-9(-9.2) mm in diam.; calyx 5-lobed, ca.
0.6-0.7 mm high; lobes separate-valvate, depressedly triangular or semicircular,
0.2-0.6 x 1-1.7 mm, rounded at apex, coriaceous, glabrous or rarely sub-
glabrous; petals subvalvate, 3.2-4(-4.5?) x 1.8-2.8 mm, apex inflexed through
Pilocarpus 155

0.6-0.7 mm, thickly coriaceous, carinate and clearly impressed above, the keel
ca. 1-2 mm long and 0.3-0.4 mm high, dull above, + shining beneath, glabrous,
venation parallelto actinodromous;filamentssomewhatadnateto the disc, some-
what flattened,trigonousat base, slightly indentedlaterally, subulatetowardstip,
1.7-2.2(-2.8) x 0.3-0.4(-0.6) mm, glabrous;anthersdorsifixedabove the middle,
heart-shaped, 1-1.5 x 0.9-1.2 mm, pale yellow, with a dorso-apical, oblong
gland; disc equalling the ovary or lower, 0.3 mm thick and 2-3.5 mm in diam.,
plicate, surroundingthe base of the filaments, tuberculatein fruit, glabrous;car-
pels competely adnateto the disc or only proximally,ca. 0.7-0.9 mm high, usually
not projectingbeyond the disc at anthesis, glabrous;ovules 2, superposed;style
insertedjust below the tips of the carpels, clavate, 0.2-0.3 mm long and 0.2-0.5
mm thick, sometimes not reaching beyond the ovary, up to 1.5 mm long after
anthesis, glabrous;stigma clavate, 0.2-0.3 x 0.4-0.5 mm. Mericarpssubobovate
in outline with ratherstraightventralaxis, 9-12 x 8-9 mm, dorso-apicallyround-
ed, obtuse or rounded at apex, slightly mucronateat the very tip, punctate with
glands to 0.4 mm in diam., glabrous, dehiscent up to /2 or 2/3below tip; seed 1
per mericarp,dischargedimmediatelywhen ripe, kidney-shaped,6-9 x 5-7 x 4-
5 mm, flattenedat base and obtuse at apex, slightlycarinatedorsally;testa black-
brown, finely reticulate-colliculatewith interspaces 0.04-0.05 mm, slightly irreg-
ularly wrinkled;hilum tear-shaped,2-3.5 x 0.7-1.4 mm; cotyledons ? unequal
with ears 0.6-1.5 mm long, radicle 0.5-0.8 mm long, and plumule 0.1 mm long.
Distribution.Caribbean.Fig. 50.

Key to the Subspecies of Pilocarpus racemosus

1. Petals purplishor darkreddish;leaves 1-2-jugate, 1-3-foliolate,or simple.
7a. subsp. racemosus.
1. Petals greenish;leaves frequently2-3-jugate. 7b. subsp. viridulus.

7a. PilocarpusracemosusVahl subsp. racemosus.

Extra-Venezuelanspecimens 2-6 m tall. Leaves 1-2-jugate, 1-3-foliolate, or
sometimes simple. Leaflets elliptic to slightly obovate, to narrowly elliptic in
Cuban and Venezuelan specimens. Petals purplish or deep red, disc becoming
so, stigma greenish, calyx, filaments, and ovary ? purplish.

Key to the Varieties of Pilocarpus racemosus subsp. racemosus

1. Perulesof terminalbuds with tuftedhairs, otherwiseglabrousor ciliate; petiole and leaflets
usually glabrous. 7a-l. var. racemosus.
1. Perules of terminalbuds densely pubescent;petiole, and base and midvein of leaflets, pu-
bescent with hairs 0.1-0.2 mm long. 7a-2. var. yucatanus.

7a-1. PilocarpusracemosusVahl subsp. and var. racemosus. Fig. 51A-H.

Pilocarpus latifolius Saint-Hilaire, Fl. Bras. mer. 1: 82. 1825, nom. nud.
Pilocarpus latifolius Saint-Hilaire ex Tulasne, Ann. Sci. Nat. Bot. ser. 3. 7: 285. 1847 (as "P.
latifolia"); Englerin Martius,Fl. bras. 12(2): 135. 1874;Duval, RecherchesJaborandis51.
1905;Englerin Englerand Prantl,Nat. Pflanzenf.(ed. 2) 19a: 280. 1931,syn. nov. TYPE.
Martins.n., French Guiana,Cayenne, fl (holotype, P, drawingin P-Herb. Sagot; isotypes,
B-Herb. Kunth ex P, destroyed, photo 12521made by F, F, NY; G-Moricand(without
collectorand no.), K ex P, photo NS 2854 madeby NY, NY; P ex Herb. Baillon,P ex Herb.
Jussieu, P 3x, L ex P (s.n., holotype of P. guyanensis van Hall)).
PilocarpuspauciflorusKnox, Cat. 86. 1857,non Saint-Hilaire,n.v., pro syn., teste Urban, Bot.
Jahrb. Syst. 21: 553. 1896.
Pilocarpus guyanensis van Hall, Ann. Hort. Bot. 3: 116. 1860, syn. nov. TYPE. (Martin s.n.),
156 Flora Neotropica

FIG. 51. Pilocarpus racemosus. A-H, var. racemosus. I, var. yucatanus. K, subsp. viridulus. A,
habit (Stehle 394, NY). B, shoot apex (Stehle 394, NY). C, inflorescence (Duss 2240, NY). D, flower
Pilocarpus 157

French Guiana, (Cayenne),fl (holotype, L ex P, =isotype of P. latifolius Saint-Hilaireex

Tulasne).
RaputiaheterophyllaGrisebach,Mem. Acad. Arts. ser. 2. 8: 170. 1861,non de Candolle;Urban,
Bot. Jahrb.Syst. 21: 553. 1896,pro syn. TYPE. C. Wright1129, Cuba. Oriente:Nr. Santa
Catalinade Guantanamo,nr. Monte Verde, Jan-Jul 1859,fr (holotype, GOET-Herb.Gri-
sebach; isotypes, CGE, G 2x, GH, K, LE, MO, NY (photo), P, W).
PilocarpusheterophyllusA. Grayin Grisebach,Mem. Acad. Arts. ser. 2. 8: 170. 1861;Grisebach,
Cat. pl. cub. 48. 1866,syn. excl.; Holmes, Pharm.J. Trans. ser. 3. 5: 581. 1875;Urban,Bot.
Jahrb.Syst. 21: 553. 1896,pro syn.; Duval, RecherchesJaborandis57. 1905.TYPE. C. Wright
1129, Cuba. Oriente:Nr. Santa Catalinade Guantanamo,nr. Monte Verde, 25 Dec 1859,fl
(holotype, GH).
Pilocarpus laurifolius Vahl ex Eggers, Bull. U.S. Natl. Mus. 13: 37. 1879, pro syn. Based on
Ryan s.n., West Indies. LeewardIslands: Montserrat,fl (C-Herb.Homemann).
Pilocarpuslongipes Rose, Contr.U.S. Natl. Herb. 5: 112. 1897;Wilson, in North Amer. Fl. 25:
199. 1911;Standley,Contr.U.S. Natl. Herb. 23: 535. 1923;Field Mus. Nat. Hist., Bot. Ser.
3: 309. 1930,pro syn.; Englerin Engler and Prantl,Nat. Pflanzenf.(ed. 2) 19a: 280. 1931.
TYPE. Palmer 514, Mexico. Guerrero:Nr. Acapulco, Feb 1895, fl & fr (lectotype, US-
256912;isolectotypes, F 2x, GH, MO, NY, UC, US ex Donnell-Smith2x (all ex US), US-
256916).
Pilocarpus insularisRose, North Amer. Fauna 14: 80. 1899;Wilson, in North Amer. Fl. 25: 199.
1911,pro syn. TYPE. Nelson 4307, Mexico. Nayarit:Las Tres Marias,Isla MariaMadre,
3-25 May 1897,fr (holotype, US-346051,fragmentNY; isotype, K).
Branchlets glabrous, perules of terminal buds with tufted brown-translucent
hairs at tip, otherwise glabrous or ciliate. Petiole glabrous or very rarely beset
with spreadinghairs 0.05 mm long. Leaflets glabrous or sometimes beset with
hairs 0.05-0.1 mm at base.
Type. Ryan s.n., West Indies. Leeward Islands: Montserrat, fl (holotype,
C-Herb. Vahl, IDC microfiche 541, 3-4; isotypes, BM, C-Herb. Homemann,
C-Herb. Schumann3x, 2 of which collected by Ryan & Lamb., P-Herb.Jussieu).
Distribution. West Coast of Mexico, West Indies from Cuba to Martinique,
Venezuela (Bolivar and rare in Lara), and French Guiana. Woods, usually on
calcareous hills, once savannas (Ekman 12473);alt. to 1200 m. FloweringAug-
Mar.
Specimens examined. MEXICO.Nayarit:Maltby82 fr (US 3x, photo, NY); Mason 1837 st (GH,
K, US). Michoacan:Hinton 16178fl (G, K, MICH, NY, UC, US).
CUBA. Pinardel Rio: Ekman 18036fr (G, LD, S), 18228 fr (S). Habana:Le6n 11412bud (GH,
NY), 14781 fr (GH); Wilson 9177 bud (GH, K, NY 5x), 9549 fl/fr (NY 3x, US); Wilson & Le6n 1949
bud (NY). Las Villas: Britton & Britton 5166 fr (NY); Britton & Wilson 5460 fr (NY 2x); Ekman
14017 fr (BM, S); Luna 124 bud (NY), 463 fr (NY), 777 bud (NY); Morton 10531 st (US), 10781 st
(US), 10821 fr (US). Oriente: Ekman 15222 fl & fr (S), It. Regn. III 3751 fl (S), 3757 bud (NY), 4208
bud (S), 4753 st (F, S); Le6n 12253 st (NY); Le6n & Alain 19176 dec. fr (GH), 19336 bud (GH); Leon
& Matos 19645 fl (GH, NY, US); Morton & Acuna 3434 fl (US 2x); Shafer 3610 fr (F, NY 2x, US),
3616 fr (F, NY 2x, US), 4219 bud (F, NY 2x, US), 8323 bud (K, NY 2x, US), 8366 bud (NY, US),
8779 fr (NY); Wright 1129 (1860) fl (GOET), (13 Oct 1860-1864, Monte .. .) fl (W), (1860-1864) fl
& fr (BM, K, L, S), (12 Dec 1865) bud (NY), s.n. (ex Herb. Olneyanum, only lowermost specimen)
fr (F), s.n. (Cuba orientalis) fl (S). Isla de Pinos: Britton et al. 15264 st (NY, US); Ekman 12473 bud
(A, S). Departmentunknown:de la Sagra 820 bud & fr (P); s.n. (Herb. Triana,rec. 1 Dec 1877)fr
(P).
HAITI: Ekman H 3051 fr (GH, S, US), H 3564 st (G, S), H 7292 st (C, K, S), H 9488 fl (A, S).
DOMINICAN REPUBLIC: Ekman H 15062 fl (S).
PUERTO RICO. Aquadilla s.n. (comm. Nov 1889) st (K); Sintenis 5649 st (BM, F, G, GH, K, L,
LD, LE, M, MO, NY, P, US, W), 5751 fr (E, G, GH, GOET, K, LD, P 2x, S 2x, Z). Vieques, Ravn
s.n. ("in Crabbeneiland") fr (C 2x); Shafer 2387 fr (NY 2x, US), 2557 fr (NY, US).

(Duss 2240, NY). E, fruit (Stehle 394, NY). F, endocarp (Stehle 394, NY). G, axial part of endocarp
with two obturatorsand one vestigial ovule (Stehle 394, NY). H, seed (Stehle 394, NY). I, shoot
apex (Gaumer 24399, type, F). K, habit (Jimenez M. 1529, type, NY).
158 Flora Neotropica

Table II
Distinguishing Characters Between Pilocarpus racemosus and P. goudotianus

P. racemosus P. goudotianus
Leaves l-5(-7)-foliolate or simple 3(-l)-foliolate or 1-jugate
Indument of leaves (nearly) absent always pubescent
Diam. of inflorescence (cm) 3.5-4(-5) 3-3.5 and up to 6
Indument of inflorescence absent or rarely pubescent (densely) pubescent with
with hairs of 0.05-0.1(-0.2) hairs of 0.1-0.4 mm
mm
Length of pedicels (mm) 8-21 10-15 and 20-24
Diam. of flowers (mm) 7.5-ca. 9 ca. 9-11
Breadth of petals (mm) 1.8-2.8 2.2-3
Length of filaments (mm) 1.7-2.2(-2.8) 2.5-3.5

VIRGIN ISLANDS: St. John. Eggers s.n. (10-12 Mar 1877) fr (C 2x, GH, M 3x).
LEEWARD ISLANDS: Montserrat. Robson s.n. (comm. 4 Jun 1908) fl & fr (K 2x); Shafer 445
bud (F, NY, US). Guadeloupe. Guadeloupe: Bena 981 fl (P), 1699 bud (P); Comm. Wright et al. s.n.
(Sep 1903) st (PHA). Desirade: Stehle 394 fr (NY). Dominica. Anderson s.n. (ex Herb. Forsyth) fr
(K).
WINDWARD ISLANDS: Martinique. Duss 1193 fl & fr (MO, NY), 1193.4558 fl (F, NY 2x, US),
2240 fl & fr (F, MO, NY 2x, US), 4558 fl (Z); Sieber s.n. (Herb. Martius suppl. Ill) bud (M). Locality
in West Indies unknown: Anderson s.n. ("insula caribua") fr (BM); I'Herminier s.n. fl & fr (G 4x,
P); L. Cl. Richard s.n. (Bouillantes) fl (P 4x), s.n. (Antill.) fl (P).
VENEZUELA. Lara: R. F. Smith V 659 fr (VEN). Bolivar: Steyermark 87998 bud (F, GH, MICH,
NY, US, VEN), 88161 fl (NY, U, VEN), 88515 bud (K, NY, U, VEN). State unknown. Herb. Nyst
(Anonymus 12) fl (BR); s.n. (17 Jan) fl (BR); s.n. fr (BR); de Ponthieu s.n. (misit Banks 1772) fl/fr
(BM, S); Comm. Roemer & Schultes s.n. ft (G).
CULTIVATED. Hort. Kew H 493(66) fl (K); Saleman G 6169 (Amani) fl bud (K 2x); Winkler &
Hanke 627 (Cameroun)fl (Z).
Local names and uses. Martinique:bois flambeaucaraibe (Duss 4558); bois
blanc (Duss 2240). For uses as drug, see PHYTOCHEMISTRY.
Pilocarpus racemosus is very near to P. goudotianus. I have toyed for a time
with the idea to unite the two but decided to distinguish two species because
there are a lot of easily recognizablemorphologicalcharacters, see Table II. The
species show parallel variation, see Table III. The separatingof the species has
a practical side since one single large species would have had eight infraspecific
taxa.
The types of Pilocarpus insularis and of P. longipes are not different from P.

Table III
Parallel Variation in Pilocarpus racemosus and P. goudotianus

P. racemosus P. goudotianus
Greenish flowers subsp. viridulus subsp. heterochromus
Long pedicels former P. latifolius var. mollis
Perules of terminal buds var. yucatanus subsp. goudotianus
densely hairy
Pilocarpus 159

racemosus var. racemosus. These names therefore, are taxonomic synonyms, as

Standley (1930: 309) already suggested.
Pilocarpus latifolius is known only from the type. The leaves at the tips of the
branchlets are simple and slightly acuminatetowards the tip, (7-)13-30 cm long
and to 9.5 cm broad. The pedicels are up to 21 mm long and become to 26 mm
postflorally.The flowers are up to 9.2 mm in diam. with petals of ca. 3.8 mm long
and 2.2-2.7 mm broad; the filamentsare 2.7-2.8 mm long.
The slightly largerflowers with longer filamentsdiffer somewhat from those of
P. racemosus, the petals however, show no differences. Because there is a wide
range as to the sizes in the racemes of P. racemosus, these minordifferencesare
not significant. Various collections of P. racemosus, especially from the West
Indies, have simple leaves, especially towards the ends of the branches. Speci-
mens of P. racemosus sometimes also show large leaves to 22 x 11.5 cm. Ped-
icels attaining a length of 26 mm after anthesis also occur in the Venezuelan
collections mentioned below (R. F. Smith V659; Steyermark 88161). The minor
differences of P. latifolius with P. racemosus could merely be due to ecological
variation:P. latifolius was collected near the marginof the area of P. racemosus.
Pilocarpus guyanensis van Hall was describedfrom a poorly labeled duplicate
of the holotype of P. latifolius and is therefore a homotypic synonym.
Three collections from Venezuela are referred here to the present variety,
althoughP. racemosus was not known from there. The specimens were collected
from remarkablytaller trees with narrowlyelliptic leaflets frequentlyslightlyacu-
minate at apex; the leaves are 3-foliolate or 1-2-jugate.Leaves of this kind were
also found in specimens from other areas; the taller growth may be attributedto
differentecological conditions.
Specimens with some 2-jugateleaves were only found in Mexico and Puerto
Rico (including Vieques). Hinton 16178, from Mexico, has 1-3-jugate leaves.
Sintenis 5649, 5751, and Ekman 4208 show vigorous shoots with long internodes
and large leaves.
Rarely some pedicels bear 3 or 4 bractlets: Leon 19645, Wright1129 (Jan-Jul
1859,GOET-Herb.Grisebach).
7a-2. PilocarpusracemosusVahl subsp. racemosusvar. yucatanusKaastra, Acta
Bot. Neerl. 26: 486. 1977. Fig. 511.
Small tree, branchletswhen young pubescent with spreadinghairs0.1-0.3 mm,
becoming glabrousin age; perules of terminalbuds densely pubescent with hairs
0.2-0.3 mm. Leaves (l-)3-foliolate or imparipinnate1-2-jugate;petiole pubescent
with hairs 0.1-0.2 mm long; leaflets (narrowly)elliptic to subovate, occasionally
subacuminateat apex, pubescent at base and on midveinon both sides with hairs
0.1-0.2 mm long.
Type. Gaumer 24399, Mexico. Yucatain,st, 1917-1921 (holotype, F-552392;
isotypes, G, GH, MO-951573,NY, US-1268394, all ex F).
Distribution.Confinedto Mexico, Yucatan;advanced deciduous forest (I rec-
ord).
Specimens examined. MEXICO. Yucatan: Progreso, Flores s.n. (Feb 1935) fr (F); nr. Yokdzonot,
Lundell & Lundell 7861 dec. fr. (MICH); exact locality unknown: Gaumer 24399 st (F, G, GH, MO,
NY, US).

Variety yucatanus is distinguishedby the persistent indument, whereas var.

racemosus is nearly always glabrous, an indument, if any, of hairs to 0.05 mm
only. Intermediatespecimens are unknown. Flowering specimens are wanted.
160 Flora Neotropica

7b. PilocarpusracemosusVahl subsp. viridulus Kaastra, Acta Bot. Neerl. 26:

486. 1977. Fig. 51K.
Small tree 6-8(-15) m tall with trunk to 30 cm in diam. and with dense, round
crown. Leaves frequently(l-)2-3-jugate, but 1-3-foliolateleaves also exist; leaf-
lets occasionally narrowlyelliptic. Flowers greenish.
Type. JimenezM. 1529, Costa Rica. Guanacaste:Liberia,bankof Rio Colorado
between Hacienda Guachipelin and Las Pailas, 4 Jan 1964, fl (holotype, CR;
isotypes, F-1698498,NY).
Distribution. El Salvador and Costa Rica; usually dry forest; alt. 500-800 m.
Flowering in Jan.
Specimensexamined. BELIZE. Countyunknown, Turner7 st (probablythis subsp.) (U).
EL SALVADOR.Ahuachapan:Sierrade Apaneca, regionof Finca Cobina,Standley20093fl (GH,
NY, US).
COSTA RICA. Guanacaste:Liberia, Bank of Rio Colorado,Rinc6n de Vieja, L. 0. Williamset
al. 26618 fl (F); Tilaran,Quir6s 1092 bud (CR 2x, F 2x), 1458fr (F); Standley & Valerio44213 bud
(US), 45662 bud (US), 45744 bud (US); Poveda 836 fr (CR).
Local names and uses. Belize: tancache (Turner 7); Costa Rica: talcacao
(Quiros 1092, 1458). The leaves and bark are aromatic; the bark anesthetizes
the tongue (Poveda 836; Standley on label of Quiros 1092). Used for making
handles of crushing-mills,accordingto Quiros 1092.

8. PilocarpusgoudotianusTulasne, Ann. Sci. Nat. Bot. s6r. 3. 7: 284. 1847 (as

"P. goudotiana"); Englerin Martius,Fl. bras 12(2): 138. 1874;Duval,
Recherches Jaborandis52. 1905.
Shrub or small tree to 7.5 m tall; branchlets 3-7 mm in diam., dull grayish- or
reddish-brown;perules of terminalbuds often ampulliformand rather abruptly
ending in a long, blunt-pointedtip. Leaves alternate, usually crowded at tip of
branchlets, 3-foliolate, occasionally some 1-foliolate,or imparipinnate1-jugate,
with stalked leaflets, subcircularor obovate when 3-foliolate, 6-18(-27) x 5-17
(-25) cm; petiole semiterete, strongly ribbed, slightly winged or not, 1.5-7 cm
long and 0.1-0.2 cm thick, wings to 0.1 mm broad; rachis, if any, 1-4(-5.5) cm
long; petiolules 0-5 mm long, of terminalleaflets to 27 mm; leaflet blades elliptic,
subovate, or occasionally oblong, 3.5-10(-17) x 1.5-5.5(-8) cm, lateral leaflets
unequalat base, roundedor obtuse at apex, the very tip emarginate,marginthick
and revolute; leaflet blades shining above, dull and pale below, (densely) pilose
on both sides, venation brochidodromous-camptodromous, prominent, midvein
plane above. Racemes solitary, subterminal,erect, 10-30 cm long and 3-6 cm
wide, with numerous flowers developing in basipetal fashion and soon all flow-
ering together; peduncle absent or obsolete; rachis 2(-4) mm in diam. at base,
(densely) pubescent with thin, spreadinghairs 0.1-0.4 mm long; bracts broadly
triangular,0.7-1.1 x ca. 1 mm; pedicels inserted at 60-90?, indumentlike that of
the rachis;bractlets2, alternate,subobliquelyinsertedjust below the calyx. Flow-
ers ca. 9-11 mm in diam.; calyx 5-toothed or 5-lobed, ca. 0.8-1.2 mm high;lobes
or teeth subvalvate, some of them overlappingor not, depressedlyovate or round-
ish, (thickly) coriaceous; petals subvalvate, 3.5-4.5 x 2.2-3 mm, apex inflexed
through0.4-0.6 mm, coriaceous, keeled and impressed above, venation parallel
to acrodromous, nerves branched towards margin, median nerve thicker; fila-
ments adnate at base to the disc, subulate, 2.5-3.5 mm long and 0.4-0.7 mm
thick, glabrous; anthers ovoid, ca. 1.4-1.7 x 0.9-1.5 mm, pale yellowish, with
an oblong, subapicaldorsal gland;disc as high as the ovary, ca. 0.7-1.2 mm high
and ca. 2.4(-4) mm in diam., (occasionally strongly)plicate, surroundingthe base
Pilocarpus 161

of the filaments, (sub)glabrous, tuberculate in fruit; carpels adnate to the disc,

0.7-1 mm high, punctate with very small outlets of internal glands; ovules 2,
superposed; style inserted among the tips of the carpels, clavate or terete, 0.3-1
mm long and 0.5 mm thick, free for 0.2-0.6 mm; stigma capitate, 0.2-0.3 x 0.5-
0.6 mm. Mericarps obovate or subelliptic in outline, 9-13 x 17-10 mm, the ventral
axis rather straight, dorso-apically rounded, slightly mucronate or rounded at
very apex, with glands 0.2 mm in diam., glabrous or rarely shortly hairy towards
tip, sterile mericarps usually hairy at tip, dehiscent up to /2-2/3 below tip; seed
1 per mericarp, kidney-shaped, 7-8.2 x 5-6.5 x 4.5-5 mm, flattened at base,
rounded and slightly keeled on the back, obtuse at apex, testa black brown,
shining, finely colliculate with collicles 0.05 mm in diam., wrinkled when imma-
ture; hilum tear-shaped; cotyledons ? unequal with ears 1 mm long, radicle 1-
1.2 mm long, plumule 0.1 mm long.
This species is near to Pilocarpus racemosus, which see.

Key to the Subspecies of Pilocarpus goudotianus

1. Tips of branchlets,perules of terminalbuds, and petioles densely pubescent; calyx lobes
0.5-0.6 x 1.3-1.7 mm; petals purplish-brown,usually hairy beneath. 8a. subsp. goudotianus.
1. Branchletsand perules of terminalbuds glabrous;petioles subglabrous;calyx teeth 0.3-
0.5 x 1-1.1 mm; petals violet-brown-orangetowards the base with yellowish-greentip.
8b. subsp. heterochromus.

8a. Pilocarpus goudotianus Tulasne subsp. goudotianus.

Branchlets densely pubescent with spreading hairs varying from 0.1-0.6 mm,
the hairs light brownish when young, becoming whitish and deciduous; perules
of terminal buds densely brown-pubescent. Petiole densely pubescent with
spreading hairs 0.1-0.2(-0.4) mm; base of leaflets shortly attenuate, shortly cu-
neate, or rounded, of terminal leaflets also to long-attenuate. Calyx lobes ca. 0.5-
0.6 x 1.3-1.7 mm, minutely puberulous with hairs 0.1-0.2 mm long; petals pur-
plish-brown when dried, glabrous above, (rather densely or sparsely) pubescent
beneath with spreading hairs 0.1-0.2 mm long, occasionally becoming glabrous
at anthesis; filaments purplish; ovary usually provided at tip with hairs 0.05-0.2
mm long, or glabrous; style pilose at base with hairs 0.1-0.2 mm long; stigma
sometimes provided with some hairs 0.01 mm long. Seeds 7-8.2 x 5-6.5 x 4.5-
5 mm; hilum 2.8-3 x 1.1-1.3 mm.

Key to the Varieties of Pilocarpus goudotianus subsp. goudotianus

1. Leaflets(sub)coriaceous,pubescenton both sides or becomingsubglabrous;raceme3-4 cm
wide, pedicels usually 10-16 mm long. 8a-l. var. goudotianus.
1. Leaflets chartaceous,ratherdensely pubescent on both sides, somewhatsoft to the touch
below; racemes up to 6 cm wide, pedicels 20-24 mm long. 8a-2. var. mollis.

8a-1. Pilocarpus goudotianus Tulasne subsp. and var. goudotianus. Fig. 52A-D.
Pilocarpus alvaradoi Pittier, Bol. Corn. Industr. (Caracas)34 (also cited: Arb. arbust. nuev.
Venez. decadas 2, 3): 27. 1923;Trab. Mus. Com. Venez. 7: 338. 1930, syn. nov. TYPE.
Alvarados.n., Venezuela. Lara:Nr. Barquisimeto,Jan 1922,fl (holotype, VEN-5844).
Pilocarpus racemosus sensu Arnoldo, Zakflora58. 1954;Zakflora(ed. 2) 190. 1964,non Vahl,
syn. nov.
Leaflets (sub)coriaceous, pubescent on both sides with spreading, thin hairs
0.2-0.4 mm, more densely pubescent at base and on midvein, or becoming sub-
glabrous. Racemes 3-3.5(-4) cm wide; pedicels (3.5-)10-16 mm long and 0.05-
162Flora Neotropica
Pilocarpus 163

0.1 mm thick, in fruit ascending and 12-25 mm long. Petals 3.5-4.5 x 2.2-2.7
mm; disc glabrous.
Type. Goudot s.n., Colombia. Tolima: Between Aipe (at Rio Magdalena)and
Ataco, Apr 1844, fl (holotype, P; isotype, K ex Herb. Hooker, photo NS 2861
made by NY, NY).
Distribution.Colombia, Venezuela, Aruba, Curagao. Deciduous forests, cha-
paral, and desert, dry, stony places on (calcareous) slopes; alt. 300-1500 m.
Usually common, but rare in Curagao.FloweringDec-Jan(-Mar), and Jun-Aug.
Fig. 50.
Specimens examined. SOUTHERN DUTCH ANTILLES. Curagao:Arnoldo 97 st (U). Aruba:
Arnoldo 231 fl (U); Boldingh 6481 st (U); Stoffers 1745-A dec. fr (U), 6259 fl & fr (U 9x).
COLOMBIA.Locality unknown:Wagner307 bud (F, NY (photos)),s.n. bud (ZT).
VENEZUELA. Falc6n: Benitez de Rojas 1808 fl & fr (U); Curran & Haman 507 fr (GH, US), 540
fl & fr (GH, NY, US 2x); Lasser 2761 fl & fr (VEN), 4335 fr (VEN); Lasser & Foldats 3156 fl (VEN);
Steyermark 99155 fr (M, S, VEN); Steyermark & Braun 94720 fl (F, NY, P, US, VEN); Tamayo 755
fl (VEN). Lara: Agostini 1962/70 fl (US, VEN); Aristeguieta 3969 bud (NY 2x, U, VEN); Benitez de
Rojas 1704 fr (U); Karsten s.n. (Tocuyo) fl & fr (LE, W); Mocquerys s.n. (1893-1894, Duaca) fl (K,
NY, P 4x, S, U, US, VEN); Pittier 13102fl (A, F, G 3x, K 2x, M, MO, NY, US, VEN); Steyermark
55569 st (F, NY); Steyermark & Carreho Espinoza 111690 bud (VEN); Steyermark et al. 110262 fl
(VEN); Tamayo 1109 fr (F, S, UC, VEN).

Local names and uses. Aruba and Curagao: burachi; palu cayente ("fiery
wood") (Arnoldo, 1954, 1964). Venezuela (Falcon): borrachera (Lasser 2761,
4335; Tamayo 755); matasarna(Tamayo 755). Arnoldo 97 mentions the use of
parts of the twigs in rum to make the drink stronger; (is this the origin of its
rareness in curacao?). The name "borrachera"("burachi")means drunkenness.
Tamayo 755 reportsthat the asses become the worse for drinkwhen they eat the
plant. Wood used for barrels(Tamayo 755).
8a-2. PilocarpusgoudotianusTulasne subsp. goudotianusvar. mollis(Cuatrecasas)
Kaastra, Acta Bot. Neerl. 26: 486. 1977. Fig. 52E-F.
Pilocarpusalvaradoi Pittiervar. mollis Cuatrecasas,Revista Acad. Colomb. Ci. Exact. 8: 467.
1952.

Shrub to more than 2 m tall. Leaflets chartaceous, rather densely pubescent

on both sides, somewhat soft to the touch below. Racemes to 6 cm wide, the
pedicels 20-24 mm long and 0.05 mm thick. Petals 4-4.3 x 2.8-3 mm; disc beset
with few hairs 0.1-0.2 mm.
Type. Haught 4480, Colombia.Guajira:70 km SE of Riohacha, along highway,
7 Dec 1944,fl (holotype, US-1709397,photo 49578 made by F, F; isotypes, COL
n.v., K, NY ex US, UC ex US).
Distribution.Known only from the type locality. Forest; alt. 80 m. Fig. 50.
This variety is distinguishedby the dense hair-coveringof the leaves (which is
sometimes also shown in var. goudotianus, however), the longer pedicels at
anthesis, and the broaderpetals. The flowers are conspicuous, very darkpurple
with yellow anthers, accordingto Haught.

FIG. 52. Pilocarpus goudotianus. A-D, var. goudotianus. E-F, var. mollis. G, subsp. hetero-
chromus. A, habit (Karsten s.n., from Tocuyo, W). B, inflorescence (Pittier 13102, M). C, flower
(Pittier 13102, M). D, mericarp(Karstens.n., from Tocuyo, W). E, indumentof lower side of leaf
(Haught 4480, type, UC). F, inflorescence (Haught 4480, type, UC). G, flower (S. Galen Smith 1136,
type, GH).
164 Flora Neotropica

8b. Pilocarpus goudotianusTulasne subsp. heterochromusKaastra, Acta Bot.

Neerl. 26: 486. 1977. Fig. 52G.
Tree ca. 7.6 m tall; branchlets and perules of terminalbuds glabrous. Petiole
subglabrous;leaflets to 14.5 x 8.3 cm, attenuate at base, coriaceous. Racemes
3-3.5 cm wide. Calyx (sub)glabrous,with teeth of 0.3-0.5 x 1-1.1 mm; petals
3.5-4.5 x 2.2-2.7 mm, brown-orangeand green-tippedwhen alive, the upperside
tinged with violet towards base and yellowish-green towards apex when dried,
quite violet below when dried, glabrous;filamentspurplish-tingedat base; gyn-
oecium glabrous. Seeds (only 1, immature known), ca. 9 x 8 mm in outline,
hilum 4.5 x 2.2 mm.
Type. S. G. Smith 1136, Colombia. Huila: Upper basin of Rio Magdalena,
CabreraLajas, ca. 11 km E of Villavieja on road to Baraya, 4 Jul 1950, fl & fr
(holotype, UC-963230;isotypes, F-1435630,GH, MO, US-2047485).
Distribution. Known only from the type locality. Dissected plateaus ("bad-
lands") with woody plants mainly restricted to gullies and stream-bottoms;alt.
460 m. Fig. 50.
This taxon is distinguished by glabrous branchlets and petioles, by smaller
calyx teeth, and by the colors of the petals. The indumentof the leaflet blade,
and the pedicels are similarto those of var. goudotianus.

9. PilocarpusgiganteusEnglerin Martius,Fl. bras. 12(2): 136, t. 29. 1874;Engler

in Engler and Prantl, Nat. Pflanzenf. 3(4): 158, fig. 93C. 1896;Duval,
Recherches Jaborandis56. 1905;Albuquerque,Anais Acad. Brasil. Ci.
40: 507, t. 1, fig. 8. 1968. Fig. 53.
Pilocarpus macrocarpus Engler in Martius, Fl. bras. 12(2): 138, t. 31, fig. 2. 1874;Engler in
Englerand Prantl, Nat. Pflanzenf.3(4): 158, fig. 93D. 1896;Duval, RecherchesJaborandis
54. 1905;Englerin Englerand Prantl,Nat. Pflanzenf.(ed. 2) 19a: 280, fig. 129D. 1931,syn.
nov. TYPE. Sellow 5998, Brazil,(Sao Paulo:Sao Pauloor Santos, 1829,accordingto Herter
(1949)),fr (holotype, B, destroyed;fragment(fruitsonly), P-type Herb. Baillonmisit Asch-
erson ex B; isotype, B, destroyed, photo 12522made by F, F, NY).
Shrub 1.5-4 m tall, indumentum,if any, yellow-brown,0.05-0.2 mm;branchlets
7-15 mm in diam., grayish-brown,minutely pubescent when young. Leaves al-
ternate, subverticillate, simple; petiole semiterete, 1.5-15 mm long and ca. 3-4
mm thick, minutely pubescent; blade narrowly obovate, (11-)20-48 x (3-)4-10
cm, tapering into the narrowly cuneate base, ? acuminate or rarely obtuse at
apex, the very tip obtuse and shallowly emarginateor retuse, marginplane or
(sub)revolute, the blade chartaceous, dull grayish-greenon both sides, glabrous
when adult, venationbrochidodromous,? prominent,costa prominulousor plane
above, very prominent below. Racemes several, erect or pendulous, inserted
axially, laterally among or below the leaves, 10-15 x 1 cm at the opening of the
first flowers, but the part that has finished flowering enlarging immediatelyto
become up to 40 x 3 cm in fruit, minutely pubescent; rachis 1-2 mm in diam.,
in fruit to 4 mm in diam., becoming transversely striped due to cracks; bracts
very broadlytriangular,ca. 0.5-0.8 x 0.8-0.9 mm; pedicels ascendingto spread-
ing, 2-3 mm long and 0.5 mm thick, in fruit to 3-11 mm long and 1.5 mm thick;
bractlets 2, alternately inserted at variable height. Flowers 6-7.5 mm in diam.;
calyx 5-lobed, 0.6-1.2 mm high; lobes valvate to subquincuncial,depressedly
ovate or triangular,0.5-0.6 x 0.8-1.4 mm, roundish to acute at apex, thickly
coriaceous, minutelypubescent without;petals subvalvate, 2.8-3.5 x 1.8-2 mm,
the apex inflexedthrough0.4-0.5 mm, marginslightlyundulate,thinlycoriaceous,
semitransparent,yellow-green, becoming purplish, adaxially slightly keeled to-
Pilocarpus 165

FIG 53. Pilocarpus giganteus. A habit Schott Herb Bras 4757 type W) B flower.
(Scho. t
Herb. Bras. 4757, type, W). C, fruit (Sucre et al. 10192, U).
166 Flora Neotropica

wards tip, abaxiallyminutelypubescent or sometimes subglabrous,venation par-

allel to subactinodromous;filamentstruncate, firm, 2.3-2.5 mm long and 0.4-0.5
mm thick, glabrous; anthers subcircular to ovate in outline, recurved, 0.9-
1.1 x 0.9-1 mm, with a dorsal ovate gland; disc + 10-lobed,0.5-1 mm high and
2.2-2.7 mm in diam., glabrous; carpels 0.7-0.9 mm high, densely minutely pu-
bescent, with internalglands; style inserted midwayon the carpels, 0.4 mm long,
the united part 0.1 mm long but to 1 mm after anthesis; stigma capitate, 0.3-
0.4 x 0.6-0.8 mm. Mericarps subobovate in profile, (10-)15-16(-19) x (8-)10-
12(-15)mm, dorso-apicallyrounded,the very apex obtuse, notched or not, dense-
ly minutelypubescent, with glands to 0.3 mm in diam.; seeds subovoid, ca. 10.5-
11 x 6-6.5 x 5-5.8 mm, flat at base, the ventral axis + straight,roundedat tip,
the micropyle below the tip, slightly keeled on the back, testa brown, finely
colliculate with interspaces 0.1 mm; hilum ca. 3-4.5 x 1-1.5 mm; cotyledons
greenish-creamish,slightlyunequalwith ears 1 mm long; radicle ca. 1.1 mm long,
enclosed between the ears; plumule0.5 mm, strigilloselike the radicle.
Type. Schott Herb. Bras. 4757, Brazil. Rio de Janeiro,fl (lectotype, W, drawn
in Fl. bras. 12(2):t. 29).
Distribution. Brazil, Minas Gerais (rare), Sao Paulo, and besides in Rio de
Janeiro. Primaryand secondary forest, once sub-bosque; alt. 140 m (I record).
FloweringJan-Feb, and May-Oct. Fig. 43A.
Specimens examined. BRAZIL. Minas Gerais: Sucre (& Martinelli & Silva) 10192 fr (U). Sao
Paulo: Gaudichaud 644 fl (P); Lofgren CGGSP 4178 fl (NY, SP); Usteri SP 19824 fl (SP). Rio de
Janeiro: Constantino 124 fl (RB), RB 15261 fl (RB 2x); Duarte 5896 fr (U); Glaziou 14587 fl & fr (C,
G, K, LE, P 2x, R); Horto Florestal 6127 fl (RB); J. G. Kuhlmann 16396 fl & fr (U); Occhioni 324
fl (RB 2x), 325 fl (RB 2x); Pereira4005 fl & fr (RB 2x), 7255fl (LP, M);Riedel 470 fl & fr (E, LE 2x).
Locality unknown:Riedel s.n. fl (K, photo, NY; P).
Pilocarpus macrocarpus was based only on an inflorescence with flowers and
fruits. Although Engler supposed it to be near to P. riedelianus, it has more in
common with P. giganteus: the inflorescence is lateral, not terminal as in P.
riedelianus; perianth, ovary, and fruits are pubescent, not glabrous;leaves and
racemes of the isotype are like those of P. giganteus. The infructescence is
curved near the base; this indicates a pendulous position as in P. giganteus. The
mericarpshowever, are somewhat larger:to 19 x 15 x 7 mm. Because all other
charactersagree well with P. giganteus I reduce P. macrocarpus to synonymy
under this species.
Planchon (1875) apparentlybased his description of Jaborandi, viz. the fruit
and one of the inflorescences, partly on the present species.
Pereira 7255 is entirely glabrousexcept for the perules. The bracts and calyx
are ciliate.
The specimens from Sao Paulo show some dorsal strigae 0.1-0.2 mm on the
anthers.
10. Pilocarpus peruvianus (Macbride) Kaastra, Acta Bot. Neerl. 26: 487.
1977. Fig. 54.
Pilocarpusspicatus Saint-Hilairevar. peruvianusMacbride,Field Mus. Nat. Hist., Bot. Ser. 13:
676. 1949.

Shrub 2-10 m tall; branchlets2-5 mm in diam., grayish-brown,shining when

young, glabrous; perules of terminal buds triangular,to 2.5 x 2 mm, acute or
truncate, strigillose. Leaves alternateto (sub)oppositeor ternate, crowdedbelow
the racemes, simple, glabrous;petiole semiterete, 2-5 mm long and 2-3 mm in
diam., scarcely or slightlywingedby the decurrentbase of the blade, transversely
Pilocarpus 167

FIG. 54. Pilocarpus peruvianus (Schunke V. 1983, F). A, habit. B, flower, C, part of infructes-
cence. D, seed.
168 Flora Neotropica

cracked; wings distally up to 0.2 mm broad; blade usually narrowly elliptic to

narrowly oblong, or obovate, 13-30 x 4.5-9 cm, just below the flowers occa-
sionally smaller, attenuate or narrowly cuneate, and decurrent at base, ? acu-
minateat apex, the very tip obtuse, emarginate,or retuse, the marginsubundulate,
thick and strongly revolute, the leaf blade subcoriaceous, deep green above,
somewhatpalerbeneath, shining,bullatebetween the secondarynerves, venation
brochidodromous,prominenton both sides, costa + prominentabove. Racemes
solitary, terminal, erect, 15-40 x 0.7-1.2 cm, with numerous flowers, subgla-
brous; base of rachis 1.5-2.5 mm thick and provided with some scars of sterile
bracts; bracts very broadly or depressedly triangular,0.4-0.7 x 0.7-1 mm, the
basal ones somewhat larger,pedicels inserted at 45-90?, 0.5-2 mm long and 0.7-
1 mm thick, in fruit 3-4 mm long; bractlets 2, alternateor suboppositenear base
of pedicel. Flowers ca. 6.5 mm in diam., glabrous, or the ovary subglabrous;
calyx 5-lobed, ca. I mm high, lobes separate-valvate, depressedly ovate, 0.5-
0.6 x 0.9-1.2 mm, rounded at tip, unequal; petals valvate to subimbricate,3-
3.2 x 1.5-1.9 mm, apex inflexed through 0.7 mm, coriaceous, semitransparent,
green-yellow, the upper side keeled towards the tip with obsolete impressions,
venation subparallel, the median nerve thicker; filaments truncate, ca. 2.7-
3 x 0.4-0.5 mm; anthers heart-shaped, recurved, 0.8-1.4 x 0.7-1 mm, with a
dorsal gland; disc 0.7-0.8 mm high and 1.8-2 mm in diam., 5-10-plicate; carpels
0.8-1.0 mm high, protrudingbeyond the disc, with internalglands, glabrous, or
with some hyaline hairs 0.05-0.1 mm near the stigma; ovule I per carpel; style
obconical, not protrudingbeyond ovary; stigma capitate, 0.3-0.4 x 0.4-0.7 mm.
Mericarps ? ellipsoidal, 12-16 x 10-13 mm, dorso-apically rounded, the very
apex obtuse with an angle of ?90?, punctate with glands 0.3-0.5 mm, glabrous,
the wall very prominentlynerved and somewhat wrinkled in the lower dorsal
part, dehiscent up to 2/5-1/2 below the tip; seed 1 per mericarp,kidney-shaped,
ca. 10 x 7-7.5 x 6-6.5 mm, flattenedat base, keeled on the back, the micropyle
ventraland not apical;testa chestnut-brown,irregularlyreticulate-colliculatewith
interspaces 0.05-0.1 mm; hilumjust above the middle, tear-shaped, 3 x 2 mm;
cotyledons unequal, auriculateto 1.8 mm, enclosing the radicle 0.7-0.8 x 0.6 mm
and the 2-leafed plumule 0.2-0.3 mm; plumule ciliate with brown, shininghairs
0.05-0.1 mm long.
Type. Ule 9500, Brazil. Acre: Rio Acre, Seringal S. Francisco, upstream of
Tacna (Bolivian border village), May-Aug 1911,fl (holotype, F? n.v.; isotypes,
G, L, U, UC ex B 2x).
Distribution.Peru (San Martin,Hufanuco,and Loreto); Brazil (Acre). Forests,
once terrafirmewith dry clay; alt. 150-400m. Floweringin the dry season, May-
Aug. Fig. 43A.
Specimens examined. PERU. Loreto: Lower Rio Huallaga,Santa Rosa, Ll. Williams4878 fr (A,
F, US). San Martin:MariscalCaceres, Tocache Nuevo, nr. Rio Tocache, SchunkeV. 3290 fr (F, G
2x); Huanuco:Pachitea, Honororia,20 km upstreamof confluenceRio Pachitea with Rio Ucayali,
Schunke V. 1983 fl & fr (F, G 2x, MO).
BRAZIL. Acre: Rio Acre, SeringalIracema,Ducke RB 23828 fl & fr (U).
Local name. Peru, Loreto: sapoto jaru (Ll. Williams4878).
Macbride(1949) was not sure whether LI. Williams4878 and Ule 9500 should
be referredto the same taxon or not. With more materialavailablenow, it is clear
that they indeed do so. They only superficiallyresemble Pilocarpus spicatus.
ThereforeI elevate var. peruvianus to species level.
The type locality is in Brazil, not in Peru as Macbride mentions (Ule, 1914:
101).
Pilocarpus 169

11. PilocarpusriedelianusEnglerin Martius, Fl. bras. 12(2): 135. 1874;Englerin

Engler and Prantl, Nat. Pflanzenf. 3(4): 158. 1896. Fig. 55.
Shrub or treelet 0.6-6 m tall; branchlets 3-10 mm in diam., dark-reddish-or
grayish-brown,shining at apex, glabrous, providedbelow the racemes with nar-
rowly triangularhypsophylls up to 5 mm long and hairtowards their tips. Leaves
alternateor subverticillate,simple, glabrous;petiole semiterete with small wings,
1-10 mm long and 2-3 mm thick, wings 0.1-0.2 mm broad; blade (narrowly)
elliptic or (narrowly)(ob)ovate, 10-37 x 4.2-11.5 cm, attenuateand decurrentat
base, subacuminate,obtuse, or acute at apex, the very tip emarginateor entire,
margin revolute, the blade (sub)coriaceous, venation brochidodromous,main
nerves prominenton both sides or impressed above, costa plane and longitudi-
nally wrinkledor not. Racemes terminal, erect but later on recurved, ca. 18-45
cm long and 0.7 cm wide, in fruit at least 65 cm long and possibly up to ca. 90
cm, with many flowers developing in basipetal fashion; rachis 1.2-2 mm thick at
base, to 5 mm thick in fruit, minutelypuberulouswith hairs 0.1 mm long, becom-
ing glabrous;bracts alternate,depressedly triangular,concave, 0.3-0.5 x 0.6-0.7
mm, glabrous; pedicels inserted at 45?, 0.5-2 mm long and 0.4 mm thick, up to
6 mm long and 1.5 mm thick in fruit, indumentlike that of the rachis; bractlets
2, alternateor subopposite. Flowers ca. 5-6 mm in diam., mostly quite glabrous
at anthesis but occasionally the perianth minutely puberulous in bud; calyx
5-lobed; lobes separate, very broadly triangular,ca. 0.5 x 0.6 mm, rounded at
apex, glabrous, or subglabrousbelow, minutelypubescent in fruit with hairs 0.1
mm; petals reflexed at anthesis, ovate, ca. 2.4-2.6 x 1-1.3 mm, apex inflexedup
to 0.2 mm, margininflexed towards tip, papery but coriaceous in age, semitrans-
parent, brown-purplishor dark red, (sub)glabrous,nerves paralleland branched
towards margin,median nerve thicker; filamentstruncate, flattenedtowards the
base, ca. 1.5-2.3 x 0.2 mm, red-purplish;anthers dorsifixed below the middle,
ovate in outline, strongly recurved, ca. 0.8-1 x 0.6-1 mm, with a dorsal gland
0.2 mm; disc 5-lobed, 0.3 mm high and 1.3-1.8 mm in diam., paler thanthe ovary,
glabrous or puberulous;carpels ca. 0.6 mm high, projecting0.4 mm beyond the
disc, glabrousor puberulous;ovule I per carpel; style obsolete, but becomingup
to 0.5 mm long after anthesis; stigma subsessile, capitate, 0.2-0.3 x 0.4-0.5 mm,
green. Mericarpsobovoid, 11-15 x 8-10 mm, dorso-apicallyrounded, obtuse at
apex, the very tip scarcely notched, punctate with dark glands up to 0.6 mm in
diam., minutely puberulouswith hairs 0.1 mm, dehiscent up to 1/2/3 below tip;
seeds obovoid to reniform, ca. 9-10 x 6 mm in outline, with a curved blunt tip,
carinate on the back; embryos (1 seed studied): 2 well developed and some ves-
tigial ones per seed, the cotyledons unequalwith ears to 1.5 mm, punctate;radicle
blunt, to ca. 1.7 x 1 mm, projectingbeyond the ears, strigillose with brownhairs
0.1 mm; plumule to 0.8 mm long, strigillose like the radicle.
Type. Riedel 438, Brazil. Bahia: Nr. Ilheos, Sep 1821, bud & fl/fr (lectotype,
LE; isolectotype, B-Herb. Engler, destroyed, photo 12525made by F, F, NY).
Distribution.Brazil, Bahia and Espirito Santo. Dry, rocky places in secondary
and tabuleiro forests, abundantin sub-bosques; alt. 65-110 m. Flowering Jan-
Jun(-Sep). Fig. 43A.
Specimens examined. BRAZIL. Pernambuco: Ducke & Andrade Lima 119 fl/fr (R). Bahia: Blanchet
1716 fl (G, P); Duarte 5987 fr (LP, U); Riedel 437 fr (LE); Velloso 994 fl (R), 1000 fl (R 7x); possibly
from Bahia: Riedel s.n. fl (E), s.n. fl/fr (K, photo, NY; P). Espirito Santo: J. G. Kuhlmann 347 fl (U);
Sucre 8335 fl/fr (U), 8706 fl/fr (U); Sucre (& Soderstrom) 8947 fl/fr (RB).

Vegetatively this species sometimes resembles Pilocarpus spicatus var. spi-

170 Flora Neotropica

FIG. 55. Pilocarpus

riedelianus. A, habit (Duarte 5987, U). B, inflorescence (Sucre 8706, U). C,
embryos (Duarte 5987, LP).
Pilocarpus 171

catus. The flowers differ in color and in some more characters; the fruits are
much larger.The indumentof fruits and calyx becomes more dense afteranthesis.

12. Pilocarpusspicatus Saint-Hilaire,Bull. Sci. Soc. Philom. Paris (1823): 131.

Sep 1823;Mem. Mus. Hist. Nat. 10: 145. between 21 Apr and 28 Jun
1823,nom. nud.; Mem. Mus. Hist. Nat. 10: 360. after 6 Nov, 1823;de
Candolle, Prodr. 1: 728. 1824; Sprengel, in Syst. veg. ed. 16. 1: 954.
1824/1825;Saint-Hilaire,Fl. Bras. mer. 1: 83. 11 Jun 1825;Hist. rem.
Bres. Par. 1: 146, t. 16. 4 Nov 1825; Spach, Hist. nat. veg. 2: 345.
1834;Dietrich, Syn. pl. 2: 1017. 1840(all citations so far, except for
Sprengel, as "P. spicata"); Engler in Martius, Fl. bras. 12(2): 133.
1874; Geiger, Ber. Deutsch. Pharm. Ges. 7: 395, 414, t. 35 a and b.
1897; Duval, Recherches Jaborandis43. 1905, pro parte; Macbride,
Field Mus. Nat. Hist., Bot. Ser. 13: 675. 1949; Albuquerque,Anais
Acad. Brasil. Ci. 40: 507. 1968.
Shrubor treelet 2-5 m tall, glabrousor pubescent;branchlets3-7 mm in diam.,
grayish- or reddish-brown,shining when young; perules of terminalbuds trian-
gular, 1.5-4 x 3 mm, strigillose with hairs 0.2-0.3 mm. Leaves alternate,
(sub)opposite or ternate, crowded towards apex of branchlets, simple; petiole
semiterete, slightly canaliculatetowards tip or not, slightly winged to 0.3 mm,
1.5-20(-50) x 1.5-2 mm, often becoming transverselycracked in age; blade (nar-
rowly) elliptic to obovate, rarely some ? ovate, 4.5-18.5(-25) x 1.5-6(-8) cm,
narrowlyor rarelybroadlycuneate or (long-) attenuateat base, ? decurrentalong
petiole, rounded, obtuse, or subacuminateat apex, the very tip sometimes to 1
cm long and emarginateor entire, marginthick and stronglyrevolute, blade char-
taceous to subcoriaceous,slightlypalerbeneath, ? shiningabove, venationbroch-
idodromous to camptodromous,principal nerves prominent, costa above plane
and wrinkled or not, prominentbelow. Racemes 1-several, subterminal,erect,
13-45 x 0.5-1.5 cm, with numerousflowers developingin basipetalfashion, gla-
brous, or rachis and pedicels pubescent with spreadinghairs 0.05-0.1(-0.3) mm
long; rachis 1-2 mm thick at base; bracts broadly to depressedly triangularor
suborbicular,0.3-0.5 x 0.3-0.5 mm, basal bracts up to more than twice as large
as the upper ones; pedicels inserted at 90?; bractlets 2, alternate, obliquely or
transversely inserted, one close to the calyx. Flowers 4.5-7 mm in diam.; calyx
4-5-lobed; lobes valvate to subquincuncial,very broadly to depressedly ovate-
triangular,0.2-0.6 x 0.5-0.8 mm, acute to rounded, coriaceous, glabrous or in
bud with some hairs0.05 mm long; petals subvalvate,ovate or very rarelyelliptic,
the tip 0.2-0.3 mm inflexed, (thinly)coriaceous and usually semitransparent,yel-
low-greenishwhen fresh, yellow-brownishwhen boiled, slightly carinatetowards
tip and with slight impressionsabove, shining below when young, glabrous, ve-
nationparallel,the mediannerve thicker;filamentssemiterete, 1.5-2(-2.8) x 0.2-
0.3 mm, glabrous; anthers suborbicular, ca. 0.6-0.9 mm, pale yellow, with a
dorsal, ovate gland; disc 0.3-0.6(-0.8) mm high and 1.4-1.8(-2.5) mm in diam.,
5- to slightly 10-plicate,paler than the ovary, glabrousor strigillose with tawny
hairs to 0.5 mm long; carpels ca. 0.5-0.7 mm high, with glands, glabrous or
strigillose like the disc; ovule 1 per carpel; style obconical, ca. 0.2-0.3 mm long,
not projectingbeyond the ovary at anthesis but growing after flowering;stigma
capitate, 0.2-0.4 x 0.2-0.5 mm. Mericarpssubovoid, 6-9 x 4.5-6 mm, dorso-api-
cally rounded, the very apex notched or not, with glands 0.3-0.4 mm, glabrous
or minutelypubescent when young, dehiscent up to /? or % below tip; seed 1 per
mericarp,expelled immediately,(ob)ovoid to reniform,5.5-6 x 4 x ca. 3.6 mm,
slightly keeled on the back and curved at tip, testa chestnut-brown,irregularly
172 Flora Neotropica

finely reticulatewith interspaces0.05-0.1 mm; hilumto 2.5 mm long; cotyledons

subequal with ears I mm; radicle thick, blunt, ca. 1.5 mm long; plumule to 1.5
mm, strigillose with tawny hairs to 0.2 mm long.
The indumentof ovary and disc is not consistently correlated with other fea-
tures, and cannot be used as a discriminatingcharacter, as Engler (1874: 133)
alreadycarriedinto effect. The leaves of some specimens are very long-attenuate
or narrowlycuneate at base and accuminate at apex. These specimens resemble
Pilocarpuspauciflorus, but the inflorescenceis sometimeslonger, and the flowers
are much smaller in diam. Other specimens have attenuate and less gradually
narrowingleaf blades, with the apex obtuse or rounded.
Three subspecies can be distinguished, one divided into two varieties. The
varieties are very close to each other and especially specimens in bud are some-
times difficultto identify.

Key to the Subspecies of Pilocarpus spicatus

1. Pedicels 4-5 mm long; racemes 1-1.5 cm wide; flowers 4.5-5.5 mm in diam.; filaments
subulate. 12c. subsp. aracatensis.
1. Pedicels 0-5 mm long; racemes 0.5-1.5 cm wide; flowers 4.5-7 mm in diam.; filaments
subulateor truncate.
2. Flowers4.5-6 mm in diam.;pedicels 0-3.5 mm long; racemes0.5-1 cm wide;petals 1.9-
3 mm long; filaments subulate or truncate; leaves subglabrousor pubescent below.
12a. subsp. spicatus.
2. Flowers 6-7 mm in diam.; pedicels 2-5 mm long; racemes0.5-1.5 cm wide; petals 2.5-
2.7 mm long; filamentssubulate;leaves densely pubescentbelow.
12b. subsp. longeracemosus.

12a. Pilocarpusspicatus Saint-Hilairesubsp. spicatus.

Shrub or treelet, minutely pubescent or subglabrous.Racemes to 45 cm long
and I cm wide; pedicels 0-3.5 mm long. Flowers 5- or 4-merous, 4.5-6 mm in
diam.; petals 1.9-3 mm long and 1-1.9 mm broad; filaments truncate or ? sub-
ulate.

Key to the Varieties of Pilocarpus spicatus subsp. spicatus

1. Flowers 5-merous;filamentstruncateor obtusish at tip; petals 2-3 mm long; leaves sub-
glabrousbelow. 12a-1.var. spicatus.
1. Flowers 4-merous;filaments ? subulate with acuminateor acute tip; petals 1.9-2.5 mm
long; leaves pubescent or subglabrousbelow. 12a-2.var. lealii.

12a-1. Pilocarpusspicatus Saint-Hilairesubsp. and var. spicatus. Fig. 56A-D.

PilocarpusparviflorusNees & Martius,Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol.Nat.
Cur. 11: 177,t. 30. after 26 Jul, 1823;de Candolle,Prodr.1: 728. 1824,pro syn.; Sprengelin
Syst. veg. ed. 16. 1: 954. 1824/1825,pro syn. TYPE. Wied-Neuwieds.n., Brazil. Rio de
Janeiro:Cabo Frio, fr & fl/fr(lectotype, BR; isolectotypes, BR 2x).

FIG. 56. Pilocarpus spicatus. A-D, var. spicatus. E-F, var. lealii. G-I, subsp. longeracemosus.
K-L, subsp. aracatensis. A, habit (Brade 12560, RB). B, flower with BI, abaxial side of stamen
(Brade 12560, RB). C, infructescence (Pereira 737, RB). D, fruit (Pereira 737, RB). E, indument of
lower side of leaf (Machado RB 75557, RB). F, abaxial side of stamen (Machado RB 75557, RB). G,
indument of lower side of leaf (Harley et al. 16737, U). H, inflorescence (Harley et al. 16737, U). I,
abaxial side of stamen (Harley et al. 16737, U). K, habit (Guedes 553, type, US). L, abaxial side of
stamen (Guedes 553, type, US).
Pilocarpus 173

~~~~~~~~~~~U

RzImm,~~~~~~~~"':.,:.~

2c'

:i %"~a:.
174 Flora Neotropica

PilocarpussubcoriaceusEnglerin Martius,Fl. bras. 12(2):134. 1874;Englerin EnglerandPrantl,

Nat. Pflanzenf.3(4): 157. 1896;Duval, RecherchesJaborandis45. 1905;Albuquerque,Anais
Acad. Brasil. Ci. 40: 508, t. 2, fig. 9. 1968, syn. nov. TYPE. Herb. Warming2454, Brazil.
Rio de Janeiro:Nr. Rio de Janeiro,Dois Irmaos, 18 Jun 1866,fl & fr (lectotype, C, nr /1;
isolectotype, C, nr 12).
Pilocarpusypanemensis Englerin Engler and Prantl,Nat. Pflanzenf.3(4): 157. Mar 1896;Bot.
Jahrb.Syst. 21: Beibl. 54: 27. 12 May 1896;Duval, RecherchesJaborandis48. 1905;Engler
in Englerand Prantl,Nat. Pflanzenf.(ed. 2) 19a: 279. 1931,syn. nov. TYPE. Sellow 2178,
Brazil. Sao Paulo(?):fr (lectotype, K ex B; isolectotype, B, destroyed);Sellow' B 2178, c
2174, same locality, now st (paratype,S ex B).
Pilocarpusspicatus Saint-Hilairevar. subcoriaceus Englerex Duval, RecherchesJaborandis45.
1905,comb. invalid. ex Art. 34(4)ICBN.
Vegetative parts glabrous, or branchlets,petioles, and (mainly)base and costa
of the leaves pubescent with spreadinghairs to 0.05(-0.1) mm. Racemes to I cm
wide, glabrous or pubescent with spreadinghairs to 0.1 mm. Pedicels 0-3 mm
long and 0.5 mm thick, in fruit up to 4(-6) mm long. Flowers 5-merous, rarely
some 4-merous, 4.5-6 mm in diam.; petals 2-3 x 1-1.9 mm; filaments truncate
or obtusish at tip, 1.5-2.4 mm long; anthers recurved;disc 0.3-0.6 mm high and
1.4-1.8 mm in diam.
Type. Saint-Hilaire 72B, Brazil. Rio de Janeiro: Nr. Rio de Janeiro, 1818, fl
(lectotype, P ("pres Laranjeira"); isolectotypes, P ("a la mont du Trapiceiro"),
P (locality not mentioned, with labels added later), F ex P (fragment)).
Distribution.Brazil, Ceara, Bahia, Minas Gerais, Sao Paulo, and mainlyin Rio
de Janeiro. Forests, occasionally also restingaforest but usually low hills to 360
m (1 record). Flowering Apr-Aug(-Sep). Fig. 43C.
Specimensexamined.BRAZIL.Ceara:Allemdo275 fl (R). Bahia:OliveiraRB 82176fr (U); Velloso
96/ fl (R 9x). Minas Gerais: J. G. Kuhlmann 29 fl/fr (RB 2x); Monteiro RB 130242 fl (U). Sao Paulo:
Hemmendorf 169 fl (S 2x): Lofgren CGGSP 4390 bud (SP). Rio de Janeiro: Brade 12560 fl (RB 2x);
Burchell 2972 fl (K, L, P); Casaretto 1260 fl (G), 1780 fr (TO 2x); Constantino RB 16394 bud (RB,
U); Duarte 310 fr (RB); Ducke RB 20479 fl (RB 2x); Dusen s.n. ("Mar 1907" Herb. d'Alleizette) fl/
fr (L); Frazdo RB 7335 fl & fr (K, S, U, US); Gardner 829 fl & fr (BM, K, PHA), s.n. (Rio de Jan.)
fl (CGE); Glaziou 14 fl/fr & fr (R), 41 fl & fr (BR 4x; F, NY (photos); P), 2958 bud (BR, C 2x, P,
PHA), /1855 fl (R), 18971 fl (C, K, LE, P, US); J. G. Kuhlmann 6159 fl (RB 2x); Lund 277 fl (C 2x,
P), s.n. (1833) fl (C); Luschnath s.n. (Rio, 1833) fl (LE), s.n. (Serra d'Estrella, May 1834) bud (BR),
s.n. (Rio de Jan.) bud & fr (OXF); Lutz 640 fl bud & fr (R 2x); Machado 285 fl (RB 2x); RB 75539
fl & fl/fr (U); Comm. Martius s.n. (1841) bud (BR); Miers 3521 fl & fr (K, P), s.n. (Corcovado) fr
(BM); Pereira 686 fl (U), 737 fr (RB 2x), 4567 fl & fr (M), 7101 fl & fr (LP, M); Riedel 78 bud (R),
162 bud (R), 169 fl (R), 466 fl & fr (LE 2x, M), s.n. (R. Jan. 1836) fl (P, U), s.n. ("ad Macahe et Rio
de Janeiro") fl & fr (E, G 2x, GH, K, photo, NY; LE 4x, M 2x, P 2x, W), s.n. fl (C, E, G, GH, K,
LE, M, NY, P 2x, S, US, W); Saint-Hilaire 274 fl (BM, MPU, P); Sampaio 1584 fl (R 2x); Sellow
s.n. (Rio, 1814-1815) fl (LE, P); Sucre 1887 fl/fr (U), 8506 fl & fl/fr (U); Vauthier 5 (?) fl (K, P), 159
fl/fr (G, P), s.n. (Rio) fl/fr (P); Widgren 679 fl (S 2x), s.n. (Rio de Jan.) fr (S); Wied-Neuwied s.n. fl
(GOET); Wilkes s.n. (nr. Rio, 1838-1842) fl & fr (NY, US). State unknown: Herb. Drake s.n. (nr.
jardin bot., Jun 1862) bud (P); Gardner s.n. fl & fr (K); Herb. Persoon 23 fl (L); Sellow 5 (?) fl/fr
(SP), 6 (?) fl (F, NY (photos)).
Local name. Near Rio de Janeiro:jaborandi-miuido-da-restinga
(Machado RB
75539).
Pilocarpus subcoriaceus Englercannot be maintainedas a separatetaxon. The
differences describedby Englervary and occur also in many specimens certainly
belongingto var. spicatus. The pedicels e.g., measurebetween 0 and 2 mm, and
these sizes are not consistently correlated with other characters. The leaves of
the syntypes of P. subcoriaceus are chartaceous or subcoriaceous, and subco-
riaceous leaves were also found in plants belongingto var. spicatus with pedicels
1.5-2 mm long. The plants grow under the same conditions and in the same
general region. Geiger (1897:414) already suggested synonymy but in absence of
ample materialhe did not combine them.
Pilocarpus 175

The original syntypes of Pilocarpus ypanemensis Engler were destroyed in

Berlin. Engler mentioned the two collections (Sellow 2174; 2178) in Fl. bras. 12(2)
under P. spicatus. Duplicates of Sellow 2174 (PR, S, UC, all ex B) however, do
not at all belong to Pilocarpus, nor to other Pilocarpinae. The duplicate in UC
was misidentified by Engler as Esenbeckia riedeliana! In my opinion mistakes
were made in B with these specimens. Sellow B 2178, c 2174 (S ex B) is now
sterile but agrees in the vegetative parts with P. ypanemensis. Sellow 2178, fr
(K ex B, 15 Nov 1907) agrees completely with the description of P. ypanemensis.
This specimen I selected as lectotype; it was identified by Engler as P. spicatus.
The description by Engler of P. ypanemensis, and the lectotype, agree fully with
my concept of P. spicatus var. spicatus. Therefore I reduce P. ypanemensis to
synonymy under this variety, as Geiger (1897: 414) suggested.
Allemdo 275, from Ceara, has large leaves with petioles to 5 cm long; this is
the northernmost collection that is known.

12a-2. Pilocarpus spicatus Saint-Hilaire subsp. spicatus var. lealii (Machado)

Kaastra, Acta Bot. Neerl. 26: 487. 1977. Fig. 56E-F.
Pilocarpus lealii Machado, Bull. Mens. Soc. Linn. Lyon 16: 145. 1947 cum photo; Albuquerque,
Anais Acad. Brasil. Ci. 40: 506, t. 1, fig. 7. 1968.
Pilocarpus minutiflorus Cowan, Bol. Mus. Nac. Rio de Janeiro. ser. 2. Bot. 27: 3, t. 1, fig. b and
c. 1961, syn. nov. TYPE. Segadas-Vianna 4194, Brazil. Rio de Janeiro: Cabo Frio Co.,
Arraial do Cabo, Pontal Beach, Feb-Mar 1951, bud (holotype, US-2279054; isotype, R n.v.).

Branchlets and petioles pubescent with hairs 0.05 mm long, or densely pubes-
cent with hairs 0.2-0.3 mm. Leaves (sub)opposite, aromatic when crushed, pu-
bescent with spreading hairs to 0.3 mm, mainly at base, on principal nerves, and
on the margin, the indument often soft to the touch below but occasionally nearly
absent there. Racemes often several near tips of branchlets, to 1 cm wide; pedicels
1.5-3(-3.5) mm long, in fruit to 6 mm. Flowers 4-merous, rarely some 5-merous,
4.5-6 mm in diam.; petals 1.9-2.2(-2.5) x 1.3-1.5(-1.9) mm; filaments + subu-
late, the tip acute or acuminate, but never truncate; anthers versatile.
Type. Machado RB 75557, Brazil. Rio de Janeiro: Restinga da Tijuca, 5 Feb
1947, fl & fr (lectotype, RB (fl); isolectotypes, RB 2x, US n.v., WU n.v.).
Distribution. Brazil, Bahia and Espirito Santo (rare), Rio de Janeiro; locally
frequent in restinga woods on sandy soil. Flowering Feb-May(-Sep). Fig. 43C.
Specimens examined. BRAZIL. Bahia: Duarte 6622 fl (U). Espirito Santo: Souza RB 90086 fl (RB
2x). Rio de Janeiro: Almeida 1304 fl (RB 2x); Almeida & Laroche 1361 fr (RB 2x); Armond 87 fl (R
2x); Duarte 9872 fl/fr (RB 2x); Duarte (& Pereira) 4643 fl (K, M, U); Freire et al. R 71074 fl/fr (R 3x);
Gardner 830 fl (BM, K); Glaziou 3919 fl (C, K, P 2x); J. G. Kuhlmann RB 53634 fl (BM, F, MO,
NY), RB 102948 fr (RB 2x); Lutz 673 bud (US), 857 fr (R); Lutz & Cochran R 29275 fr (US); Machado
RB 75532 fr (RB 2x), RB 75537 fl (BM, F, MO, U), RB 75538 fr (U), RB 75549 fl & fr (RB 2x), RB
75552 fl (NY, U); Mello Filho 1207 fl (R 2x); Miers 3593 fl (K, P); Peckolt et al. R 71075 fl/fr (R 3x);
Pereira 546 fl (U); Restinga I 659 fl/fr (R 2x); Ribeiro R 71092 fl (R 3x, of doubtful origin); Riedel 35
(?) fl (LE); Rizzini RB 152807 fl (U); Schott Herb. Bras. 4458 fl (NY, W); Vidal III 1652 fl (R 3x).
Local name. Near Rio de Janeiro: jaborandi-gratido-da-restinga (Machado RB
75532, RB 75557).
The flowers of var. lealii are nearly always 4-merous and always stalked. In
these characters this variety differs from var. spicatus, which has usually 5-mer-
ous stalked or subsessile flowers. Variety lealii is typically a plant from the
restinga's of the Brazilian eastern coasts, whereas var. spicatus chiefly grows
more inland.
Some specimens are in their leaf shape rather similar to subsp. longeracemo-
sus. These specimens are densely pubescent below. Others resemble var. spi-
176 Flora Neotropica

catus in the indument of the leaves. On the same locality specimens were col-
lected with the leaves subglabrousbelow and others with densely pubescent
leaves. Intermediatespecimens are also known: Lutz 673 and J. G. Kuhlmann
RB 53634, for instance. The presence of hairs on disc and ovary is not correlated
with the indumentof the leaves, neither could other correlationswith the indu-
ment of the leaves be found. The habitat of hairy and subglabrousspecimens is
always the same too. Therefore in my opinion there is no reason to create a
separate taxon for subglabrousspecimens.
Cowan based Pilocarpus minutiflorus on two subglabrous collections from
Cabo Frio. The pedicels of the holotype are rather long: 3-3.5 mm; this is 0.5
mm longer than the range of P. spicatus var. lealii. In the paratypes however,
they are postflorally only 2.5 mm long. In other characters P. minutiflorusis
similar to specimens of var. lealii with subglabrousleaves; the petals of P. mi-
nutiflorusare ca. 2 mm long and therefore not differenteither.
The originaldescription of P. lealii: "Petala cum diametro mm 3 usque 3,5.
Petala ovatae ..." apparently is an error and should be read as "Flores cum
diametro . .. " (these measurements apparently were made without spreading
the petals). This can be observed from the photo in the protologueand also from
the paratypes. Hairy discs occur in subglabrousand in pubescent forms of var.
lealii, and the same applies to glabrous discs. The only difference of P. minuti-
florus with var. lealii, therefore, is the slightly longer pedicels in one of the
originalspecimens. I suppose that if Cowan had studied authenticmaterialof P.
lealii he would not have publishedhis P. minutiflorus.Now I have to reduce this
species to synonymy.
Machado did not designate a type of var. lealii, he only gave a type locality.
From that place he collected several specimens and identifiedthem as P. lealii
or P. longeracemosus. ApparentlyI did not see all the collections, because Al-
buquerque(1968)mentions some more specimens;Machadocould also have stud-
ied additionalcollections from other collectors. Because Machado did not des-
ignate a type, and since Albuquerque only made mention of a "cotypus":
Machado s.n., Restingada Tijuca, 27 Feb 1947(RB), Kaastra(1977)selected one
of the originalspecimens of Machadoin RB as the lectotype.

12b. Pilocarpusspicatus Saint-Hilairesubsp. longeracemosus(Martiusex Engler

in Martius)Kaastra, Acta Bot. Neerl. 26: 487. 1977. Fig. 56G-I.
Pilocarpus longeracemosusMartiusex Engler in Martius,Fl. bras. 12(2): 135. 1874;Engler in
Englerand Prantl,Nat. Pflanzenf.3(4): 158. 1896;Duval, RecherchesJaborandis58. 1905.
Pilocarpus longeracemosus Martius ex Engler in Martius var. breviusculus Rizzini, Leandra 4-
5: 13. 1974,syn. nov. TYPE.Ramalho204, Brazil. Bahia:Morrodo Chapeu,FazendaMaria
Vermelha, 15 May 1973,fl (holotype, RB-148266).
Branchlets and petioles beset with spreading hairs 0.3-0.4 mm. Leaves pu-
berulous above with spreadinghairs to 0.5 mm long, densely pubescent and soft
to the touch on the costa and below. Racemes to 1.5 cm wide, pubescent with
spreading hairs 0.1-0.3 mm long; pedicels 2-5 mm long. Flowers 6-7 mm in
diam., perfumed(Ramalho204) and much visited by bees (Harley 16737);petals
2.5-2.7 x 1.7-1.8 mm; filaments subulate, 2.2-2.8 mm long, anthers versatile;
disc 0.5-0.8 mm high and 2-2.5 mm in diam.
Type. Blanchet 3263, Brazil. Bahia: IgaljaVelha, 1841,fl (holotype, BR-Herb.
Martius;isotypes, F ex G, G-Herb. Delessert, G-Herb. Moricand,photo 26426
made by F, NY; GOET-Herb.Luca, K, LE, P 3x, U, W).
Pilocarpus 177

Distribution.Brazil, confinedto Bahia; alt. up to 1000m. FloweringJan-May.

Fig. 43C.
Specimensexamined.BRAZIL.Bahia:Serrado Rio de Contas,8 km N of Rio de Contas,41?47'W,
13?33'S,Harley et al. 15249bud (K 6x, U); Serra do CurralFeio, 3 km NW of Lagoinha,41016'W,
10?27'S,Harley et al. 16737fl & fr (K 9x, U). Exact locality unknown:Blanchet339 fl (G, P), 6025
(?) fl (G 2x); Herb. Glaziou s.n. leaf (F ex P); Glocker 339 fl (BM); Comm. Martius s.n. (anno 1862)
fl (L). State unknown: Herb. Martius s.n. fl (BR 3x); Mooy (?) (probably from Austria) s.n. (anno
1817) fl (BR).

Local name. Jaborandi(Ramalho204).

This subspecies is distinguishedby its large flowers with subulatefilamentsand
versatile anthers, and by the long hairs. It is most near to subsp. aracatensis.
The 8-10 cm long raceme of some of the syntypes equals that of Pilocarpus
longeracemosus var. breviusculusRizzini in length. Because the type of the latter
variety shows no differences with subsp. longeracemosus, I have reduced var.
breviusculus to synonymy.

12c. Pilocarpus spicatus Saint-Hilaire subsp. aracatensis Kaastra, Acta Bot.

Neerl. 26: 487. 1977;Holmes, Pharm.J. Trans. ser. 4. 1: 501. 14 Dec
1895;idem, ibidem: 521, fig. D, Dl, D2. 21 Dec 1895;idem, ibidem:
540. 28 Dec 1895, sine nom. Fig. 56K-L.
Tree, minutelypubescent with hairs to 0.1 mm long, the leaves mainly at base
and on costa, or subglabrous.Racemes to 20 cm long and 1-1.5 cm wide; pedicels
4-5 mm long. Flowers 5-merous, 4.5-5.5 mm in diam.; petals 2-2.3 x 1.2-1.5
mm; filamentssubulate, anthersversatile.
Type. Guedes 553, Brazil. Ceara: Serra da Aratanha, 27 Mar 1958, fl & fr
(holotype, US-2484399;isotypes, NY, RB-114215(s.n., dated 26 Mar 1958,det.
Ducke 1957(sic!), all ex IAN)).
Distribution. Brazil, Pernambuco,Ceara, and Bahia (not mapped). Collected
in flower in Mar-Jul Fig. 43C.
Specimensexamined.BRAZIL.Ceara:Serrada Aratanha,AllemdoR 71071fl (R 6x). Exact locality
unknown: Allemdo 276 fl (P, R 5x), R 71077 fl (R 2x); Glaziou 11855 fl (C, K, P); Herb. Saldanha
7582 fl (R). Pernambuco: Serra do Araripina, Andrade Lima & Magalhdes 52-1069 bud & fl/fr (R).
Bahia: Itiruqu-Maracas,
Pinheiro 1872 fl (U); 12 km from Valen9a, nr. Guaiubi,Santos 2646 fl (U).
Northern Brazil, of commercial origin: Comm. Bragg & Co. s.n. (Jun 1896, bale 312) st (PHA);
Comm. Holmes s.n. (1 Jan 1895) st (BM), s.n. (Nov 1895) st (K 2x); Herb. Holmes s.n. st (PHA 2x).

Local name and use. Jaborandi-de-madeira (Guedes553). Leaves importedinto

Europe (Liverpool) in 1895 and possibly some more years afterwardsas jabo-
randi. Chemicalexaminationshowever, proved that they probablydo not contain
pilocarpine, the main active alkaloid (Holmes, 1895a),see also PHYTOCHEM-
ISTRY.
Geiger (1897:414) supposed synonymy of this subspecies with Pilocarpus spi-
catus (subsp. spicatus). The floweringspecimens available now, however, show
clear differences.
The subspecies was discovered by Holmes in 1895,who obtained some leaves
and fruits out of commerce. He called it "aracatijaborandi," after the harborin
Ceara from which it was shipped.
Glaziou 11855 in P is labeled: "Rio de Janeiro, Morro da Viuva, 2 Janvier
1870." This is a similarannotationas at Glaziou3919 in C. Accordingto Glaziou
(1905), the two collections were made on the Morroda Viuva, too. In my opinion
however, the label on the sheet of Glaziou 11855in P has been wronglyattached
178 Flora Neotropica

to a collection communicated by Allemao, cf. Wurdack (1970). Allemao collected

specimens of the same taxon, as appears from his collection 276 in P and R.

13. Pilocarpus pauciflorus Saint-Hilaire, Bull. Sci. Soc. Philom. Paris (1823): 131.
Sep 1823; Mem. Mus. Hist. Nat. 10: 145. 21 Apr-28 Jun 1823, nom.
nud.; Mem. Mus. Hist. Nat. 10: 361. after 6 Nov, 1823; de Candolle,
Prodr. 1: 728. 1824; Sprengel in Syst. veg. ed. 16. 1: 954. 1824/1825;
Saint-Hilaire, Fl. Bras. mer. 1: 83, t. 17. 11 Jun 1825; Hist. rem. Br6s.
Par. 1: 147. 4 Nov 1825; Spach, Hist. nat. veg. 2: 345. 1834; Dietrich,
Syn. pl. 2: 1017. 1840 (all citations so far, except for Sprengel, as "P.
pauciflora"); Engler in Martius, Fl. bras. 12(2): 134. 1874; Albuquer-
que, Anais Acad. Brasil. Ci. 40: 507. 1968, pro parte; Cowan and Smith
in Reitz, Fl. Ilustr. Cat. 1 (RUTA) 30, t. 7. 1973. Fig. 57.
Pilocarpusfuminensis Casarettoex Englerin Martius,Fl. bras. 12(2): 134. 1874,pro syn. Based
on: Casaretto 1479, Brazil. Rio de Janeiro:Nr. Rio de Janeiro,fr (G-Herb.DC-Delessertex
TO, 2x).
PilocarpusbreviracemosusCowan, Sellowia 12: 86. 1960;Cowan and Smith in Reitz, Fl. Ilustr.
Cat. 1 (RUTA) 30. 1973,pro syn. TYPE. Reitz & Klein 4685, Brazil. Santa Catarina:San
Franciscodo Sul, Garuva,MinaVelha, 24 Aug 1957,fl & fr (holotype,US-2278675;isotypes,
G, NY, S, UC, Z, all ex HBR).
Pilocarpus organensis Occhioni & Rizzini, Leandra 4-5: 101, t. 2. 1974, syn. nov. TYPE.
Occhioni5970, Brazil. Rio de Janeiro:Serrados Orgaos,nr. Limoeiro, May 1974,fl (holo-
type, RFA-15132,n.v.); Occhioni6088, Brazil. Rio de Janeiro:Serrados Orgaos,nr. Guapi-
Mirim,21 Aug 1974,fr (paratype,RFA-15629,n.v.).
Pilocarpus pauciflorus auct. non Knox, Cat. 86, 1857:Urban, Bot. Jahrb.Syst. 21: 553. 1896.
Shrub or small tree 1-6 m tall with trunk reported as 7 cm in diam. (once
reported from Sao Paulo as a tall tree); dead bark very thin, gray-purplish-brown
with minute elongate-reticulate cracks, falling off in minute chips, with green
boundary layer and a living bark 2.5 mm thick (according to Lindeman); branch-
lets 2-4 mm thick, grayish-green-brown, shining when young, pubescent with
hairs 0.05-0.1 mm long, becoming glabrous in age; perules of terminal buds tri-
angular, strigillose with yellow-tawny hairs ca. 0.3 mm long. Leaves alternate or
subopposite, crowded or not at the apex of branchlets, simple or indistinctly
1-foliolate; petiole semiterete, canaliculate towards base by erect wings, distally
sometimes turgid-geniculate, 4-25(-40) mm long and I mm thick, glabrous or
pubescent with hairs 0.05-0.1 mm; the wings formed by the decurrent base of
the blade, up to 0.2-0.3 mm broad; blade (narrowly) elliptic to (narrowly) ob-
ovate, 5.5-16 cm long (-30 cm near ends of branchlets) and 1.8-5.5 cm broad
(-9 cm near ends of branchlets), long-attenuate or narrowly cuneate towards
base, decurrent along petiole, obtuse or acuminate at apex, the very tip retuse,
emarginate, or entire, margin revolute; the blade chartaceous or subcoriaceous,
grayish-green, shining, glabrous or pubescent with spreading hairs 0.05-0.1(-0.2)
mm at base and midvein below, venation brochidodromous, midvein usually plane
and longitudinally wrinkled above, principal veins prominulous. Racemes I or 2
per branch, subterminal, erect, 5-40 x 1.5-2.5(-3) cm, in fruit to 45 cm long,
many-flowered, developing acri- and basipetally; rachis 1-1.5 mm thick, glabrous
or puberulous with hairs 0.1 mm long; bracts and bractlets depressedly triangular,
to 0.7 mm long, subglabrous; pedicels inserted at 70-90?, 2.5-9(-11) x 0.5-1 mm,
indument like that of rachis; bractlets (2-)3-4, alternate or subopposite at variable
height. Flowers 7-9 mm in diam.; calyx 5-lobed; lobes separate, depressedly
ovate or suborbicular, 0.4-0.7 x 0.9-1 mm, rounded, thinly coriaceous, glabrous;
petals cochlear to subvalvate, 3-3.9 x 1.7-2.4 mm, at tip inflexed through 0.7
mm, slightly carinate above, coriaceous, yellowish-green, shining underneath,
Pilocarpus 179

FIG. 57. Pilocarpus pauciflorus. A, habit (Klein & Bresolin 9469, US). B, flower with B 1, abaxial
side of stamen (Klein & Bresolin 9469, US). C, mericarp (Dusen 16453, MO).
180 Flora Neotropica

glabrous but beneath in bud often beset with hairs 0.05 mm long, venation acti-
nodromous, median nerve thicker; filaments truncate, flattened, 2-3 x 0.3-0.5
mm, yellowish-green; anthers ovoid, heart-shaped or suborbicular, recurved,
(1-)1.2-1.5 x (0.8-)1-1.5 mm, with a dorsal gland 0.2-0.5 mm; disc 0.5-1 mm
high and 2-3 mm in diam., irregularly10-plicate,glabrousto mostly strigose with
yellowish-tawny, hyaline hairs 0.1-0.4 mm; carpels 0.7-0.9 mm high, protruding
0.3-0.6 mm beyond the disc, indumentlike that of the disc, with internalglands;
ovule 1 per carpel; style at anthesis obsolete to 0.1 mm long and 0.7 mm thick,
after anthesis to 0.2 mm long; stigma subsessile, capitate, 0.2-0.5 x 0.5-0.7 mm.
Mericarpsellipsoidalor obovoid, (6-)7-15 x (5-)7-14 mm, dorso-apicallyround-
ed, blunt or rounded at very apex, with glands to 0.5 mm, glabrousbut young
and sterile ones usually sparsely strigillose, dehiscent to 3-12?below tip; seed I
per mericarp,ellipsoidal, 7.7-12 x 4.3-8.5 x 3.8-4 mm, at apex curved or not,
ventral axis straight,slightlykeeled on the back, testa externallyflatly colliculate
with angularinterspaces0.05-0.1 mm; hilumin the upperpart of the ventralaxis,
ca. 2.5-3 x 1.2-1.3 mm; embryo 1 or 2, if 1: providedwith 2 or 3 cotyledons (or
the third cotyledon belonging to a second, incomplete embryo?), if 2: provided
with 4 or 2 cotyledons, dirty greenish, (one of them) large, with or without an
internal, small, thin cotyledon and an external, large cotyledon ca. 8 x 4.6 x 3
mm with ears 0.4-1 mm long, radicle 1.1-1.2 mm long, projectingor enclosed,
plumule 0.3-0.7 mm long; radicle and especially the plumule (rather densely)
strigose with tawny, hyaline hairs 0.05-0.4 mm, or subglabrous;the second em-
bryo, if any, smaller, with an external cotyledon ca. 6.5 x 3.5 x 1.5 mm, gla-
brous.
Type. Saint-Hilaire 1734(o?, Brazil. Santa Catarina: Nr. Itapocoroi, Mar 1820,
fl (lectotype, P ("forets vierges"); isolectotypes, MPU, P ("sylvis primaevis")).
Distribution.Brazil, Bahia (not mapped),Rio de Janeiro, and Sao Paulo (rare),
Parana,and Santa Catarina.Forests and cerrado;alt. 10-850 m. FloweringMar-
Oct. Fig. 43C.
Specimens examined. BRAZIL. Bahia:Mattos Silva et al. 178 fl (U). Sao Paulo:EdwallCGGSP
3408 fl (SP); Gehrt CGGSP 39556 fl (SP); W. Hoehne SP 30868 fl/fr (F, NY 2x, P, SP, US); M.
Kuhlmann 872 fl/fr (SP); Mello s.n. (23 Jul 1872) fl (S 3x). Rio de Janeiro: Casaretto 1862 fr (TO);
Glaziou 1070 bud (BR, C, P 2x, PHA); Occhioni & Rizzini RB 168199 fl & fr (RB 2x). Parana: Dusen
10468 bud (GH, S), 16453 fl & fr (F, GH, K, MICH, MO, NY, S), 16745 fl (S); Gomes & Mattos
Filho 253 fl & fr (U); Hatschbach 3149 fl (US), 14428 bud (K, P, US), 16510 fl (WIS), 19490 fl (C, L,
NY, UC), 22082 fl & fr (F, K, MO, UC); Lindeman& de Haas 2728 bud & fr (U). Santa Catarina:
Hetschko 225 IV fr (R); Klein 10142 fl (US); Klein & Bresolin 9469 fl (US 2x); Reitz & Klein 5914
bud (UC, US); Saint-Hilaire 1735(0) fl (P), s.n. leaf (F ex P); Ule 349 fl (HBG).

Local name. Santa Catarina:pitaguara-do-branco(Reitz & Klein 5914).

This species sometimes superficiallyresembles Pilocarpus spicatus var. spi-
catus. In that case P. spicatus has strigose ovaries. The diameterof the flowers
of P. spicatus is smaller, however.
Pilocarpuspauciflorusoccasioned much brain-rackingto me. At firstI intended
to place it as a variety of P. spicatus. This was due to a misinterpretationof the
descriptionby Saint-Hilaire"pedicelli 2-3 longi." The measurement"linea" was
omitted here, not "mm"! However, when I studied the type in P, I realized that
the present species is identicalwith P. breviracemosus.Cowan, in his "Rutaceae
of Santa Catarina"(1960) did not mention P. pauciflorus, but he is of the same
opinion now, see Cowan and Smith (1973).
The "type" of Pilocarpusfluminensis has poorly developed fruits with meri-
carps ca. 6 x 5 mm. The sterile mericarpsare strigose, the pedicels 7 mm long,
and the persistent petals ca. 2.4 mm long when dry. The racemes are short.
Acknowledgments 181

Although the mericarpsare small, the "type" does not differ from other speci-
mens of P. pauciflorus: the isotype of P. breviracemosus, Reitz & Klein 4685
(NY), also has some small fruits.
In specimens from outside Rio de Janeiro the racemes are 5-17 cm long, and
just after floweringup to 24 cm. The racemes of P. organensis, as far as I was
able to study them, are 30-35 cm long, in fruit40 cm (accordingto the protologue
still 5 cm longer). These long racemes, together with the large apical leaves (20-
30 x 5-9 cm according to the protologue), recently gave rise to the publication
of the new species P. organensis by Occhioni and Rizzini. However, the collec-
tion Glaziou 1070, from Mt. Corcovado near Rio de Janeiro, shows racemes with
flowers still in bud 22-35 cm long, and on the sheet in BR also one flowering
raceme 12 cm only. The leaves are up to 16 x 5.5 cm. In other charactersneither
Glaziou 1070 nor P. organensis differ from P. pauciflorus. Apparently these
collections from Rio de Janeiroare somewhat more luxuriant,which may be due
to the differentclimatologicalconditions. The same phenomenon occurs in Me-
trodorea nigra. Therefore I reduce P. organensis to synonymy under P. pauci-
florus.
Seeds are rarely found in herbariumspecimens.

Excluded from Pilocarpus

Pilocarpusamarus Blanco, Fl. Filip. (ed. 2) 540. 1845(as "P. amara"). =Lunasia
amara Blanco, Fl. Filip. 783. 1837, according to Merrill, Bur. Gov.
Lab. (Manila)27: 27. 1905.
Pilocarpus atropurpureus(Fischer ex Lemaire) Koch in Koch and Fintelman,
Wochenschr. GairtnereiPflanzenk. 2: 152, 419. 1959. =Metrodorea
nigra Saint-Hilaire(see this study, page 125).
Pilocarpus humboldtiiSprengelin Syst. veg. ed. 16. 4(2): 126. 1827,nom. illegit.
=Esenbeckia pilocarpoides Kunth (see this study, page 84).

ACKNOWLEDGMENTS
I wish to acknowledge Mr. L. Y. Th. Westra for spending many hours in
polishing the text and for valuable discussions which promoted this study. The
criticalreadingof the manuscriptby Dr. A. L. Stoffersis also greatlyappreciated.
I am indebtedto Dr. J. C. Lindemanfor informationabout Brazilianmattersand
for advice about ecological data; to Dr. F. A. Stafleu for nomenclaturalconsul-
tations; to Dr. J. L. van Eijk, Mr. W. F. van der Giesen, and Mr. J. K. J. Kuiper
for pharmaceuticaldata; to Mr. F. G. W. Schouten for classical information;to
Ing. P. N. H6weler for lending his maps; to Dr. E. Forero and Padre Lorenzo
Uribe-Uribefor providinga copy of a part of a rare book; to Dr. K. U. Kramer,
Dr. P. J. M. Maas, and Mr. W. G. Driehuis c.s. for their help in the loan of ca.
7500 specimens; to Mr. E. A. Mennega and Ing. B. J. ter Welle for technical
assistance; to Mr. A. J. Goslinga, Mrs. J. R. A. J. Hendriks-Holla,Mrs. Trix
Pelupessi, and Mrs. K. de Rooij-Stekelenburgfor the typing of the manuscript;
to the directorsand curatorsof the many herbariacited in the introduction,who
providedinformationand lent materialfor study;and to the directorsand curators
of the herbariaand botanicalgardens in Meise (BR), Geneva, Ghent, Kew, Lei-
den, London (British Museum, Natural History), and Paris for makingtheir fa-
cilities availableto me duringmy visit. To Mr. H. RijpkemaI express my sincere
thanks for the 55 drawingsand maps, which contributeto a better understanding
182 Flora Neotropica

of the subtribe and which make this monographmore pleasant. This work was
supportedin part by grant nr. R 85-126 by the Netherlands Organizationfor the
Advancementof Pure Research (Z. W. 0.).

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286-325.
Numerical List of Taxa 185

Troll, W. 1969. Die Infloreszenzen2(1). Gustav Fischer, Jena.

Ule, E. 1914. Bericht iiber den Verlaufder zweiten Expeditionin das Gebiet des Amazonenstromes
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Wurdack,J. J. 1970. Erroneousdata in Glazioucollections of Melastomataceae.Taxon. 19:911-913.

NUMERICAL LIST OF TAXA

1. Esenbeckia
subgenus I. Esenbeckia
sect Ia. Pachypetalae Engler in Martius
1. E. pentaphylla (Macfadyen) Grisebach
a. subsp. pentaphylla
b. subsp. belizensis (Lundell) Kaastra
c. subsp. australensis Kaastra
2. E. berlandieri Baillon ex Hemsley
a. subsp. berlandieri
b. subsp. litoralis (Donnell-Smith) Kaastra
c. subsp. acapulcensis (Rose) Kaastra
3. E. runyonii Morton
4. E. feddemae Kaastra
5. E. collina Brandegee
a. subsp. collina
b. subsp. conspecta Kaastra
6. E. irwiniana Kaastra
7. E. pumila Pohl
8. E. nesiotica Standley
9. E. grandiflora Martius
a- 1. subsp. and var. grandiflora
a-2. subsp. grandiflora var. intermedia (Martius ex Engler) Kaastra
b. subsp. brevipetiolata Kaastra
10. E. echinoidea Standley & Steyermark
11. E. macrantha Rose
12. E. flava Brandegee
13. E. hartmanii Robinson & Fernald
14. E. leiocarpa Engler
15. E. cornuta Engler
16. E. oligantha Kaastra
sect Ib. Esenbeckia
17. E. pilocarpoides Kunth
a. subsp. pilocarpoides
b. subsp. maurioides (Martius) Kaastra
18. E. amazonica Kaastra
19. E. alata (Karsten & Triana) Triana & Planchon
20. E. warscewiczii Engler
21. E. scrotiformis Kaastra
186 Flora Neotropica

subgenus II. Oppositifolia Kaastra

22. E. febrifuga (Saint-Hilaire)Jussieu ex Martius
23. E. densiflora (Chodat & Hassler) Hassler
24. E. hieronymi Engler
subgenusIII. LateriflorensKaastra
25. E. almawillia Kaastra
26. E. cowanii Kaastra
2. Metrodorea
1. M. maracasana Kaastra
2. M. flavida Krause
3. M. mollis Taubert
4. M. nigra Saint-Hilaire
5. M. stipularis Martius
3. Raulinoa
1. R. echinata Cowan
4. Pilocarpus
1. P. pennatifolius Lemaire
a. var. pennatifolius
b. var. pilosus Kaastra
2. P. demerarae Sandwith
3. P. grandiflorus Engler
4. P. jaborandi Holmes
5. P. trachylophus Holmes
6. P. microphyllus Stapf ex Wardleworth
7. P. racemosus Vahl
a-1. subsp. and var. racemosus
a-2. subsp. racemosus var. yucatanus Kaastra
b. subsp. viridulusKaastra
8. P. goudotianus Tulasne
a-1. subsp. and var. goudotianus
a-2. subsp. goudotianusvar. mollis (Cuatrecasas)Kaastra
b. subsp. heterochromus Kaastra
9. P. giganteus Engler
10. P. peruvianus(Macbride)Kaastra
11. P. riedelianus Engler
12. P. spicatus Saint-Hilaire
a-1. subsp. and var. spicatus
a-2. subsp. spicatus var. lealii (Machado)Kaastra
b. subsp. longeracemosus(Martiusex Engler)Kaastra
c. subsp. aracatensis Kaastra
13. P. pauciflorus Saint-Hilaire

LIST OF EXSICCATAE
Ackerman, s.n. (Herb. Martius 1065) (1-9a2).
Agostini, G., 1962/70(4-8a1).
Agostini, G. et al., 1750(1-17a).
Aguilar, J. I., 95(1-10).
Alboff, N. M., LP 38517(4-la).
Alexander (Prior), R. C., s.n.(l-la).
Allemao e Cysneiros, F., 274(4-4); 275(4-12al);
276(4-12c); 277(2-3); 279(1-9al); 281(1-22); R
7107, R 71077(4-12c).
Allen, P. H., 982(1-2b).
Almeida, J. (et al.), 1304, 1361(4-12a2).
Alvarado, L., s.n.(Jan 1922)(4-8al).
Amaral, E. do, SP 35609(2-4).
Anderson, s.n.(West Indies), s.n.(Herb. For-
syth)(4-7a 1).
Anderson, W. R. (et al.), 7924(1-7); 8850(1-6);
9041(1-7); 9199, 9300(4-5); 36545, 36546(1-
16); 37017(4-5).
Andrade, N. de, EFSP 24(2-4).
Andrade Lima, D. de & M. Magalhaes, 52-
1069(4-12c); 52-1070(1-25); 52-1092(2-3).
Anisits, J. D., 2609(4-la).
Anonymus, 1(R 70805)(4-la); 29(SP 19838)(2-5);
62 (LPS 22286)(4-1a); 150(2-4); 885(anno
1890)(4-la); 1938(1-14); CGGSP 113(1-9al);
R 71068(1-24); s.n.(Ilheos 1855)(2-4); s.n.
(Pharmacie Centr. Paris 1880), s.n.(4-4), s.n.
(4-5).
Aquadilla, s.n.(Nov 1889)(4-7al).
Araujo, A., RB 168527(4-4).
Archer, W. A., 3355(1-19); 4905(4-1a).
Aristeguieta, L., 3969(4-8al); 5370(1-17a).
Aristeguieta, L. & G. Agostini, 4736(1-17a).
Armond, N., 87(4-12a2).
Arnoldo, M., 97, 231(4-8al).
Assis, V., 178(1-22).
Badini, J., 3182(4-la).
Herb. Bailey, L. H., 234(1-12).
List of Exsiccatae187
Balansa, B., 2514(4-la); 2516, 2518(1-22); 2518a,
2518b(1-23); 2519, 2519a, 3261, 3262(1-9al).
Barreto, M., 163, 6103(1-22); 6104(2-5); 8960,
10334, 10373(1-22).
Bartlett, H. H. (et al.), 10696, 10805(1-3); 16643(1-
2b).
Bauer (?) s.n. (4-la).
Belem, R. P., 1848(1-9b); 1929(1-7).
Bel6m, R. P. & G. M. Barroso, 4002(1-7).
Belem, R. P. & M. Magalhaes, 914(1-9b).
Belem, R. P. & R. S. Pinheiro, 2548(1-9al);
2815(1-14).
Bena, P., 981, 1699(4-7a1).
Benitez de Rojas, C. E., 1633(1-17a); 1704,
1808(4-8a1).
Berg, C. C. et al., 749(1-17a).
Berg, C. C. et al. (in Prance), 18590(1-17a);
18595, 19842(2-2).
Herb. Berlandier, J. L., 3125, s.n.(1-2a); ? s.n.
(Dec 1831)(1-3).
Bernardi, A. L., 6925(1-17a).
Comm. Bernhardi, Th., s.n.(anno 1886)(2-4).
Bertoni, 1403, 1651, 2233, 2869, 3657, 3698, 3721,
3739(4- la).
Black, G. A. & R. E. Schultes, 46-259(1-18).
Blanchet, J. S., 339(4-12b); 1716(4-11); 2378(2-4);
2779, 2780(1-7); 3263, 6025(?)(4-12b); s.n.(l-
7).
Blanco, C., 376(1-17a); 507(1-9al).
Bockermann, W., 79(1-22).
Boldingh, I., 6481(4-8a1).
Bonpland, A. J. A., 1226(4-la).
Bornmiiller, A., 725(4-la).
Bowie, J. & A. Cunningham, s.n.(2-4).
Brade, A. C. (et al.), 10643(1-9a2); 12560(4-12al);
1
17886(4-la).
Braga, R. & R. Lange, 78(2-4).
Comm. Bragg & Co., messrs. W. J., s.n. (Feb
1894)(4-5); s.n.(1895-6)(4-6); s.n.(Jun 1896)(4-
12c).
Brandegee, T. S., 89, s.n.(1890), s.n.(Sep 1891),
s.n.(17 Oct 1899), s.n.(1901), s.n.(Nov
1902)(1-12).
Bresolin, A., 558(1-24).
Brito (Venezuela), 14(1-17a).
Britton, N. L. (et al.), 577(1-la); 2268(1-la);
3693(1-la); 5166, 5460.
Britton, N. L., E. G. Britton & W. G. Freeman,
2161, 2716(1-17a).
Britton, N. L., P. Wilson & J. S. S. Le6n,
15264(4-7a1).
Broadway, W. E., 384, 3843, 4192, 4257(1-17a);
TRIN 5249(l-9a1).
Burchell, W. J., 1423(1-9a2); 2972(4-12al);
3582(2-4); 4935, 5045(1-22); 5051(2-4); 6190(1-
7); 6240(2-5).
Burkart, A. E., 18896(4-1a).
Cabrera, A. L. et al., 74(1-9al); 262(4-la).
Calder6n, S., 2462(1-2b).
Camargo, O., 61595(1-9al).
Camp, W. H., E 2976(1-20).
Campos Porto, P., RB 15255(2-5); RB 20481(2-4).
Capucho, P., 475(2-2).
Cardenas, M., 992(2-2).
Carter,A. et al., 3275, 3439, 3613(1-12).
Carter,A., I. L. Wiggins& W. R. Ernst, 501(1-
12).
Casaretto,J., 541(2-4); 1260(4-12al);1479(4-13);
1780(4-12al);1862(4-13).
Castellanos,A., 21831(1-7).
Ceroni, Z. et al., ICN 5908(4-la).
Chiang,50(1-2a), 114, 294(1-2a).
Clissold, J., s.n. (4-6).
Comm. Geogr. Geol. S. P., 92(2-5); 113(1-9al);
140(2-4).
Constantino, D., 124(4-9); RB 2549(2-4); RB
15261(4-9);RB 16394(4-12al).
Contreras,E., 6406, 7358, 7370(1-lb).
Conzatti, C., 4031, 4106(1-11).
Cowan,R. S. (et al.) (in Maguire),38757,38833(1-
26); 39049-A,39194(1-17a).
Comm. Craig,W., s.n.(13 May 1875)(4-4).
Croizat,L., 592, 592A(1-17a).
Crovetto,R. M., 5940(4-1a).
Criiger,H., s.n.(l-9al); s.n.(1-17a).
Crutchfield, J. et al., 5178, 5594(1-3); 5686,
5715(1-2a);6080(1-3);6100(1-2a).
Curran,H. M., 18(1-22);33, 682(4-la).
Curran,H. M. & M. Haman, 507, 540(4-8al).
Daveau, J., s.n.(27 Nov 1894),s.n.(1908)(4-la).
Dedecca, D. & C. Teixeira, IAC 8165(4-lb).
Delgado, E., 298(1-9a1).
Devoto, F. E., BAB 90738(1-9al);BAB 90810(1-
22); Herb. Direcc. Forestal 1495,Herb. Di-
recc. Forestal 1876(4-1a).
Dorantes,J. et al., 1250, 1325(1-2a).
Herb. Drake del Castillo, E., s.n.(Jun 1862)(4-
12al).
Duarte, A. P. (et al.), 310(4-12al); 1632(1-22);
1665(4-la);4643(4-12a2);5026(2-4);5562(4-
lb); 5896(4-9); 5987(4-11); 6622(4-12a2);
6841(1-9al);9423(1-7);9872(4-12a2).
Ducke, A., 1266(1-18);MG 1549, MG 2011(1-
9al); MG 15116,MG 15872(2-2);RB 8556(1-
14); RB 13621, RB 17734, RB 17735, RB
17736, RB 20474(2-2); RB 20479(4-12al); RB
23828(4-10).
Ducke, A. & D. de Andrade Lima, 48(1-9b);
119(4-11).
Ducke, A. & J. G. Kuhlmann,RB 16511(2-4).
DugandG., A., 536(1-lc).
Dusen, P., 22/33(1-22);9478(1-9al); 10468(4-13);
11198(1-22); 11201(4-la); 11259(1-22); 11788(1-
9al); 11817(4-la); 13215, 13788, 14946,
14964(1-9al); 16453(4-13); 16554(1-9al);
16745(4-13); 16818(1-22); 17798(1-9al); s.n.
(Herb. d'Alleizette, Feb 1904)(2-4); s.n.
(Herb. d'Alleizette, Mar 1907)(4-12al);s.n.
(1908-1912), s.n.(14 Jul 1911), s.n.(6 Oct
1911)(l-9al).
Duss, P. A., 1193, 2240, 4558(4-7al).
Dutra, J., R 71508(1-9al).
Duval, A. A. P., s.n.(4-5).
Edwall, G., CGGSP 3408(4-13); CGGSP 3409(1-
22); Exp. R.F. 4(4-1b); Exp. R.F. 60(2-4);
Inst. Biol. 19816(4-lb).
Egerton, E. A., s.n.(1881)(4-la).
188
Eggers, H. F. A. von, s.n.(10-12 Mar 1877)(4-
7al).
Egler, W. A. & H. S. Irwin, 45962(2-2).
Ehrhardt, W., 13(1-9al).
Ekman, E. L., 1965, 1966(4-la); 12473, 14017,
15222, 18036, 18228, H 3051, H 3564, H
7292, H 9488, H 15062, It. Regn. III 3751, It.
Regn. III 3757, It. Regn. III 4208, It. Regn.
III 4753(4-7al).
Emrick, G. M., 122(1-4).
Herb. Engelmann, 12084(4-4).
Enriquez, 0. G., 330(1-2a).
Falcao, J. I., 70(4-la); 93-51(1-22).
Fanshawe, D. B., 2065, 2811(4-2).
Faria, R., 11(1-14).
Feddema, C., 2699(1-4).
Ferris, R. S., 5699(1-8).
Fiebrig, K., 19(4-la); 518a(1-23); 765(1-9al);
5383(4-la); 6331(1-22).
Finlay, K., TRIN 3063(1-17a); TRIN 3064(1-9al,
1-17a).
Fisher, G. L., 3375(1-3); 35497(1-5a).
Flores, R. S., s.n.(1931)(1-2a); s.n.(Feb 1935)(4-
7a2).
Forero, E. et al., 6335(2-2).
Frazao, A., RB 7335(4-12al); RB 15258(2-5).
Freire, C. V. et al., R 71074(4-12a2).
Froes, R. L., 33, 19971, 19983(2-4).
Fr6es, R. L. (in Krukoff), 1710(1-17b); 9568,
9573(4-6); 11666(1-17b); 11668, 11762(4-6);
11797(1-17b); 12667(2-4).
Funck, N., 26(1-17a).
Funck, N. & L. J. Schlim, 675, 678bis(l-17a).
Galvao, R., s.n.(=Herb. Glaziou 11853)(1-22).
Garcia Barriga, H., 4312(1-19).
Gardner, G., 829(4-12al); 830(4-12a2); 3028,
3828(1-7); 4439(1-22); s.n.(4-12al).
Gaudichaud, Ch., 16(1-9al); 644(4-9); 752(1-9al);
795(2-4).
Gaumer, G. F., 752(1-2a); 24399(4-7a2); s.n.
(Comm. Godman Aug 1886)(1-2a).
Gehrt, A., SP 19855(1-22); SP 29999(1-14); SP
37019(1-22); SP 39556(4-13).
Geier, S., s.n.(19 Dec 1960)(4-la).
G(entle?), A. P. F., 193(1-lb).
Gentle, P. H., 2934, 3069, 4758, 4797, 5130,
8897(1- Ib).
Gentry, H. S., 2254, 6773, 7068(1-13); 11233,
12343(1-12); 14384(1-13).
Gevieski, A., 60(1-9al).
Gibert, 55(4-la).
Gifford, D. R. & S. Fonseca, G 275, G 323(2-3).
Gines, Bro., 3355(1-17a); 3577(1-9al); 3770(1-
17a).
Herb. Glaziou, A. F. M., 14, 41(4-12al); 675(1-
14); 678(1-9a2); 1070(4-13); 1390(1-9a2);
1391(2-4); 1392(1-14); 2525(1-9a2); 2526(2-4);
2958(4-12al); 3917(1-14); 3918(1-9a2); 3919(4-
12a2); 3959(4-1b); 4780(2-4); 8330(1-9al);
10452(2-3); 10458(1-9al); 10460(1-14);11850(2-
3); 11851, 11852(2-4); 11853(1-22); 11855(4-
12al; 4-12c); 11856(1-9al); 12526(2-5);
13417(4-6); 13651(2-4); 14587(4-9); 14588(2-
Flora Neotropica
3); 14589(2-4); 15887(2-3); 18172(1-9al);
18173, 18179(2-5); 18971(4-12al); 18973,
20246(1-22); 20798(1-7); 20798a(4-1a);
20799a(4-lb);s.n.(4-12b).
Glocker, E. F. von, 339(4-12b).
Goeze, E., s.n.(Apr 1876)(4-la).
Goldman,E. A., 246(1-13).
Gomes, J. C. & Mattos Filho, 253(4-13);1049(4-
la).
Goudot, J., 1(1-19);2(1-17a); s.n.(Apr 1844)(4-
8al); s.n.(1844)(1-19).
Greenway,P. J., 2576(4-la).
Grondona& Piccinini,3228(4-la).
Comm. Gubler,A., s.n.(Apr 1876)(4-4).
Guedes, T. N., 553(4-12c).
Guillemin,J. B. A., 1033(2-4).
Herb. Haenke, T., s.n.(PR 28)(1-2c).
Handro,O., s.n.(Jan 1944)(1-14).
Harley, R. M. et al., 15249(4-12b);15253(1-9al);
16737(4-12b); 17558(1-9al).
Harris, W., 5629, 5664, 7057, 8209, 9074, 10285,
11195, 12367(1-la).
Harrola, T., 1264(4-la).
Arr. Hart, J. H., TRIN 1324(1-17a); TRIN 1399,
TRIN 1411(1-9al);TRIN 3063(1-17a).
Hartman,C. V., 240(1-13).
Herb. Harvey, W. H., s.n.(l-9a2).
Hassler, E. (see also Rojas), 416(4-la); 512G(1-
9al); 892(4-la); 1439(1-22); 1579, 1695, 2107,
2107a(1-23);3046(4-la); 3401(1-23);4019(1-
9al); 4222(4-la); 4905(1-22); 5129, 6857(1-
9al); 8334(4-la); 8936, 8936a(l-22); 9337,
9415, 11282(4-1a); 11439(1-22); 11798,
11798a(4-1a); 12076(1-22); 12217(1-9al);
12240(4-la); 12358(1-23); 12951(4-la).
Hastings,J. R. & R. M. Turner,64-207,64-252(1-
12); 65-88, 65-174, 69-176(1-13).
Hatschbach, G. (et al.), 3149(4-13); 3795(1-9al);
7568(1-22); 7703, 8656(1-9al); 9400(4-la);
9411, 9446(1-9al); 10390, 11215(1-22);
11335(4-la); 12937(2-4); 12946(4-1a); 13238(1-
22); 13248-B, 13908(1-9al); 14428(4-13);
15740(1-22); 16583(4-la);16638(1-
16510(4-13);
22); 16935, 16948, 17006, 17055(2-4);17991,
18587(1-9al); 19077(1-22); 19152, 19303(4-
la); 19490(4-13); 21113(1-22); 21514(4-la);
21517(2-4); 22082(4-13); 26685(1-22); 26916(4-
la); 28378(1-9al); 28587(1-24).
Haught, O., 4480(4-8a2).
Heiner, A., 334(2-5);s.n. (1-22).
Helmreich(en?),s.n.(2-5).
Helmreichen, 17(2-4); 29(1-14); 64(2-5); s.n.(l-
14); s.n.(2-5).
Hemmendorf,E., 43(2-4); 169(4-12al).
Heringer,E. P., 8742(1-7).
l'Herminier, F. L., s.n.(4-7al).
Hetschko, 225(coll. IV)(4-13).
Hinton, G. B. et al., 10924(1-2c); 16178(4-7al).
Hoehne, F. C. (et al.), 318(2-4); Comm. Rondon
994(2-2); CPEF II 163(1-14); SP 251, SP
1335, SP 1492(1-9al); SP 1934(2-5); SP
2847(1-9al); SP 3456, SP 24859(2-4); SP
List of Exsiccatae
28424(1-9al);SP 29729(2-5);SP 31389(1-22):
SP 31430(1-9al);SP 39536(1-22).
Hoehne, W., FFO 2995(1-14);SP 30868(4-13);SP
30869(1-9al);SP 54145(1-22);SP 54147(2-4).
Comm. Holmes, E. M., s.n.(12 Jan 1875+ Feb
1875)(4-4); s.n.(8 Nov 1893)(4-6); s.n.
(1894)(4-4);s.n.(I Jan 1895)(4-12c);s.n.(Nov
1895)(4-4);s.n.(Nov 1895)(4-12c);s.n.(Apr
1896 + Feb 1897), s.n.(Mar 1897)(4-4);
idem(4-6);s.n.(4-4).
Herb.Holmes, E. M., s.n.(1898)(4-4); s.n.(1898)(4-
5); s.n.(1905)(4-6);s.n.(4-4); s.n.(4-12c).
Holton, I. F., 66("815")(1-19).
Homrich,M. H. et al., ICN 5164(4-la).
Horto Florestal (Rio de Janeiro), 6127(RB
43635)(4-9);RB 100583,RB 100585(1-14).
Hort. BO, 15, VII E 6(2-4); 149, s.n.(4-1a).
Hort. Butantan, 1492(1-9a1).
Hort. C, s.n.(Jan 1876)(2-4).
Hort. CGE, s.n.(30 Jan 1894)(4-4).
Hort. Cork, s.n.(Nov 1893)(4-la).
Hort. E, s.n.(Jun 1888)(4-4).
Hort. Hanbury(see Hort. GE).
Hort. GE, s.n.(1892), s.n.(13 May 1893),
s.n.(1894),s.n.(Jun1898),s.n.(20Oct 1920)(4-
la).
Hort. K, (1854), s.n.(1856)(2-4);s.n.(May 1857),
s.n.(1859)(1-9al);s.n.(Apr 1862)(2-4); s.n.(Jun
1888), s.n.(Nov 1891)(4-1a); s.n.(9 Jan
1896)(4-4);H 493/66(4-7al).
Hort. LE, s.n.(4-la).
Hort. LP, LPS 22287(4-la).
Hort. M, s.n.(Oct 1903),s.n.(4-la).
Hort. Orot(?),s.n.(Jan 1882)(4-la).
Hort. P, s.n.(Dec 1861), s.n.(P. simplex)(4-la).
Hort. Sch6nbrunn,s.n.(17 Dec 1907)(4-l1a).
Hort. STR, s.n.(Jun 1893)(4-1a).
Hort. TO, s.n.(1889)(4-la).
Hort. U, s.n.(4-la).
Hort. W, s.n.(1860)(4-1a); s.n.(1861)(2-4);
s.n.(1861)(4-la); s.n.(1862)(2-4); s.n.(Nov
1863)(4-1a); s.n.(1864)(2-4); s.n.(23 Nov
1868)(4-la);s.n.(22 May 1878/1875)(2-4).
Howell, J. T., 10235(1-2b).
Humboldt, F. W. H. A. von & A. J. A. Bon-
pland, 276(1-17a).
Hummel,C., 38(1-lb).
Hunger Filho, M., EFSP 301, s.n.(Jul 1928)(1-
14);s.n.(Jul 1928)(2-4);s.n.(Jul 1928)(2-5).
Inst. Malariologia,44(1-9al).
Irwin, H. S. et al., 5993, 8236, 9710, 10788,
10826a, 12175, 12459a, 14400(1-7); 15603,
18048(2-5); 23415(1-6); 26481, 26647(1-7);
31543(1-16); 34444(1-7).
Irwin, H. S. et al. (in Maguire), 55309(1-17a).
Janzen, D. M., s.n.(1-2a).
Jesus, J. A. de, 400(2-1).
Jimenez M., A., 1529(4-7b).
Jobert, C., s.n.(4-6).
Johnston,I. M., 4087(1-12).
Jones, M. E., 22355, 27433, 27433a(1-12).
de Jonghe, 479(1-9al).
Jonsson, I. G., 245a(l-9al).
189
Jorgensen,P., 163(2Sep 1909), 2588(May1919)
(4-la); 3975(5Oct or Nov 1928),3975(20Oct
or Nov 1928), 3975(19 Oct 1929)(1-22);
3975(Oct 1930)(1-23); 3975(Sep 1931),
3975(Oct 1931)(1-22); 3975(Oct 1931)(1-23);
3977(4-la);4793(F only)(l-23).
Karsten, H., s.n.(Oct 1872), s.n.(l-19); s.n.(4-
8al).
Kermes, 505(4-la).
Kerr, A., s.n.(Jan 1863)(1-9al).
Khek, E. J., s.n.(1896)(4-la).
Kiehl, J., SP 181(4-1b).
Killip, E. P. et al., 38300(1-19).
King, R. M., 760, 1483, 1562, 1652-A(l-2b);
4012(1-2a).
Klein, R. M. (et al.), 493, 600(1-9al);675(4-la);
761, 785, 1073(1-9al); 1212(1-24); 1256,
1564(1-9al); 2019(4-la); 6926, 6927(1-23);
7175(1-9al); 7378, 7445(4-la); 9373(1-24);
9469, 10142(4-13); IM 44(9 Apr 1951), IM
44(24 Apr 1951)(1-9al).
Klug, G., 1990, 3275, 3380(1-18); 3722(1-9al).
Koscinsky, M. de, 144(1-9al); 184(2-4);295(1-
22).
Krapovickas,A. et al., 13318(4-la);13697(1-9al);
14261(1-22);15068(4-1a);22959(1-9al).
Comm. Kreuzpointner,s.n.(1878)(4-4).
Krukoff,B. A. (see also Fr6es), 1667,5517(2-2).
Kuhlmann, J. G., 29(4-12al); 211(1-22);301(1-
9al); 347(4-11);6159(4-12al);6613(1-14);RB
15384(1-9a2);RB 16396(4-9);RB 46768(4-4);
RB 52281(4-1a); RB 53634(4-12a2); RB
73689(1-23);RB 102948(4-12a2).
Kuhlmann, M., 728(2-4); 755(1-9al); 782(1-22);
849(1-14); 872(4-13); 904(1-22); 1160(2-5);
1162(1-9al);4507(1-22);4508(1-14).
Kuntze, O., s.n.(Sep 1892)(4-1a).
Langsdorff, G. H. Baron von, s.n.(Dec 1825,
Itu)(1-22); s.n.(Ipanema)(l-9al).
Lasser, T. (et al.), 1483(1-9al); 2761(4-8al);
3027(1-17a); 3156, 4335(4-8al).
Earl Lathrop,5936, 5941(1-2b).
Lavalree, A., 16993(4-1a).
LBB (Lands Bosbeheer Suriname),3220(1-9al).
Leite, J. E., 567(1-9al);2173(4-la);2582(1-9al).
Le6n, J. S. S. (et al.), 11412,12253,14781,19176,
19336, 19645(4-7al).
Liebmann, F. M., 4198, 6414, 6614, 15005, 15015,
s.n.(Aug 1841), s.n.(l-2a).
Lillieskold,W., s.n.(4-la).
Lillo, M., 10593(1-22).
Lindeman, J. C. & J. H. de Haas, 577(2-4); 650,
650a, 710, 799(4-la); 916, 1480(1-22); 1482(4-
la); 1525(2-4);1526(4-1a);1569(1-22);1699(2-
4); 1705(1-22);2378(1-9al); 2728(4-13); 2841(4-
la); 4250, 4351, 5377(1-22); 5454(4-la);
5518(1-22); 5524(2-4).
Lindeman,J. C., A. L. Stoffers et al., 180(4-6);
493(1-9al).
Lindeman, J. C. et al., ICN 8925(4-1a);ICN
21599(1-9al).
Lindeman, C. A. M., It. Regn. I 35891/2,It. Regn.
I A3589(4-la).
190
Lofgren,A. (et al.), CGGSP34(2-4);CGGSP420,
CGGSP 496, CGGSP 623(1-22); CGGSP
668(4- b); CGGSP946(2-4);CGGSP 1342(2-
5); CGGSP 1363(1-9al); CGGSP 1390,
CGGSP 2729, CGGSP 2740(2-4); CGGSP
4178(4-9); CGGSP 4390(4-12al); CGGSP
4416(1-14);CGGSP4417(1-9al).
Lourteig,A., 1940(4-la).
Herb. LPS, 22278(1-22).
Luna, A., 124,463, 777(4-7al).
Lund, P. W., 277, s.n.(1833)(4-12al);s.n.(Feb
1834, =Herb. Warming2462)(1-22,2-4).
Lundell, C. L. & A. A. Lundell, 7532, 7831(1-
2a); 7861(4-7a2);7983(1-2a).
Luschnath, B., s.n.(Feb 1833)(1-9a2);s.n.(Nov
1833)(l-9al); s.n.(Rio, 1833)(2-4);s.n.(Rio,
1833), s.n.(May 1834)(4-12al); s.n.(2-4);
s.n.(4-12al).
Lutz, B. M. J., 640(4-12al); 673, 857(4-12a2);
1094, 1423(1-9al).
Lutz, B. M. J. & D. Cochran,R 29275(4-12a2).
Lutzelburg,Ph. von, 75, 975(4-5);1912/284,1912/
295(1-7); 13058(2-4).
Lynch, R. I., s.n.(Oct 1894),s.n.(1904),s.n.(4-4).
Llamas, A. de, 769(4-la).
Macedo, A., 384(4-5);4652(1-7).
Macfadyen,J., s.n.(Dec 1843),s.n.(l-la).
Machado,0. X. de Brito, 285(4-12al);RB 75532,
RB 75537, RB 75538(4-12a2);RB 75539(4-
12al); RB 75549, RB 75552, RB 75557(4-
12a2).
McLean, R. C., R 79(2-4).
McVaugh,R., 23032(1-2c).
McVaugh,R. & W. N. Koelz, 1672(1-2c).
Herb. Magnus, P. W., s.n.(20 Mar 1867)(2-4);
s.n.(20 Feb 1870),s.n.(10 Nov 1892)(4-la).
Maguire,B., 35808(1-9al).
Malme,G. 0. A., It. Regn. I/222(1-9al);It. Regn.
1/850, It. Regn. 1/850-b(4-1a);It. Regn. II
867(1-9a1).
Maltby, F. S., 82(4-7al).
Manning,W. E. & M. S. Manning,53404(1-3).
Manso, A. L. P. da Silva, 23(1-9al); 245(2-5);
246(4-1b); 247, 248(1-9al); s.n.(1835)(2-5);
s.n.(184.)(1-22);(in Riedel) s.n.(1835)(1-14).
March,W., s.n.(1888)(1-la).
Marshall,R. C., TRIN 12248(1-17a).
Martin,J., s.n.(4-7al).
Martius, K. F. P. von, 472, 614(1-9al); 1821(1-
22); Coll. Lign. 12(1-9al); s.n.(Chapadado
Parana, Sep)(1-7); s.n.(l-9al); s.n.(1-14);
s.n.(1-22).
Comm. Martius,K. F. P. von, 1 (anno 1841)(2-
4); s.n.(anno 1841)(4-12al);s.n.(anno 1862)(4-
12b).
Herb. Martius, K. F. P. von, 79(2-4); 181(2-5);
464(1-9al, 1-9a2); 1065(1-9a2);1069(1-14);
s.n.(4-12b).
Marufiak,V., 380(4-1a).
Mason Jr., C. T. & W. R. Brewer, 1731(1-13).
Mason, H. L., 1837(4-7al);1878(1-12).
Matthews,A., 3045(1-20).
Mattos Silva, L. A. et al., 178(4-13).
Matuda,E., 596(1-2b).
Flora Neotropica
Meaca, 31/138(1-9a1).
Mello, J. C. de, s.n.(23 Jul 1872)(4-13).
Mello Filho, L. E. de, 1107(1-9a2);1207(4-12a2).
Mennega, E. A., 66-1774(1-22).
Meyer, T., 1093, 5306(4-la); 5350(1-22);5478(4-
la); 5489(1-9a1).
Miers, J., 3333x(1-9a2);3370(2-4);3521(4-12al);
3593(4-12a2);4485, s.n.(Dec 1837)(2-4);s.n.
(1-9a2);s.n.(2-4); s.n.(4-12al).
Mikan,J. C., s.n.(2-4).
Milano, V. A. & J. M. Buceta, BAB 90812(1-
9al).
Miller, 0. 0. & J. R. Johnston,224(1-17a).
Millspaugh,C. F., 1475(1-2a).
Miranda,Faust., 5487(1-2a);8199(1-2b).
Mocquerys, A., s.n.(1893-1894)(1-17a); s.n.
(1893-1894)(4-8al).
MolinaR., A., 168, 181, 502, 4017, 7484, 10079,
14338(1-2b).
Molina R., A. & A. R. Molina, 25769(1-2b).
Monteiro, S. V., RB 130242(4-12al).
Montes, J. E., 72, 429, 780, 940, 2115, 2196, 2339,
2486, 4136, 14773, 15752(4-la).
Moore Jr., H. E. & C. W. Wood Jr., 3848(1-3).
Moore, Th., s.n.(Dec 1852)(2-4).
Mooy (?) (probablyfrom Austria, 1817)s.n.(4-
12b).
Moraes, J. C. de, EAN 2165(1-9b).
Moran, R. V., 3806, 3881, 6899, 9344, 11774,
11793(1-12).
Morel, I., 3486, 3532, 4262(4-la).
Morin, Cl. V. P., 510(4-la); 716(1-23).
Morong, Th., 635, s.n.(4-la).
Morton, C. V. (et al.), 3434, 10531, 10781, 10821(4-
7al).
Mosen, Hj., 1184(1-22); 1185(2-5); 2444(2-4):
2445(1-9a2); 2804, 3165(2-4); 3954(4-1b);
3968, 4059(1-9al); 4060(2-5); 4061(2-4).
Muiller,C. H., 2085, 2980(1-3).
Muller, 164(anno1880)(1-9al);s.n.(1-22).
Muniez, A., BAB 90811(4-1a).
Mutis, J. C., 4033(1-19).
Nadeaud, J., s.n.(30 May 1862)(1-9a2); s.n.(7
Nov 1862)(2-4); s.n.(1-9al).
Nelson, E. W., 1831(1-11); 4307(4-7a1).
Nelson, E. W. & E. A. Goldman,7258(1-12).
Netto, L. do S. M., 117(2-4).
Herb. Neyraut, E. J., s.n.(5 May 1930)(4-la).
Novaes, J. de C., 517(1-14); 521(2-4); 740
(=CGGSP 3305)(1-9al);CGGSP3216(1-14);
CGGSP 3799(2-5); SP 1950(1-9al); SP 2032(1-
22).
Nunes, G. M., EFCB 17, SP 20923(1-14).
Herb. Nyst, P., Anonymus 12, s.n.(17 Jan), s.n.
(4-7a1).
Occhioni, P., 324, 325(4-9).
Occhioni, P. & C. T. Rizzini, RB 168199(4-13).
Olboze (?), 486(1-9a2).
Oldenburger,F. H. F. et al., ON 508, ON 534(2-
2).
Oliveira, E. P. de, RB 82176(4-12al).
d'Orbigny,A. C. V., 764(1-25).
Orsted, 6(1-2b).
Ortega, J. G., 1212, 4204, 4621, 6348, 6398(1-13).
List of Exsiccatae 191

Osten, C., 8120, 8815(4-la). Herb. Richard, 138(2-4).

Otto, E., 1015(1-17a). Riedel, L., 3(2-5); 35(?)(1-14;4-12a2); 54(2-4);
Pabst, G. F. J., 5077(1-7); 8490(1-9b); 8545(2-1). 78(4-12al); 84(1-9al); 126(2-5); 162, 169(4-
Pacifico, V., CEFSP 280(2-4). 12al); 324(1-22);437, 438(4-11);466(4-12al);
Palmer, E., 175(1-2c); 181(1-lb); 477(1-3); 497(1- 469(2-4); 470(4-9); 1394, 1439, 1851(1-22);
2a); 514(4-7al); 1801(1-13). 2511(1-7);s.n.(1829)(4-12al);s.n.(1836)(2-4);
Peckolt, 0. et al., R 71075(4-12a2). s.n.(1836)(4-12al); s.n.(1-9al); s.n.(l-9a2);
Pedersen, T. M., 4210(1-23); 4213(4-la); 5137(1- s.n.(1-14);s.n.(1-22);s.n.(Hort. LE), s.n.(2-
23); 6039(4-la); 9379(1-22). 4); s.n.(2-5); s.n.(2-9); s.n.(4-11); s.n.(4-
Pereira, E. (et al.), 546(4-12a2); 686, 737(4-12al); 12al).
3968(1-9al); 4005(4-9); 4217(2-4); 4567(4- Riedel, L. & G. H. Baronvon Langsdorff,857(1-
12al); 4752(1-7); 7097(1-9al); 7101(4-12al); 9al).
Rizzini, C. T., RB 152807(4-12a2).
7255(4-9); 7774(1-22); 9003(1-7); 9601(1-9b);
9656(2-1). RN (anno 1892), 124(4-la).
Herb. Persoon, C. H., 23(4-12al). Robson, W., s.n.(4 Jun 1908)(4-7al).
Peters, R. E. K., 94(1-12). Rodrigues,W. & D. Wilson, 3426(2-2).
Pickel, B., 5450(2-4); 5451(4-1b); 5551(2-5). Rodriguez, F. M., 293(4-la); 433, 436(1-22);
Pinheiro, R. S., 1251(2-4); 1865(2-1); 1872(4-12c). 495(1-9al).
Pires, J. M., 58085(1-9al). Comm. Roemer, J. J. & J. A. Schultes, s.n.(4-
Pires, J. M. et al., 9496(2-5). 7al).
Pissis, D., 12(1-9a1). Rohrer,F., s.n.(16 Jul 1909)(4-1a).
Pittier, H. (et al.), 34(1-2b); 113(1-10); 5799, Rojas, T., 1579(1-23);8162(4-la); 12002(1-22).
6377(1-17a);8098(1-2b);8961, 10823(1-17a); Rojas, T. (in E. Hassler), 10539a(4-1a);10659(1-
13102(4-8al); 15435(1-9al); 15554(1-17a); CR 23).
2777(1-2b). RomeroC., R., 297, 9062(1-lc).
Plaumann,F., 117(4-1a). Rose, J. N. (et al.), 13609, 14735(1-13);16345(1-
Plee, A., 652(1-17a). 12); 22197(1-20).
Comm. Poehl, A., s.n.(4-4). Rosendahl, H. D., s.n.(29 May 1900),s.n.(May
Pohl, J. E., Herb. Bras. 27(2-4);Herb. Bras. 739, 1903)(4-1a).
750, 750 delta, s.n.(1839)(1-7);s.n.(2-5). Ross, H., s.n.(Oct 1902)(4-la).
Ponthieu,de, s.n.(misit Banksanno 1772)(4-7al). Rothkugel& F. E. Devoto, BAB 54371(1-22).
Poveda, L. J., 218(1-2b);836(4-7b). Runyon, R., 177, 2173, 3084, s.n.(6 Jul 1930),
Prance, G. T. et al., 8339, 8344, 18225, 18595(2- s.n.(1943)(1-3).
2); 18946(1-7); 19842, 58694, 58975(2-2); Rusby, H. H., 2617, 2663(2-2).
59058(1-9a 1). Rusby, H. H. & R. W. Squires,s.n.(1896)(1-9al).
Prata,G. da Rocha, 27(4-la). Ryan, J., s.n.(4-7al).
Pringle, C. G., s.n.(ll Oct 1890)(1-3). Ryan, J. & Lamb., s.n.(4-7al).
Pulle, A., 3053(4-la). Rzedowski, J., 5688(1-3);26572(1-12).
Pulle, A. & B. M. J. Lutz (in Lutz), 1094(1-9al). Sagra, R. de la, 820, s.n.(4-7al).
Purdie, W., 64(1-17a); 65(1-9al; 1-17a); s.n., s.n. Saint-Hilaire,A. F. C. P. de, 72B, 274(4-12al);
(1-la); s.n.(l-9al); s.n.(l-17a). 669(2-4); 717(1-22); 1734(o, 1735(0), s.n.(4-
Purpus, C. A., 4196(1-11); 6161(1-2a); 7140(1-5a); 13).
8419, 8428, 8651, 8895(1-2a). Herb. Saldanha, J. de, 7582(4-12c);7583(2-3);
Quir6s C., M., 1092, 1458(4-7b). 7584(1-9a1).
Ramalho, F. B., 204(4-12b). Saleman,R., G 6169(4-7al);G 6170(4-la).
Rambo, B., 413, 29104, 30942, 41841, 42012(1- Sampaio,A. J. de, 1584(4-12al);3909(4-la).
9al); 42102, 42376(4-la); 42733(1-9al); 42891, Sandeman,Chr., 4729, 4780(1-22).
43449, 47204(4-la). Santos, T. S. dos, 1158(2-1);1522(4-3);2646(4-
Herb. Raphelis, 1075(4-la). 12c);2934(2-1).
Ravn, P., s.n. (4-7al). Sargent,C. S., s.n.(Apr 1887)(1-3).
Record, S. J., G 20(1-lb). Scala, A. C., 138(?),275(?)(LP31753)(4-la).
Regnell, A. F., I 273x(2-4);II 53(1-22);II 54(2-5); Schenck, J. H. R., 2330(1-9a2)
III 373(1-9a1). Schiller, I., 668, 803(1-3).
Reitz, R., 1727(4-la); 3141(1-9al); 4355(1-23); Schinini,A., 4881, 5095(4-la).
6044(1-9a1). Schipp, W. A., 248, S-643(1-lb).
Reitz, R. & R. M. Klein, 826(1-24); 1116, 2004(1- Schomburgk,R., 153.5(1-17a).
9a1); 3828(3-1); 4104(1-24); 4685(4-13); 4688, Schott, H. W., 69v(1-22); 7496(=Herb. Bras.
4805(1-9al); 5914(4-13); 6198(1-9al); 7359(3- 4754)(1-9al); Herb. Bras. 4458(4-12a2); Herb.
1); 7401, 7578(4-la); 8279(1-24); 8818(4-la); Bras. 4757(4-9).
9040, 9461, 9595, 9633, 12492(1-9al). Schott, H. W. & J. E. Pohl, D 999(=Herb. Bras.
Restinga I (see also Segadas-Vianna), 659(4- 56118)(1-14).
12a2). Schreiner,R 71497(1-22).
Ribeiro, R 71092(4-12a2). Schiicht, J., s.n.(2-4).
Richard, L. Cl. M., s.n. (4-7al). Schulz, A. G., 3948(4-la).
192
Schunke V., J., 1983, 3290(4-10); 5818(1-18).
Schwacke, W., 1535(2-4); II 164, R 71066(1-9al).
Schwarz, G. J., 1208, 2593, 2634, 2792, 2835,
2915, 3020, 3083, 3105, 3132, 3254, 4725,
4766(4-1a); 7415, 7489, 7490(1-22); 7884,
7909, 10491, 10647, 10785(4-la).
Schwindt, E., 734, 4887(4-la).
Segadas-Vianna, F., 4194(4-12a2).
Segadas-Vianna, F. et al., 913(1-9al).
Sehnem, A., 1537(4-la).
Sellow, F., 5(?), 6(?)(4-12al); 144, 563, 676,
1406(2-4); 1424(1-22); 1645(1-9al); 2170(2-5);
2171, B 2171.c 2168(1-9al); 2172, B 2172,
563(2172.c 2169), B 2173(2-4); 2177, B 2177.c
2173(1-22); 2178, B 2178.c 2174(4-12al);
4021(4-la); 5310(1-22); 5495(1-9al); 5998(4-
9); s.n.(l-9al); s.n.(1-22); s.n.(2-4); s.n.(2-5);
s.n.(4-3); s.n.(4-12al).
Shafer, J. A., 445, 2387, 2557, 3610, 3616, 4219,
8323, 8366, 8779(4-7al).
Shreve, F., 6765(1-13); 7241(1-12).
Sieber, F. W., 34(1-17a); s.n.(4-7al).
Silva, A. S. et al., 17(1-9al).
Silva, M. & R. Souza, 2266(2-2).
Silva, N. T., 181, 57778(2-2).
Sintensis, P., 5649, 5751(4-7al).
Smith, A. C., 2361(1-17a).
Smith Jr., C. E. et al., 3721(1-11).
Smith, L. B. (et al.), 1238(2-4); 7389(1-9al);
11784(4-la); 12410(3-1); 12416, 12696,
12747(4-la); 12931(1-9al); 12940, 13110(4-
la); 14447, 15389(1-9al).
Smith, L. B. & A. Macedo (in Macedo), 4652(1-
7).
Smith, R. F., V659(4-7al); V886(1-17a).
Smith, S. G., 1136(4-8b).
Souza, A. B. de, RB 90086(4-12a2).
Stahel, G., 361(1-17a).
Standley, P. C., 9363, 9377, 9495, 15577(1-2b);
20093(4-7b); 61395(1-10); 73711, 73742, 73781,
74036, 74303(1-2b); 74413, 74456(1-10).
Standley, P. C. & J. Valerio, 44213, 45662,
45744(4-7b).
Standley, P. C., L. Williams & P. Allen,
548(1-2b).
Steere, W. C., 2956(1-2a).
Steh6le,H., 394(4-7al).
Stephan, s.n.(1843)(1-22).
Stern, W. L. et al., 1822(1-lc).
Steyermark, J. A. (et al.), 29345(1-2b);46452(1-10); Wilkes, Ch., s.n.(1838-1842)(4-12al).
55569(4-8al); 57682, 62839, 87967(1-17a);
87998, 88161(4-7al); 88198(1-17a); 88515(4-
7al); 88527(1-9al); 94720, 99155(4-8al);
106496(1-9al); 107802(1-17a); 110262,
111690(4-8a1).
Stiles, H. (in Runyon), 177(1-3).
Stoffers, A. L., 1745-A, 6259(4-8al).
Sucre, D. (et al.), 1887(4-12al); 8335(4-11); 8506(4-
12al); 8706, 8947(4-11); 9446, 9448(4-6);
10192(4-9).
Sullivan, J. R., 639(1-3).
Swabey, C., TRIN 12525(1-17a).
Herb. Swartz, 0. P., s.n.(2-4).
Flora Neotropica
Tamayo, F., 755, 1109(4-8al).
Triana, J. J., s.n.(3741, 600 m), s.n.(3741, 700 m),
s.n.(3742, 600 m), s.n.(3762?, 600 m),
s.n.(3841?, 600 m), s.n.(Feb 1854, 700 m),
s.n.(500 m), s.n.(600 m), s.n.(700 m),
s.n.(800 m), s.n.(1-19).
Herb. TRIN (see also Finlay & Hart), 3853(1-
9al).
Tube, s.n.(5998?), s.n.(Apr 1871)(2-4).
Turner, B. L., 7(4-7b); 2086(1-5b).
Ule, E., 349(4-13); 500(1-24); 1297(4-la); 3556(2-
4); 5806(1-9al); 9491, 9492(2-2); 9493, 9494(1-
25); 9500(4-10); 9501(1-21); s.n.(Jul 1891)(1-
9al).
Usteri, A., 40a(l-9al); SP 19824(4-9).
Vauthier, 5(?), 159, s.n.(4-12al).
Vecchi, O., CPEF 162(2-5); CPEF 163(1-14);
CPEF 282(1-9al); ESP 212(1-22).
Veith-Rohrer, J., s.n.(9 Mar 1910)(4-la).
Velloso, H. P., 961(4-12al); 994, 1000(4-11).
Ventura A., F., 2843, 3935, 4331(1-2a).
Vidal, J., 1712(1-9al); III 1652(4-12a2).
Vi6gas, A. P. (et al.), IAC 2376, IAC 3000(2-4).
Virlet d'Aoust, P. T., 1884(1-3).
Vogel, s.n.(30 Jan 1897)(2-4).
Wagner, M. F., 307, s.n.(4-8al).
Comm. Wardleworth (Evans, Sons & Co., Liv-
erpool), s.n.(31 Oct 1893), s.n.(8 Nov
1893)(4-6).
Warming, J. E. B., 510(Herb. 2467/2), 549(Herb.
2467/3), 869(?)(Herb. 2467/1), s.n.(2 Jan
1865), s.n.(2-5); s.n.(Lagoa Santa)(1-22).
Herb. Warming, J. E. B., 2451/1(1-9a2); 2454(4-
12al); 2463(1-22).
Warscewicz, J. von, s.n.(Ja6n de Bracamoras)(l-
15); s.n.(Sonda)(l-20).
Weberbauer, A., 6584, 7110, 7645(1-20).
Webster, G. L. et al., 11232(1-3); 12012, 12031,
13006(1-2b).
Weddell, H. A., 2967(1-7); 3448(1-25).
White, S. S., 2921(1-13).
Whitehead, J., 841(1-12).
Widgren, J. F., 483(2-4); 679(4-12al); 1077(2-4);
1078(1-22); s.n.(1845)(1-9al); s.n.(1845)(2-
4); s.n.(1845)(2-5); s.n.(4-12al).
Wied-Neuwied, M. A. Ph. Prinz zu, s.n.(1828)(2-
4); s.n.(l-17b); s.n.(2-5); s.n.(4-12al).
Wiggins, I. L., 5590, 5671(1-12); 7459(1-13);
11503, 14543, 15015, 15425(1-12).
Williams, L. 0. et al., 6093(1-22); 10228, 11050,
14321, 16716(1-2b); 26618(4-7b).
Williams, LI., 4878(4-10); 9681(l-2b).
Williams, R. O., TRIN 12169(1-17a).
Williams, R. S., 482(2-2).
Wilson, P. 9177, 9549(4-7al).
Wilson, P. & J. S. S. Le6n, 1949(4-7al).
Winkler & Hanke, 627(4-7al).
Woloszczak, 4047(?)(1884)(4- la).
Woolston, A. L., 256, 256-c(l-22); 724(4-la).
Woytkowski, F., 7646(1-18).
Wright, C., 1129, s.n.(4-7al).
Index of Local Names
Wright,Wm., 12(1-la).
Comm. Wright et al., s.n.(Sep 1903)(4-7al); s.n.
(Jan 1905)(4-6).
INDEX OF LOCAL NAMES
Amar6, excluded from Metrodorea.
Amari, excluded from Metrodorea.
Angohuara, E. warscewiczii, 1-20.
Apochi-taguara, E. grandiflora var. grandiflora,
l-9al.
Apojitaguara, E. grandiflora var. grandiflora,
l-9al.
Arco-de-pipa, M. stipularis, 2-5.
Arruda, P. microphyllus, 4-6.
Arruda-brava, P. microphyllus, 4-6.
Arruda-do-mato, P. microphyllus, 4-6; P. trachy-
lophus, 4-5.
Bananeira-do-mato, M. stipularis, 2-5.
Barbasco caspi, E. amazonica, 1-18.
Bois blanc, P. racemosus var. racemosus, 4-7al.
Bois flambeau caraibe, P. racemosus var. race-
mosus, 4-7a1.
Boragica, E. pilocarpoides subsp. maurioides,
1-17b.
Borrachera, P. goudotianus var. goudotianus,
4-8a1.
Burachi, P. goudotianus var. goudotianus,
4-8a1.
Camba canilla, P. pennatifolius var. pennatifol-
ius, 4-la.
Candlewood, E. pentaphylla subsp. belizensis,
I-lb.
Canela-(de-)cutia, E. grandiflora var. grandi-
flora, 1-9a1.
Canela-de-cutia, P. pennatifolius var. pennati-
folius, 4-la.
Canela-de-veado, E. densiflora, 1-23.
Canelero-del-monte, E. grandiflora var. grandi-
flora, l-9a1.
Caputuna, E. grandiflora var. grandiflora, 1-9al;
M. stipularis, 2-5.
Carrapateirao, E. grandifiora var. grandiflora,
1-9al.
Carrapateiro, M. nigra, 2-4.
Catagua, M. stipularis, 2-5.
Catigua, P. trachylophus, 2-5(?).
Chupa-ferro, M. nigra, 2-4; M. stipularis, 2-5(?).
Cocao (?), E. grandiflora var. grandiflora, 1-9al.
Coya, E. alata, 1-19.
Crucecilla (?), E. hartmanii, 1-13.
Cuala (-cuala), E. alata, 1-13.
Cutia, E. grandiflora var. grandiflora, 1-9al.
Cutia-amarela, E. grandiflora var. grandiflora, 1-
9al.
Cutia-branca, P. pennatifolius var. pennatifolius,
4-la.
Gasparee, E. pilocarpoides subsp. pilocarpoides,
1-17a.
Gaspari(llo), E. pilocarpoides subsp. pilocar-
poides, 1-17a.
193
Xantus de Vesey, L. J., s.n.(Aug 1859-Jan
1860)(1-12).
Zehntner, 75, 975(4-5).
Ziircher, L., 69(4-1a); 165(52.36)(1-23).
Gaspari colorado, E. grandifiora var. grandi-
flora, 1-9al.
Gasparil, E. grandiflora var. grandiflora, 1-9al.
Gasparillo colorado, E. grandiflora var. grandi-
flora, I-9a1.
Grumarim (-de-campos), E. febrifuga, 1-22.
Guarantan, E. leiocarpa, 1-14.
Guarataia-vermelha, E. leiocarpa, 1-14.
Guatambu, P. pennatifolius var. pennatifolius,
4-la.
Guaxupita, E. grandiflora var. grandiflora,
1-9al.
Hokob, E. berlandieri subsp. berlandieri, l-2a.
Huacac, E. runyonii, 1-3.
Ibira-tai, P. pennatifolius var. pennatifolius,
4-la.
Ivira-nfieti-(m)i,E. febrifuga, 1-22.
Ivira-ovi-guacu, E. densiflora, 1-23.
Ivira-tai, P. pennatifolius var. pennatifolius,
4-la.
Laranjeira (-do-mato), M. stipularis, 2-5.
Laranjeiro-do-mato, E. febrifuga, 1-22.
Limoeira-do-mato, M. stipularis, 2-5.
Limonillo, E. runyonii, 1-3.
Loro, E. pentaphylla subsp. australensis, 1-lc.
Mamoninha, E. febrifuga, 1-22.
Matasarna, P. goudotianus var. goudotianus,
4-8a1.
Mendanha, E. febrifuga, 1-22.
Palo amarillo, E. flava, 1-12.
Palo morio, E. flava, 1-12.
Palu cayente, P. goudotianus var. goudotianus,
4-8a1.
Pau-cutia, E. grandiflora var. grandiflora, 1-9al.
Pipoca, E. pilocarpoides subsp. maurioides,
1-17b.
Pitaguara-do-branco, P. paucifiorus, 4-13.
Samota, E. hartmanii, 1-13.
Sapoto jaru, P. peruvianus, 4-10.
Talcacao, P. racemosus subsp. viridulus, 4-7b.
Tancache, P. racemosus subsp. viridulus, 4-7b.
Tres-folhas, E. febrifuga, 1-22.
Tres-folhas-do-mato, E. febrifuga, 1-22.
Tres-folhas-vermelha, E. febrifuga, 1-22.
Verde lucero, E. pentaphylla subsp. belizensis,
I-lb.
Vira sarere, M. nigra, 2-4.
Wild orange, E. pentaphylla subsp. pentaphylla,
1-la.
Yax-hokob, E. berlandieri subsp. berlandieri,
1-2a.
Yerba-de-cutia, P. pennatifolius var. pennatifol-
ius, 4-la.
194 Flora Neotropica

INDEX OF TAXA
Page numbersin bold face indicateprimarypage references. Page numberswith
an asterisk (*) indicate pages with illustrationsor maps.
Adiscanthus 24 cowanii 8, 26, 110*, 112, 113*
Angostura 5, 8, 20, 103 cuspidata 114
Apoplanesia 45 densiflora 4, 27, 86*, 103, 104*, 106, 108
Balfourodendron 6 dielsiana 94, 97
eburneum 116 echinoidea 27, 35*, 66, 67*, 68
riedelianum 116 sect. Esenbeckia 7, 21, 22, 23*, 25, 29, 83
Bocoa 153 subgen. Esenbeckia 6, 23*, 26, 29
Bombacaceae 25 sect. Euesenbeckia 29
Boronia 17 fasciculata 58, 62, 63
Boronieae 17 febrifuga 10, 11, 12, 15, 17, 20, 27, 54, 62,
Bursera 45 86*, 99, 100*, 101, 103, 106, 108, 130
Carovaglia 25 var. densiflora 103
Casimiroa 21 var. fruticosa 101
Citrus 9 feddemae 7, 29, 35*, 43, 44*
medica 40 flava 6, 8, 10, 27, 70, 72*, 73*
Colythrum 25 glaziovii 115
alatum 94 gracilis 103, 106
febrifugum 101 grandiflora 5, 8, 9, 16, 20, 22, 27, 29, 57, 58,
grandiflorum 58 62, 63, 65, 68
latifolium 53 subsp. brevipetiolata 58, 59*, 66
maurioides 88 subsp. grandiflora 58, 66
puberulum 25, 53, 54 subsp. grandiflora var. grandiflora 58,
pumilum 53 59*, 60*, 62, 63, 65, 66
Conomorpha subsp. grandiflora var. intermedia 58,
verticillata 115 59*, 63, 64*, 65
Correa 17 var. macrophylla 58, 63
Cusparieae 1, 2, 3, 20, 21, 22 var. peruviana 61, 63
Cuspariinae 1, 3, 6, 8, 17, 20, 21, 24 hartmanii 3, 8, 17, 27, 72*, 74, 75*
Diosmeae 2, 3, 21 hieronymi 8, 9, 27, 29, 86*, 103, 106, 107*
Erythrochiton sect. Hymenopetalae 29, 83
brasiliensis 17 intermededia 63
Esenbeckia 1, 2, 3, 4, 5, 6, 7, 8, 10, 12, 17, 20, intermedia 20, 63, 65
21, 22, 23*, 24, 25, 26, 29, 35*, 40, 43, 59*, 72*, irwiniana 28, 48, 50*, 59*
76, 86*, 99, 103, 110*, 114, Addenda laevicarpa 79
acapulcensis 40 subgen. Lateriflorens 6, 10, 21, 22, 23*, 26,
alata 7, 27, 28, 83, 86*, 91, 92*, Addenda 108, 112
var. laevis 91, 94 latifolia 51, 54, 55
almawillia 9, 11*, 26, 108, 110*, 111* leiocarpa 5, 6, 7, 8, 9, 12, 17, 26, 29, 43, 72*,
altissima 114 76, 77*, 79, 81, 83, 112
amazonica 7, 22, 27, 86*, 89, 90*, 91 leucophylla 10, 53
atata 12, 114, 115 litoralis 38, Addenda
attenuata 29, 58, 62, 63 lucida 115
belizensis 31 macrantha 8, 28, 35*, 68, 69*
berlandieri 3, 22, 28, 33, 34, 36, 37*, 38, 41, maurioides 88
43, 45, 114 sect. Metrodorea 29, 116
subsp. acapulcensis 10, 35*, 36, 37*, 40 mollis 115
subsp. berlandieri 27, 29, 30, 33, 35*, 36, nesiotica 27, 35*, 55, 56*
37*, 38, 39, 40, 43 obovalifolia 61, 63
subsp. litoralis 8, 28, 35*, 36, 37*, 38, oligantha 6, 7, 9, 26, 72*, 81, 82*
39, 40, Addenda subgen. Oppositifolia 4, 6, 9, 21, 23*, 25, 26,
castanocarpa 29, 83, 84 99
choisyoides 53, 54 ovata 36
collina 28, 29, 45, 46 sect. Pachypetalae 23*, 29, Addenda
subsp. collina 35*, 46, 47*, 48 panamensis 26, Addenda
subsp. conspecta 35*, 46, 48, 49* pentaphylla 22, 28, 29, 30, 31, 32*, 33, 38
coriacea 114, 119 subsp. australis 27, 31, 32*, 33, 35*, Ad-
cornuta 7, 9, 26, 29, 72*, 78, 79, 80, 81, 83, denda
89, 91 subsp. belizensis 3, 31, 32*, 33, 34, 35*
Index of Taxa 195

subsp. pentaphylla 22, 28, 31, 32*, 33, Metrodorea 3, 4, 5, 6, 7, 8, 9, 10, 12, 17, 21, 22,
34, 35* 24, 29, 110*, 116, 117, 127, 130
pilocarpoides 6, 8, 9, 12, 15*, 22, 25, 27, 29, atropurpurea 123
62, 83, 84, 85, 91, 93, 114, 181 brevifolia 125, 127
var. guianensis 114 excelsa 130
subsp. maurioides 27, 83, 84, 86*, 88 flavida 3, 10, 12, 22, 110*, 114, 117, 119, 120*,
var. maurioides 88 122
subsp. pilocarpoides 84, 86*, 87* gracilis 101, 103, 106, 130
pilosa 39, 40 maracasana 110*, 117, 118*, 119, 127
pittieri 115 mollis 110*, 117, 118*, 122
plicata 116 var. glabrata 122, 123
sect. Polembryum 29, 83 nigra 12, 110*, 117, 119, 122, 123, 124*, 127,
pumila 3, 4, 10, 15, 20, 27, 28, 29, 51, 52*, 130, 181
54, 55, 59*, 103 var. brevifolia 125, 127
var. genuina 53 var. nigra 127
var. latifolia 53 pubescens 11, 20, 123, 127, 130
var. leucophylla 53, 54 selloana 125, 127
var. pumila 54 stipularis 11, 20, 110*, 117, 119, 127, 128*
pusilla 51 Monnieria
riedeliana 116, 175 trifolia Addenda
rigida 61, 63 Musci 25
runyonii 5, 22, 28, 35*, 38, 41, 42*, 43 Myrsinaceae 115
scrotiformis 5-6, 7, 10, 28, 86*, 97, 98* Neesia 25
venezuelensis 85 altissima 114
venulosa 94, 97 Phebalium 17
warscewiczii 6, 9, 28, 29, 48, 86*, 94, 95*, Picrella 38
97 trifoliata 38
yaaxhokob 36, 38 Pilocarpea 3, 22
Eucusparieae 3 Pilocarpeae 3, 22
Euodia 25 Pilocarpinae 1, 2, 3, 6, 8, 9, 11, 17, 20, 21, 22,
febrifuga 99 24, 63, 175, Addenda
Euonymus Pilocarpus 1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13,
latifolius, racemosus 2, 132 15, 16, 17, 18, 19, 20, 21, 22, 23*, 24, 132, 133,
Evodia 25 138*, 175, 181, Addenda
febrifuga 99 alvaradoi 161
Evonymus var. mollis 163
lati-folius, racemosus 2, 132 amarus 181
Fraxinellae 2 atropurpureus 125, 181
Galipea breviracemosus 178, 180, 181
lucida 115 cearensis 145, 148
pentaphylla 30 demerarae 9, 13, 15*, 133, 138*, 141, 142*,
Garovaglia 25 145
plicata 116 fluminensis 178, 180
Glycosmis 9 giganteus 5, 8, 9, 15, 16, 18, 19, 134, 138*,
Helietta 6, 20, 38, 114, 115 164, 165*, 166
apiculata 115 goudotianus 15, 22, 133, 154*, 158, 160, 161,
cuspidata 115 162*
longifoliata 114, 115 subsp. goudotianus 158, 161
mollis 115, 123 subsp. goudotianus var. goudotianus
multiflora 116 154*, 161, 162*, 163, 164
parvifolia 38, 114, 115 subsp. goudotianus var. mollis 154*, 158,
plaeana 12, 114, 115 161, 162*, 163
Herpestis 17 subsp. heterochromus 154*, 158, 161,
Kuala 25 162*, 164
alata 25, 91, 93, 94 grandiflorus 7, 9, 133, 138*, 143, 144*, 145
laevis 91, 92*, 93, 94 guyanensis 155, 159
Leguminosae 153 heterophyllus 18, 19, 157
Leptothyrsa 24 humboldtii 84, 181
Lonchocarpus 45 insularis 157, 158
Lunasia jaborandi 1, 2, 5, 10, 11-12, 12, 13, 15, 16,
amara 181 17, 18, 19, 133, 138*, 145, 146*, 148, 150
Lysiloma 45 latifolius 15, 155, 158, 159
Meliaceae 9, 150 laurifolius 157
196 Flora Neotropica

lealii 175, 176 forma brevipedicellata 136

longeracemosus 176 var. gracilis 136
var. breviusculus 176, 177 simplex 136, 140
longipes 10, 157, 158 spicatus 6, 7, 10, 11, 12, 13, 15, 16, 133, 134,
macrocarpus 18, 19, 164, 166 168, 171, 172, 173*, 175, 180
macrophyllus 19 subsp. aracatensis 19, 115, 138*, 172,
microphyllus 1, 2, 10, 13, 15, 16, 17, 18, 19, 173*, 177
115, 133, 138*, 150, 151, 152* subsp. longeracemosus 138*, 172, 173*,
minutiflorus 175, 176 175, 176, 177
officinalis 145 var. peruvianus 166, 168
organensis 178, 181 subsp. spicatus 172, 177
parviflorus 10, 172 subsp. spicatus var. lealii 6, 138*, 172,
pauciflorus 9, 133, 134, 138*, 155, 172, 178, 173*, 175, 176
179*, 180, 181 subsp. spicatus var. spicatus 6, 138*,
pennatifolius 5, 6, 9, 10, 12, 13, 15, 16, 17, 169-171, 172, 173*, 174, 175, 175-176,
18, 19, 20, 133, 134, 135, 136, 139*, 141, 180
143, 145, 148 var. subcoriaceus 174
var. genuinus 136 subcoriaceus 10, 174
var. genuinus forma gracilis 136 trachylophus 2, 5, 7, 10, 12, 13, 15, 16, 20,
var. genuinus forma gracilis subforma 133, 138*, 148, 149*, 151
deorsum-bracteolatus 136 trijugatus 136
var. genuinus forma gracilis subforma ypanemensis 174, 175
sursum-bracteolatus 136 Piper 17
var. genuinus forma latifoliolatus 136 Polembrium 25
var. genuinus forma typicus 136 Polembryum 25
var. pennatifolius 135, 136, 138*, 139*, castaneaecarpum 58
140 castanocarpum 25, 58
var. pilosus 135, 138*, 139*, 140, 141 jussieui 58, 62, 63
var. selloanus 136 Prunus
var. selloanus forma brasiliensis 136 floribus racemosis 132
var. selloanus forma intermedius 136 Raputia
var. selloanus forma paraguariensis 136 heterophylla 157
peruvianus 134, 138*, 166, 167* Raulinoa 3, 4, 5, 6, 7, 8, 10, 17, 21, 22, 24, 110*,
pinnatifidus 134 130
pinnatifolius 134 echinata 110*, 130, 131*
pinnatus 136 Ravenia
racemosus 2, 3, 4, 10, 11, 12, 13, 15, 16, 18, spectabilis 17
19, 22, 133, 134, 153, 154*, 155, 156*, 158, Ruta 9
159, 161 Rutaceae 2, 3, 5, 7, 9, 17, 20, 21, 22
subsp. racemosus 155 Rutoideae 22
subsp. racemosus var. racemosus 154*, Simaroubaceae 7, 9
155, 156*, 158-159, 159 Swartzia
subsp. racemosus var. yucatanus 154*, decipiens 153
155, 156*, 158, 159 Xanthoxyleae 2
subsp. viridulus 154*, 155, 156*, 158, 160 Zanthoxyleae 3, 21
riedelianus 9, 10, 133, 134, 138*, 166, 169, Zanthoxylum 7
170* fagara 30
selloanus 18, 136, 140, 148
Addenda 197

ADDENDA
1. Recently some new collections, mainlyfrom Bahia, came to my notice. They
are mentionedif they provide new data about distribution,but are not mapped.
2. After the completionof my manuscript,Porter, D. M. & Th. S. Elias, Family
89. Rutaceae. In: Woodson Jr., R. E. et al., Flora of Panama. Ann. Missouri
Bot. Gard.66: 123-164. 1979,was published.Of the Pilocarpinaeonly Esenbeckia
is described (by Elias), with three species: 1) Esenbeckia litoralis = E. berlandieri
Baillon ex Hemsley subsp. litoralis (Donnell-Smith)Kaastra; 2) E. alata = E.
pentaphylla (Macfadyen)Grisebachsubsp. australensis Kaastra. See e.g. Stern
et al. 1522, cited by Elias and me, and the descriptionby Elias of the size of the
tree and of the fruitswith secondaryapophyses;and 3) E. panamensis Elias. Some
measurements appear to differ, and I disagree with Elias in the phyllotaxy,
which is essentially alternate (not opposite!), to ternate or aggregate towards
the tip of the branches. The simple leaves are attenuate at the base. The in-
florescence of the one floweringspecimen seen (the type) is minutelypubescent,
with alternate side-branchlets. The flowers are ca. 11 mm in diam., entirely
glabrous(includingthe calyx!);petals ca. 4.5! x 3 mm, roundedat apex, ? papery;
filamentsca. 2.3 mm, lackingan appendage;anthersca. 1.2 mm long; style ca. 0.7
mm long. Fruits 1.4-2 x 2-3 cm, the wrinkledloculi provided with a blunt, ? re-
curved apophysis 3-7 mm long. Distribution:E Panama (Colon, Darien) and
Colombia (N Choco). Wet forest; alt. 300-1100 m. Flowering Jul (one record).
Specimens examined: Panama. Same as cited in protologue; except the type all
fr. In addition:Darien:trailalong ridge S of Rio Setigandi,nr. Colombianborder,
A. H. Gentry et al. 28599 dec. fr (MO). Colombia. None seen. Esenbeckia
panamensis belongs to sect. Pachypetalae of subgen. Esenbeckia.
3. Holmstedt et al. (1979) gave a review of the etymology of Jaborandi,its
Indian and western medicinaluse, the identificationof Folia Jaborandi,and the
pharmacologyof pilocarpine. I disagree with the authors in the identificationof
Marcgrave'sJaborandi(fig. 1) as a Pilocarpus species. The bifurcate inflores-
cence with "bracts" (=bractlike sepals), the indument,and the size of the plant
suggest Monnieriatrifolia Linnaeus.The other cited Jaborandi'sfrom Marcgrave
and Piso are even more unrelatedto Pilocarpus.