Organization for Flora Neotropica

Balanophoraceae
Author(s): Bertel Hansen
Source: Flora Neotropica, Vol. 23, Balanophoraceae (Jun. 16, 1980), pp. 1-80
Published by: New York Botanical Garden Press on behalf of Organization for Flora
Neotropica
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FLORA NEOTROPIC

MONOGRAPH NUMBER 23

BALANOPHORACEAE

by
Bertel Hansen

FLORA/
,>
NEOTROPICA'

, I
I I-i Op pl AN

Published for

Organization for Flora Neotropica

by
The New York Botanical Garden

New York
1980
June 16, 1980
 Copyright ( 1980

The New York Botanical Garden

Library of Congress Cataloging in Publication Data

Hansen, Bertel.
Balanophoraceae.

(Flora neotropica ; monograph no. 23)

Bibliography: p.
Includes index.
1. Balanophoraceae. 2. Botany--America.
3. Botany--Tropics. I. Title. II. Series.
QyK495.B23H34 583 ' 94 79-28385
ISBN 0-89327-195-0

Published by
The New York Botanical Garden
Bronx, New York 10458

International Standard Serial Number 0071-5794

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

BERTELHANSEN'

INTRODUCTION

The systematic part of the present treatment is confined to the 15 species,

2 subspecies and 2 varieties found in America within 7 genera of the
Balanophoraceae. They are distributed from Mexico and the Galapagos to SE
Brazil, NW Argentina and Chile. Six genera are confined to America, while
Langsdorffia has one widespread species in America, 1 in Madagascar and 1 in
New Guinea. The family in its broadest sense contains 43 species in 18 genera,
distributed mainly in the tropics, occasionally in the subtropics, rarely
reaching the temperate regions.

HISTORY OF AMERICAN BALANOPHORACEAE

Browne (1756) mentioned a Cynomorium from Jamaica, quite obviously

belonging to what is now known as Scybalium jamaicense. However, binary
nomenclature was not used in his work, and the first valid publication of
American species in Cynomorium was by Swartz (1788), ie C. cayennense and
C. jamaicense. These taxa were mentioned again by Swartz (1797) under the
same binomials.
Martius (1818) described the first Langsdorffia, ie L. hypogaea.
Richard (1822) posthumously described the family Balanophoraceae
("Balanophoreae") nom. cons., and gave an account of the genera and species
then known, ie Helosis guyannensis, a superfluous name for Cynomorium
cayennense, and H. jamaicensis transferred from Swartz's combination with
Cynomorium, Langsdorffia janeirensis, Balanophora fungosa, and Cyno-
morium coccineum.
Sprengel (1826) made the valid combination, Helosis cayennensis and also
listed H. jamaicensis.
Martius (1832) described and illustrated in detail Langsdorffia hypogaea
and Helosis guyanensis, correctly placing Langsdorffia janeirensis into
synonymy under L. hypogaea.
Schott & Endlicher (1832) described Lophophytum mirabile and
Scybalium fungiforme, thus erecting two new genera in the family. The
descriptions were accompanied by surprisingly detailed illustrations. Further-
more they described briefly Helosis brasiliensis ("brasileensis") and the com-
bination Scybalium jamaicense was made, based on Cynomorium jamaicense.
Poeppig (1833), in a review, mentioned a new generic name, Om-
brophytum, without giving a description. The name was validly published by
Endlicher (1836).

'Curator of Phanerogams, Botanical Museum, University of Copenhagen, Gothersgade 130,

DK-1123 Copenhagen K, Denmark.
1
2 Flora Neotropica

Saint-Hilaire (1837) published the generic name Archimedea and combin-

ed it with the epithet pyramidalis to name an entity apparently belonging with
Lophophytum mirabile as far as the description goes, see also p. 47.
Poeppig & Endlicher (1838) described and illustrated Ombrophytum
peruvianum. In a note they mentioned Poeppig's observation of another
species with smaller, violet colored plants.
Liebmann (1847) described Helosis mexicana and Thonningia mexicana
based on material collected by himself. He had two excellent plates made in
colour, but they were never printed. They are now in the collections of the
Copenhagen herbarium.
Klotzsch (1847) described Langsdorffia moritziana from Colonia Tovar
and Weddell (1850) published illustrations with descriptions of Langsdorffia
rubiginosa, Lophophytum bolivianum and Ombrophytum zamioides together
with accounts of already known taxa of Balanophoraceae, Hydnoraceae and
Rafflesiaceae.
J. D. Hooker (1856) gave a detailed and illustrated account of the
Balanophoraceae known up to then. He described the genera Sphaerorhizon
and Phyllocoryne with one species each, and Corynaea with three species. He
correctly transferred Liebmann's Thonningia mexicana to Langsdorffia and
placed it as a synonym of L. hypogaea. Lophophytum mirabile was partly con-
fused with a hitherto undescribed entity, partly due to scanty material, partly
due to Saint-Hilaire's and Weddell's earlier confusion, see p. 47. For some
strange reason Sphaerorhizon appears with one specific name in the text and
another in the plate.
Eichler (1868) described and illustrated a remarkable new genus and
species, Lathrophytum peckoltii, completely destitute of leaves and bracts. As
indicated later this statement, often repeated in subsequent literature, was due
to the use of very old, fruiting material as a basis for his description. In 1869
Eichler's treatment of the Balanophoraceae in Flora Brasiliensis was published
containing a wealth of detailed information on this group of plants.
Langsdorffia leandri was described here. Finally Eichler (1873) treated the
family in de Candolle's Prodromus. He described Scybalium glaziovii and
united Hooker's Phyllocoryne and Sphaerorhizon in Scybalium. In Corynaea
he described a fourth species, C. sprucei. The Langsdorffia species known un-
til then were all united under L. hypogaea.
Engler (1889) treated the family in Die Natiirlichen Pflanzenfamlien. He
proposed four subfamilies of which Scybalioideae is entirely American and
Balanophoroideae includes the partly American genus Langsdorffia.
Baker (1890) published Helosis whymperi from Ecuador, an entity shown
here to belong in Corynaea, see p. 44.
Asplund (1928) described and illustrated Juelia subterranea, a new genus
and species from the Bolivian high Andes.
Standley (1930, 1931) treated the family in Yucatan and Honduras, and
Lanjouw in Pulle (1932) gave an account of the family in Suriname.
Harms (1935) in the second edition of Die Natiirlichen Pflanzenfamilien
slightly remodelled Engler's subfamilies. He particularly stressed a more
remote relation between Corynaea - Helosis - Scybalium and Lathrophytum
-Lophophytum - Ombrophytum - Juelia, and he placed the two groups in dif-
Balanophoraceae 3

ferent subfamilies. The binomial Ombrophytum ulei was proposed without a

latin description.
Macbride (1936) in the Flora of Peru included six genera containing eight
species.
Standley & Record (1936), Standley (1937), and Standley & Steyermark
(1946) treated the family in Honduras, Costa Rica and Guatemala.
Le6n & Alain (1951) in the Flora de Cuba mentioned Helosis guyanensis
and Scybalium jamaicense.
Sleumer (1954) treated the Balanophoraceae of Argentina and described
two new species in Juelia, ie J. lilloana and J. meyeri.
Lemee (1955) and Steyermark (1957) treated the family in French Guyana
and Venezuela respectively.
Howard (1959) dealt with the variation of Helosis populations in the
Caribbean flora and concluded that at present there was no sound basis for
splitting them into two or more species.
Nevling (1960) treated the family for the Flora of Panama including three
genera, Helosis, Corynaea, and Langsdorffia, each with one species.
Falcao (1975) treated the Balanophoraceae of the state of Santa Catarina,
Brazil.
Adsersen (1976) reported on the collection of an Ombrophytum peru-
vianum from the Galapagos Islands.
Hansen (1976a) emended the description of Lathrophytum peckoltii,
showing the presence of bracts, and (1977) published and illustrated a new
species, Ombrophytum violaceum, probably observed for the first time by
Poeppig (Poeppig & Endlicher 1838). In 1978 Hansen published changes in the
status of two taxa, ie Helosis cayennensis var mexicana and Corynaea crassa
var sprucei.

SYSTEMATIC POSITION OF THE BALANOPHORACEAE

Among local inhabitants throughout the distribution range of the

Balanophoraceae these plants are often not recognized as independent
organisms; according to Asplund (1928) Ombrophytum subterraneum is look-
ed upon as a kind of subterranean flower appearing from the roots of woody
plants otherwise bearing normal, supraterranean flowers. This view prevailed
even among biologists in the early 19th century, eg Trattinick (1828) and
Junghuhn (1841).
Richard (1822), when giving the first description of the family, placed its
members at the transition of monocotyledons and dicotyledons between the
Hydrocharitaceae and the Aristolochiaceae. Also Beccari (1869) considered
them close to the monocotyledons.
Blume (1828) published the class Rhizantheae for the parasitic genera
Rafflesia and Brugmansia, now Rafflesiaceae. Endlicher (1836) placed the
Rhizantheae together with the Balanophoreae, the Cytineae and the Raf-
flesieae between ferns and monocotyledons.
Griffith (1845) abandoned the Rhizantheae and moved the Balan-
4 Flora Neotropica

ophoraceae into the dicotyledons close to Urticaceae but without stating his
reasons.
J. D. Hooker (1856) discussed the position of the Balanophoraceae in
detail and pointed out the strong similarities in flower characters between
Cynomorium and Hippuris and particularly between the bistylar forms, such
as Lophophytum (he did not know Ombrophytum very well) and Gunnera.
Eichler (1868) also placed the family in the dicotyledons and found their
closest relatives to be in the Santalaceae because of the strong similarity of the
reductions of the female flower.
J. D. Hooker, in Bentham & Hooker f. (1880), apparently influenced by
Eichler, placed Loranthaceae, Santalaceae and Balanophoraceae together.
The position of the Balanophoraceae close to the Santalaceae has been
followed by most subsequent authors. It was strongly advocated by Fagerlind
(1945b), who discussed in detail and illustrated the similarities of the much
reduced female flowers in Santalaceae, Myzodendraceae, Loranthaceae and
Balanophoraceae.
However, Kuijt (1969) warned against attaching too much importance to
the reduction in the gynoecium. He pointed out, that in most parasites the
gynoecium is subject to reduction, and similarities may easily be due to con-
vergence rather than to relationship.
Dahlgren (1975) in his angiosperm system places the Balanophorales (ie
Balanophoraceae s.l.) close to Gunnerales and Haloragales.

INFLORESCENCES

In members of American Balanophoraceae the inflorescences are either

once branched as in subfam. Lophophytoideae or spadix-like and apparently
not branched as in subfam. Scybalioideae. In Langsdorffia (subfam.
Balanophoroideae) the female inflorescences are spadix-like while the male
ones are obviously racemose. Hansen & Engell (1978) treated in detail the
structure of the inflorescences of these subfamilies.
Lophophytum has an elongated inflorescence covered by the angularly-
ovate, imbricate peltas of peltate bracts, Fig 1, 1A and Fig 20. Each bract
subtends a branch of the first order of the inflorescence, Fig 1, 1B and 1C, the
female branches being in the lower part of the inflorescence and the male ones
in the upper part. The female flowers cover even the top of the branch, and
their two styles diverge transversally, Fig 1, 1B and 1C. In Ombrophytum
violaceum, Fig 24, the inflorescence is covered by imbricate, hexagonal peltas
of peltate bracts, Fig 1, 2A. Each bract subtends a branch of the first order
with the same distribution of sexes as in Lophophytum. The branch, however,
is peltately widened in its sterile top part, Fig 1, 2C and Fig 24.
In Scybalium glaziovii, Fig 11, the inflorescence is covered by imbricate,
triangular scales with transversely extended petioles attached along the entire
lower, twice bent margin, Fig 1, 3A. Each bract subtends a much depressed
branch of the first order. In fact the branches are depressed to an extent, that
the entire inflorescence has become quite spadix-like in appearance, when the
bracts are shed. Each branch is bisexual with female flowers on its peripheral
Balanophoraceae 5

IAA"

A IC

4 A & kB CS

FIG 1. Schematic representation of the structure of inflorescences in 1, Lophophvuin weddellii, 2,

Ombrophytum violaceutm, 3, Scybalium glaziovii, 4, Scvbalium depressum and 5, Helosis caven-
nensis.-A, four neighboring bracts in front view, the lowermost darkest; B, a single bract in
front view and the corresponding, axillary inflorescence branch; C, longitudinal section of B, in 1,
2 and 5 the position of the next higher bract shown by dotted lines.-Male flowers as open rings in
3B-5B or otherwise open structures in 3C-5C; female flowers black.- x 1.6.
6 FloraNeotropica

(ie lower)part and male ones on its central(ie upper)part, Fig 1, 3B and 3C,
Fig 2A. The female flowers are compressedand face with their flat sides
towardsthe petioles of the subtendingbracts;the two styles divergeover the
edgesof the ovaries.Note that the branchin Lophophytum,Fig 1, 1B and 1C
by being depressedwould transforminto a branchmuch like the one in Fig
1, 3B and 3C with regardto the arrangementof styles.
In Scybaliumdepressum,Fig 1, 4 and Fig 10, the imbricatebractscover-
ing the inflorescenceare distinctlypeltatewith a narrow,teretepetioleand an
angularly-ovatepelta. The branchof the firstorderis not even flat, but slightly
concavewith the male flowersinsertedat a lower level than the female ones,
Fig 1, 4C. A peculiar tendency in the female flowers to point their edges
towardsthe petioles of the subtendingbractsshould be noted, Fig 2B.
In Helosis, Fig 15, the bractscoveringthe inflorescenceare peltatewith
regular,hexagonal, valvate peltas, Fig 1, 5A. Each bract subtendsa totally
compressedbranchof the first order, Fig 1, 5B, which is obviouslyconcave,
Fig 1, 5C. The arrangementof the femaleflowersis the sameas in Scybalium
depressum,Fig 2C and D.
Corynaeahas the same arrangementas Helosis, but the branchesof the
first orderare even more concave,whichcausesthe centralmale flowersto be
partlycoveredby the peripheralfemale flowers, Fig 2E.
The bracts described above are considered homologous, the most
modified ones are those in Helosis and Corynaea.In fact the regular,hex-
agonalshapeof thesebractshaveoften misledearlierauthorsto considerthem
homologous to the sterile, apical, peltate part of the branches in Om-
brophytum,see Eichler (1868) and Fagerlind(1945b). In Ombrophytumthe
caducous bracts fall very early. As they are important charactersfor the
specific delimitation,collectors should always search for young specimens,
wherebractsare still present.

POLLENMORPHOLOGY

The family is stronglyeurypalynouswith colpate, colporateand porate

grains found in the neotropicalrepresentatives;even atremegrains occur in
subgenusBalaniaof the paleotropicalBalanophora,Hansen(1972).Generally
the exine structureand sculpturingis weakly developed but quite variable,
althoughin some taxa quite smooth grainsprevail.This variationcalls for a
descriptionat generic level and sometimes even at specific level. Another
fascinating feature is the occurence of binucleatepollen grains as well as
trinucleateones, even withinthe samegenus, Scybalium,and obviouslylinked
with the syndromesof secretoryversus smooth stigmas as demonstratedby
Hansen(1976b).
The pollen materialwas acetolyzedand chlorinatedusing the standard
methoddescribedby Erdtman(1952). A small amountof Coryluspollen was
always added, and thus each preparationcontains chlorinatedas well as un-
chlorinatedpollen grains of Corylusand the taxon under investigation.The
materialwas mounted in glycerinejelly and the cover glass was sealed with
paraffin. In some cases subsequentstainingwith feulgen was carriedout to
Balanophoraceae 7

"
00 OOo""
00
'^ 0?^ O 0^eO^
o&?g
WEjC ) ,, O 0J O00oo -%q
0
Coo

00_
- ,/- o, _ooQ Ze Qjl2 ooooooo

00 ~ B
0000L0

DOj5 OOUO
c?
1.-Cameralucida drawings, x5.

10 grainsof the taxon under investigation were measured. Averages were cor-

ohavebeen prepared to throw light upon minute exine details. However, the

grains appear 15-25 Co reduced in SEM-micrographs compared with LM

measurements. Consequently details measured on SEM-micrographs have
been corrected also to Corylus diameter 29.5 t~m.

Scybalieae-- Scybalium
Scybalieae Scybalium (Fig 3)
(Fig 3)
8 Flora Neotropica

Pollen grains 6(-7-8)-pororate, pantotreme, spheroid, medium-sized to

large. Pore and os circular to elliptic, sometimes irregularly shaped, mostly
with crenulate borders. Exine 1-layered, almost smooth. NPC 6(-7)66
Scybalium fungiforme. Pollen grains large, diam. 54 /m. Pores 8-18 /,m,
os 21-30 /m. Exine 1.8-2.1 /tm thick. Binucleate. Material studied: Gehrt sn
(Brazil).
Scybalium jamaicense. Pollen grains medium-sized, diam. 40-45 /zm.
Pores 7-11 tLm, os 14.5-16.5 pm. Exine 2 /zm thick. Binucleate. Material
studied: Alexander sn (Jamaica), Eggers sn (Santo Domingo).
Scybalium glaziovii. Pollen grains medium-sized, diam. 45 Lm. Pores
10-15.5 tim, os 12-18 tm. Exine 1.5 ptm thick. Material studied: Brade sn
(Brazil).
Scybalium depressum. Pollen grains medium-sized, diam. 26 pm. Pores
6-7 pm, os 8.5-10.5 /m. Exine 0.5-0.7 /m. Trinucleate. Material studied:
Holm-Nielsen et al 1267 (Ecuador), Steyermark 52634 (Ecuador).

Helosieae - Helosis, Corynaea.

Pollen grains isopolar, tricolpate, peritreme, suboblate, medium-sized.

Colpi with parallel borders, semicircularly rounded at apocolpi, often with a
circular pore at the end of the colpus. Exine apparently 1-layered, almost
smooth or faintly rugulate (Fig 4G). Trinucleate. NPC 343.
Helosis cayennensis (Fig 4). Pollen grains with polar axis P: 24-32 /m and
equatorial diam. E: 30-39 pm; P/E 0.79-0.83. Apocolpium diam. 4-5 Am. Col-
pi 3-4 pm wide. Exine 0.9-1.1 zm thick, towards the colpi up to 1.7 pm thick.
Material studied: var cayennensis: Holm-Nielsen et al 4155 (Ecuador),
Hatschbach 20981 (Brazil), Prance et al 16309 (Brazil); var mexicana: Hawkes,
Hjerting et Lester 2152 (Nicaragua), Liebmann sn (Mexico).
Corynaea crassa (Fig 5). Pollen grains with polar axis P: 22-26 lm and
equatorial diam. 26-32 tzm; P/E: 0.77-0.82. Apocolpium diam. 3-4 tm or
grains syncolpate in some specimens (Fig 5F, I). Colpi 3 /zm wide. Exine
0.7-1.1 /m thick. Material studied: var crassa: Asplund 10010 (Ecuador), Core
331a (Colombia), Forero 1051 (Colombia); var sprucei: Holm-Nielsen et al
6910 (Ecuador), Harling et Anderson 13209 (Ecuador), Gollmer sn (Costa
Rica).

Lophophytoideae - Lophophytum, Lathrophytum, Ombrophytum.

Pollen grains isopolar, tricolporate, medium-sized or poly-panto-colpate

and large. Exine 2-layered, smooth or variously sculptured. Binucleate. NPC
typically 345 in all taxa, with curious exceptions in individuals of
Lophophytum mirabile (665), Ombrophytum peruvianum (665) and 0. subter-
raneum (443 and 763).
Lophophytum leandri (Fig 6A-E). Pollen grains spheroid-subprolate
with polar axis P: 27-28 Atmand equatorial diam. E: 24 am; P/E: 1.1-1.2.

FIG 3. Pollen grains of Scybalium; A, B, D-F, H, I all LM x 1000; C, SEM x900, G, SEM
x 1290, K, SEM x 1910.-S. Jungiforme; A, high focus; B, optical section; C, surface view of
grain with irregular apertures.-S. glaziovii; D, optical section.-S. jamaicense; E, high focus; F,
Balanophoraceae 9

.... .? ~' ' jE ?

"
. _ ' .f fi i
%^

"IFF~

cal section; K surface view with one aperture open.-A-C, Ge.hr sn 1938; D, Brade sn 1948; E
and F, Alexander sn; G, Eggers sn 1887; H-K, Steyermark 52634.
10 Flora Neotropica

r4 e
x 1000;
FIG 4. Pollen grainsof Helosis cayennensis; A-F all LM G' SEM x 1275, H-K, SEM

.
ra;,i:6.._
., -r-!. _

? F.i

nensis var mexicana; . I, surface polar view; K, surface equatorial view.-, B, E, F and H, Hom-

Apocolpium diam.up.
9 m. Copi to 6 wide; os very distinct,
circular-.

mesocolpia reticulate with lumina about 0.1 /m diam., at apocolpia rugulate.

Material studied: Hatschbach 9842 (Brazil), Krapovickas 15509 (Argentina).
Balanophoraceae 11

.
....

'."i ? .i.,

..

:.-~~,, .::...

"q:*"
:j*" _. _::"':'
% i';Z:
nexine!); P/E: 1.24. Apocolpium
'm.diam. 7- . Colpi narrow, 1. wide; os

FIG 5. Pollen grains of Corynaea crassa; A, B, D-H all LM x 1000; C, SEM x 1660, I, SEM
x2040.-C. crassa var crassa; A, polar view in high focus; B, optical cross section; C, surface
polar view, one colpus artificially opened.-C. crassa var sprucei; D, polar view in high focus; E,
optical cross section; F, polar view in high focus of syncolpate grain; G, equatorial view in high
focus; H, optical longitudinal section; I, surface polar view of syncolpate grain, two colpi cracked
and interjacent mesocolpium slightly sunk into grain.-A-C, Asplund 10010; D, E, G, H, Holm-
Nielsen et al 6910; F, I, Haring & al 13209.

Lophophytum weddellii (Fig 6F-L). Pollen grains spheroid-subprolate

with polar axis P: 30.4 /m and equatorial diam. E: 24.5 /m (measured over

more or less distinct, lalongate, 9 x 1.8 Am. Exine 1.7 Mmthick, smooth or with
irregularly distributed nanoverrucae. Sexine 0.3 Am, irregularly loosened from
nexine except in the apocolpi and the central parts of the mesocolpi (Fig 6H-L)
(artefact ?). Material studied: Schunke 5675 (Peru).
Lophophytum mirabile (Fig 7A-F). Pollen grains usually tricolporate;
12 Flora Neotropica

1.
A :~ C S

I r
. .:'.it1

, .
wD. surface
i section;
focus; C optical' longitudinal . :H
r.. 'view os open; E, oblique~surface
equatorial
. . ':...:":
F'~
_ ":i ,. . v"
:!i ;
i
.._
_!I
G _ : ,

FIG 6. Pollen grains of Lophophytum; A-C, F-I all LM x .000 D SEM x 1365; E, SEM x 2300;
Balanophoraceae 13

however, in some collections 5-25% of the grains are hexacolporate.

Tricolporate grains (Fig 7A, B, D, E, G): (spheroid-)subprolate-
-prolate(-perprolate) with polar axis P: 32-36,m and equatorial diam. E:
20-26 /m; P/E: (1.06-)1.2-1.8(-2.58). Apocolpium diam. 6-8 /im. Colpi up to
1.5 /m wide; os indistinct, about 7 x 5 /m; colpus membranes with nanover-
rucae about 0.1 M/mdiam. Exine 0.8-0.9 Mmthick, at mesocolpi smooth or
faintly reticulate with lumina about 0.05 /m diam., at apocolpi rugulate.
Hexacolporate grains (Fig 7C, F): asymmetrical tetrahedrons with 4 polar
regions connected by 6 colpi of 3 different lengths, eg 24.5 /m, 26.5 /m, and
29 /m, opposite colpi always of the same length and forming angles to each
other less than 90?, typically about 70?-80?; longest axes (27-)29.5(-34) mim.
Exine patterns as in tricolporate grains. Material studied: Curran 22 (Brazil),
Glaziou sn (Brazil), Krapovickas 1687 (Argentina), Lourteig 690 (Argentina),
Weddell sn (Bolivia).
Lathrophytum peckoltii (Fig 8A-D). Pollen grains spheroid-suboblate
with polar axis P: 27.1 /m and equatorial diam. E: 31.4 /m; P/E: 0.86.
Apocolpium diam. 8-9 /m. Colpi up to 4.5 /m wide; os very distinct, almost
quadrangular, 7 x 7 /m; colpus membranes with nanoverrucae about 0.7 /m
wide. Exine about 1 M/mthick, at mesocolpi rugulate. Material studied:
Glaziou sn, 1886 (Brazil), Peckolt sn, 1867, 1886 (Brazil).
Ombrophytum violaceum (Fig 8E-I, 9A). Pollen grains spheroid with
polar axis P: 24-28 M/mand equatorial diam. E: 22-27 /m; P/E: 1.08.
Apocolpium diam. 7 /m. Colpi up to 3.5 /m wide; os indistinct; colpus mem-
branes with verrucae up to 1.2 /m wide. Exine 1.8 /m thick, at mesocolpi ver-
rucate, at apocolpi verrucate-rugulate; mesocolpi becoming almost smooth
towards colpi and apocolpia. Material studied: Mexia 6337a (Peru), Sparre
13049 (Ecuador), Ule sn, 1911 (Brazil), Wurdack 2360 (Peru).
Ombrophytum microlepis (Fig 8K-M.) Pollen grains tricolporate, occa-
sionally hexacolporate tetrahedrons, spheroid with polar axis P: 22.1 /m and
equatorial diam. E: 22.2 /m; P/E: 1.00. Apocolpium diam. 3.6-4.5 /m. Os
lalongate or almost tetrangular, 1.8-4.1 x3.6-6.8 M/m.Exine 0.9 M/mthick,
rugulate at apocolpi, almost smooth at mesocolpi. Material studied: Schunke
3943 (Peru), Schultes 8615 (Brazil).
Ombrophytum peruvianum (Fig 9B-D). Pollen grains tricolporate, occa-
sionally hexacolporate tetrahedrons, spheroid to prolate with polar axis P: 28
/m and equatorial diam. E: 18-27 /m; P/E: 1.04-1.59. Apocolpium diam. 6.5
/m. Os subcircular, 11 x7-8 /m. Exine 1 M/mthick, coarsely rugulate at
apocolpia, faintly reticulate to rugulate at mesocolpia with minute lumina less
than 0.1 /m. Material studied: Harling et al 7026 (Ecuador), Prance et al 7664
(Brazil).
Ombrophytum subterraneum (Fig 9B-K).
1. Tricolporate grains normally predominant (Fig 9B-H): spheroid with
polar axis P: 28-32 Am and equatorial diam. E: 28-30 mti; P/E: 1.01-1.07.

view in high focus; G, focus between apocolpium and equator; H, equatorial view in high focus; I,
optical longitudinal section; K, surface equatorial view showing lalongate os; L, slightly oblique
surface equatorial view showing loose, undulating sexine which adheres to nexine in central part of
mesocolpium, compare I left side.-A-E, Krapovickas 15509; F-L, Schunke 5675.
14 Flora Neotropica

A:

.. A... g.

@B

FIG 7. Pollen grains of Lophophytum and Langsdorffia; A-C, G-L all LM x 1000; D, SEM
x1390; E, SEM x 1420; F SEM x1660; M, SEM x1775.-
Lophophytum mirabile subsp mirabile; A, polar view in high focus; B, optical longitudinal sec-
 Balanophoraceae 15

o}~~~~~~~..~A 0

::
.....

K Ls
FIG 8. Pollen grains of Lathrophytum Ombrophytum; A-C, E-G, K, L all LM x 1000; D,
Lathrophytumnand Ombrophy/um;
SEM x 1225; H, SEM x 1425; I, SEM x 1400; M, SEM x 1990.-Lathrophkfum peckoltii; A,
1990.-Lathrophvtum pekoltii;
equatorial view in high focus showing colpus and os; B, optical longitudinal section; C, optical
cross section; D, slightly oblique surface view with triangular apocolpium upwards and equator at
lower
Loureig 690 H-L, Reinhard
rim.-Ombrophytum violaceum E
sn; M, Luezelburg
violaceum; sn. view in high focus; F, optical longitudinal
E, equatorial
section; G, optical cross section; H, surface polar view; 1, I, slightly oblique surface view with
apocolpium upwards.-Ombrophytum microlepis; K, optical longitudinal section; L, optical cross
section; M, oblique surface view with apocolpium downwards.-A-D, Peckolt sn 1886; E, G-l,
Sparre 13049; F, Wurdack2360; K-M, Schunke 3943.

tion; C, hexacolpate grain with three colpi focussed, compare F; D, surface polar view; E, surface
equatorial view; F, surface view of hexacolpate grain, slightly tilted over NW-SE-axis compared
with C.-Lophophytum mirabile subsp bolivianurn; G, optical longitudinal section.--Lan?s-
section.-Langs-
dorffia hypogaea; H, optical cross section; l,1, polar view in high focus; K, optical longitudinal sec-
tion; L, equatorial view in high focus; M, surface polar view, compare I.-A-E, .-A-E, Curran 22;22, G,
Lourteig 690; H-L, Reinhardt sn; M, Luetzelburg sn.
16 Flora Neotropica

Apocolpium diam. 10 im. Os subcircular to quadrangular, 4 /m diam.; colp

membrane with numerous nanoverrucae. Exine 1 I/m thick, almost smooth to
more or less verrucate to rugulate at mesocolpi and smooth or faintly rugulate
at apocolpi. Tetraloxocolporate spheroid grains and hexacolporate tetrahedral
grains have been observed. Material studied: Cabezas 20341 (Argentina),
Sleumer 3054 (Argentina), Viirsoo sn, 1956 (Argentina).
2. Pentadecacolpate grains (Fig 9I,K): spheroid to ellipsoid or quite ir-
regularly shaped, up to 54.4 x 62.2 /tm. Exine almost smooth. Material
studied: Cabezas 20263 (Argentina), Shepard 246 (Bolivia).
3. Galapagos population: form with apparently quite normally shaped an-
thers. The development of the pollen is arrested at an early stage, and no
pollen grains are formed. The empty anthers open as usual with longitudinal
slits. Material studied: Adsersen 243, 244, 1323, Coppois sn.

Balanophoroideae - Langsdorffia hypogaea (Fig. 7H-M).

Pollen grains isopolar, tetraporate, rarely pentaporate. Suboblate with

polar axis P: 22 /rm and equatorial diam. E: 27 trm;P/E: 0.82. Apertures
semicircular pores 5 t/m diam. Exine 1.5 ,/m thick with spines up to 0.8 /m high
and numerous very fine pores 0.1 /m diam. Spines confined to 4 of the 8 areas
delimitated by the equator and meridians through the apertures, always found
in diametrically opposite areas whether seen in polar view or in equatorial view
(Fig 71, L and M). NPC 444. Material studied: Luetzelburg sn (Brazil), Mosen
4404 (Brazil), Reinhardt sn (Brazil).

EMBRYOLOGY

Apart from Fagerlind's (1938a, 1945c) and Umiker's (1920) investigations

very little is known of the embryology of neotropical taxa. More literature is
available concerning palaeotropical taxa. However, much of the information
concerning Balanophora is contradictory according to Fagerlind (1945a), who
reviewed earlier investigations in detail.
No definite ovarial cavity is found in the members of this family, except in
Cynomorium, nor is a well defined placenta seen. Consequently in most
genera normal ovules are not developed, and the ovules may be regarded as
ategmic. Cynomorium has distinctly unitegmic ovules.
In Helosis a central, basal papilla of undifferentiated placental tissue

FIG 9. Pollen grains of Ombrophytum, A, B, E, F, I, K all LM x 1000; C, SEM x 1410; D, SEM

x 1380; G, SEM x 3525; H, SEM x 1410.-0. violaceum; A, oblique equatorial view in high
focus.-O. peruvianum; B, equatorial view focussed slightly above optical section, showing two
colpi, compare C; C and D, surface equatorial,views.-O. subterraneum; E, polar view in high
focus; F, optical cross section; G, oblique surface view showing colpus and os; H, surface polar
view, compareE; I, high focus of pentadecacolpate grainshowingseveralcolpi; K, opticalcross
sectionof pentadecacolpategrain,five uppercolpislightlyout of focus, fivecolpiperpendicular
to
the paperand five lowercolpicompletelyout of focus.-A, Mexia6337a;B-D, Harlinget al 7026;
E-H, Sleumer 3054; I, K, Cabezas SI 20263.
Balanophoraceac 17

ffi *

: I .t

?;=- ~~~~~-

I_~ iW
X,, ?,,, .
18 Flora Neotropica

develops. Eventually the outer cells of the papilla seem to fuse with the inner
cells of the carpellary tissue, and no cavity can be demonstrated. Fagerlind
(1938a) found two megaspore mother cells in the papilla. They both enter
meiosis, and thus two dyads are formed. The upper cell in each dyad
degenerates and very soon also the lower cell in one of the dyads. The lower
cell in the second dyad quickly becomes 8-nucleate through three subsequent
divisions. The embryo sac development is thus bisporic and of the Allium-
type. The four upper nuclei form the egg-cell, two synergids and one polar
nucleus. The four lower, chalazal nuclei form 1-3 antipodal cells and 3-1 free
nuclei, which eventually fuse with the upper polar nucleus to become the cen-
tral nucleus. By this stage the hexagonal bracts, covering the inflorescence, are
shed. Now follows the fertilization and the formation of a cellular endosperm,
and finally a few-celled embryo develops. Fagerlind (1938b) demonstrated
much the same development in Ditepalanthus from Madagascar. Judging
from illustrations this development is likely to be found also in Scybalium
(Eichler 1869) and Rhopalocnemis (Lotsy 1901). It may thus be the rule in sub-
family Scybalioideae, but further research is needed to confirm this point.
Within subfamily Lophophytoideae much less is known. Eichler (1867)
investigated Lophophytum mirabile. The development of the female flower
shows obvious similarities to that of Helosis. A central, basal papilla, pro-
bably again an undifferentiated placenta, is formed. Also here a complete fu-
sion between placenta tissues and carpel walls takes place at later stages. The
further interpretation of Eichler's investigations is difficult. However, his il-
lustrations clearly show that two megaspore mother cells are developed in the
central papilla and that finally one embryo is developed, surrounded at
maturity by layers of stone cells. In Lathrophytum peckoltii only one embryo
is developed in the ovary as is shown in the illustrations of Eichler (1868,
1869). Poeppig & Endlicher (1838) described the ovary of Ombrophytum peru-
vianum as uniovular, possibly referring to the few-celled embryo. Asplund
(1928) and Sleumer (1954) found in Ombrophytum subterraneum an ovary
with the placental tissues at later stages completely adnate to the walls and only
one embryo developed. Further investigations are needed to throw light upon
details in the embryology of the Lophophytoideae, particularly the early stages
and the details of embryo sac development. However, even with the present
knowledge apparent similarities to the development within Scybalioideae may
be inferred.
Within subfamily Balanophoroideae Fagerlind (1945a, b and c) described
the embryology of Balanophora elongata and Langsdorffia hypogaea in
detail, reviewing earlier investigations. In Balanophora elongata he described
the development of the extremely undifferentiated ovary without cavity and
placenta (1945b). The centrally placed megaspore mother cell, however, has a
normal tetrad formation (1945a). The apical spore develops into an 8-nucleate
embryo sac. Already at the 2-nucleate stage a caecum develops from the basal
part of the embryo sac and grows quickly upwards, containing the basal
nucleus, eventually surpassing the morphological apex of the embryo sac. The
two nuclei each divide twice, and each end of the embryo sac thus contains a
group of 4 nuclei. Those at the morphological apex degenerate, while those at
the caecum develop into 1 egg cell, 2 synergids and 1 polar nucleus. The em-
Balanophoraceae 19

bryo sac is thus monosporic, developed from the apical spore, and subsequent-
ly 8-nucleate. A special feature is the development of the basal caecum, in
which the formation of the 8-celled embryo surrounded by a few-celled en-
dosperm takes place (Zweifel 1939).
Also in Langsdorffia hypogaea, Fagerlind (1945c) showed a monosporic
embryo sac development from the apical spore of the tetrad. At the 4-nucleate
stage a caecum is formed subbasally, growing quickly upwards as in
Balanophora elongata, containing the two basal nuclei. The further develop-
ment proceeds as in Balanophora elongata. Geesink (1972) described and il-
lustrated exactly the same kind of embryo sac in Langsdorffia papuana from
New Guinea.

POLLINATION, DISPERSAL AND GERMINATION

Knuth (1904) reported that pollination is effected by small Diptera in

Balanophora elongata and another species. Also in Lophophytum and Sar-
cophyte visiting Diptera were observed. Zweifel (1939) observed Diptera
visiting Balanophora abbreviata, and the present author (Hansen, 1972) found
small Hymenoptera busy on male inflorescences of Balanophora fungosa
subsp. indica. Visits of Diptera to Dactylanthus taylorii were reported by
Moore (1940). Govindappa & Shivamurthy (1975) observed pollination in
Balanophora abbreviata by various species of small Hymenoptera. Insect
pollination is highly probable in many species of the Balanophoraceae, the
pollinators being attracted by sugary secretions in the inflorescences, as
observed in Rhopalocnemis by van Steenis (1932).
Pollen tubes have been observed penetrating the stylar tissues in many
species. In Balanophora abbreviata, Zweifel (1939) observed pollen tubes
entering the embryo sac, and he also found obvious cases of double fertiliza-
tion. Fagerlind (1938a) demonstrated a quite normal process of fertilization in
Helosis contrary to the statements by Umiker (1920). In fact apomixis has been
proved only in Balanophora globosa and B. japonica, although it is highly
probable in Ombrophytum subterraneum, especially in populations destitute
of pollen or with abnormal pollen grains (p. 16).
In most genera, including all neotropical ones, the fruits can be described
as small achenes with one to a few layers of stone cells surrounding the em-
bryo. However, in Balanophora the fruit wall finally becomes 2-layered. All
walls of the inner layer and the inner, tangential walls and radial walls of the
outer layer are strongly lignified and cutinized as the cytoplasm disappears.
Thus the fruit of Balanophora becomes a nut with a very rough surface.
Little is known about the dispersal of fruits in the Balanophoraceae. In
Balanophora the fruits are about the same size and weight as orchid seeds, and
dispersal by wind is a possibility. Their rough fruit wall makes them easily at-
tachable to the bodies of visiting insects, but this remains to be observed in the
field. Also rainwash must be considered. In South American taxa the fruits are
much bigger, and wind dispersal is unlikely. However, high contents of starch
in the tissues of most species may attract herbivores, whereby dispersal could
be effected. The dispersal of Ombrophytum subterraneum to the Galapagos
20 Flora Neotropica

Islands is not easily explained, unless migrating birds were involved.

Govindappa & Shivamurthy (1976) described the germination of fruits in
Balanophora abbreviata as follows. Fruits, having lost the style, are detached
from the infructescence and dispersed. Once settled on the soil surface they are
brought down to the rhizosphere of the host probably by means of percolating
rainwater or by digging animals. Having absorbed water the fruit germinates
and breaks open at the proximal end. The endosperm cells closest to the rup-
ture elongate and send out 4-8 narrow, tubular and apparently sticky exten-
sions. The sticky endosperm tubules anchor the fruit to a young host rootlet.
The few celled embryo sends out 1-4 thick tubular processes containing a dense
cytoplasm and comparatively large nuclei, the primary haustorium.
Simultaneously the upper embryo cells through a series of divisions produce a
whitish nodule of parasite parenchyma. The growth of the primary
haustorium into the host rootlet right through to the vascular tissue induces
the formation and growth in the opposite direction of a lateral branch of host
root vascular tissue. This tissue embeds the primary haustorium and enters the
nodule of parasite parenchyma, where it at later stages will branch
dichotomously. At the apical zone of contact between this tissue and the
parasite parenchyma, the formation of a parasite meristem takes place; lateral-
ly a host root meristem is formed. The parasite meristem is responsible for the
formation of long chains of large-celled secondary haustoria. The host root
meristem forms the tissue of the host surrounding the secondary haustoria of
the parasite. It is supposed that a flow of material in this way is ensured from
the vascular tissues of the host root to the haustorial chains of the parasite.
Later stages of this development were described by Fagerlind (1948).

ANATOMY

The tubers of the Balanophoraceae are to a varying degree dual structures

containing host root tissue as well as parasite tissue. In the genera Helosis
(Eichler 1869, Umiker 1920), Scybalium and Lophophytum (Eichler 1869),
and Dactylanthus (Moore 1940) the host root tissue meets the parasite tissue in
the peripheral layers of the tuber along a rather simple surface of connection.
A direct connection of host xylem and parasite xylem has been demonstrated
in Helosis by Umiker (1920). A highly complicated connection has been
demonstrated in Thonningia by Mangenot (1947) and in Balanophora by
various authors, see Fagerlind (1948). In Balanophora and Thonningia there is
no direct connection between host root bundles and parasite bundles.
Transportation of water and nutrients between the two bundle systems is sup-
posed to take place in the haustorial cells sensu Govindappa & Shivamurthy
(1976), see above, ie "chain cells or column cells" sensu Fagerlind (1948),
"Heinricher cells" sensu Mangenot (1947), "thallus cells" sensu Heinricher
(1907), "vesicular cells" sensu Solms Laubach (1867-1868). It should be
remembered, however, that the parasite part of the tuber functions as a root
for the parasite and also that several anatomical details point to interpreting it
as root. Due to their dual construction these organs will be referred to as
tubers in the following systematic treatment. Judging from Eichler's (1869) il-
Balanophoraceae 21

lustrations the connection between host and parasite is quite complicated also
in Langsdorffia where the underground tubers are much elongated and
rhizome-like. They consist of a parenchymatous tissue and 20-30 collateral
bundles. The vessels are reticulate or striate and have scalariform perforations.
The sieve tubes have perforated sieve plates.
In Helosis and Scybalium jamaicense subterranean, horizontal, rhizome-
like structures arise from the "primary" tuber. Their anatomy in Helosis was
described by Eichler (1869) and Umiker (1920). The parenchymatic cortex con-
tains numerous groups of stone cells. Centrally a single ring of 4-10 collateral
vascular bundles intermixed with medullary bundles is found. The vascular
bundles are accompanied by layers of sclerenchymatous cells.
The scaly leaves are destitute of stomata except in Cynomorium. Three
vascular bundles are found basally in the leaves of Langsdorffia hypogaea and
Lophophytum mirabile and one bundle in Scybalium fungiforme.
The inflorescence bearing stems in Helosis contain 12-20 vascular bundles
in two basal rings; the number is greater above by ramification, and the ar-
rangement is irregular; profuse branching occurs in the inflorescence. In
Scybalium fungiforme and Lophophytum mirabile up to 300 bundles are
found in the stems (Eichler 1869).

CHROMOSOMES

Chromosome counts in Balanophora were reviewed by Hansen (1972). In

B. abbreviata, B. elongata and B. fungosa subsp. indica n= ca. 16 or ca. 18
and 2n = ca. 36 had been reported. The obviously apomictic B. japonica had
2n = 56 or 94-112. Reports on B. dioica were due to misidentification. In
Helosis Fagerlind (1938a) established n= ca. 18. Mangenot (1947) found in
Thonningia n= ca. 18. Much more work with more reliable counts is needed.

HOST PLANTS

In Table 1 the host plants so far recorded for neotropical Balano-

phoraceae are listed. In 6 of 18 taxa no host is yet on record. The remaining 12
taxa parasitize 32 host species in 15 families. Two large families, Compositae
and Leguminosae are represented with 8 and 10 host species respectively. Om-
brophytum subterraneum parasitizes 10 different hosts in 4 families.
Langsdorffia hypogaea parasitizes 6 hosts in 5 families. There is nothing in
this material to support the idea that the Balanophoraceae are specific in their
choice of host. On the contrary, the picture is much the same as in
Balanophora (Hansen, 1972) which parasitizes the largest systematic groups
most often. However, much more information concerning the hosts of
neotropical Balanophoraceae is needed, and collectors are strongly urged to
trace and collect the host by following the host root whenever possible.
 22 Flora Neotropica

Table 1. Host plants so far recorded of the following neotropical Balanophoraceae.- 1. Corynaea crassa
var sprucei.- 2. Helosis cayennensis var cayennensis.- 3. H.c. var mexicana.- 4. Langsdorffia
hypogaea.- 5. Lathrophytum peckoltii.- 6. Lophophytum leandri.- 7. L.mirabile ssp mirobile.- 8. L.m.
ssp boliviona.- 9. Ombrophytum subterraneum.- lo. Ombrophytum violaceum.- 11. Scybollum glaziovil.-
12. S. jamaicense.

1 2 3 4 5 6 7 8 9 lo 11 12
Arecac.: Geonoma sp. 4
- : Iriarteo sp. 4
Bignon.: unidentified taxon lo
Compos.: Baccharis petiolata DC. 9
- Eupatorium angulare B.L.Robinson 1
- - bupleurifolium DC. 9
- Heterothalamus spartioides Hook. & Am. 9
- Lepidophyllum quadrangulare (Meyen)Benth. & Hook. 9
- Scalesia pedunculota Hook.f. 9
- Tessaria absinthioides DC. 9
- Vguieria mollis Gris. 9
Dioscor.: Dioscorea megalantha Gris. 9
Euphorb.: Senefeldera sp. 5
Legum.: Anadenanthera macrocarpa (Benth.) Brenon 6
- Apuleia sp. 6
- Cassia emorginata L. 12
- Enterolobium sp. 8
- Inga sp. 2 7
- Medicago sativa L. 9
- Mimosa sp. 4
Piptadenia rigida Benth. 6
- p. 8
Pithecolobium sp. 8
Molpigh.: Byrsonima sp. 4
Meelastom.: unidentified taxon 11
Meliac.: Trichilia sp. 4
Moroc.: ticus sp. 3 4
Myrist.: Myristica bicuhyba Schott 5
Myrsir .: unidentified taxon 11
Sapot.; Achras sapota L. 3
Solon.: Nicotiana glauca Grah. 9
Urticac.: Boehmeria sp. 2
Number of hosts per taxon of parasites 1 2 2 6 2 3 1 3 lo 1 2

SYSTEMATICTREATMENT

BALANOPHORACEAERichard,Mem. Mus. Hist. Nat. Paris 8: 429. 1822

nom. cons. ("Balanophoreae");Eichler in DC., Prod. 17: 117. 1873;
Engler, Nat. Pflanzenf. ed. 1. 3. 1: 243. 1889;Tieghem,Ann. Sci. Nat.
Bot. Ser. 9. 6: 141. 1907;Fawcett& Rendle, Fl. Jamaica3: 103. 1914;
Britton& Wilson, Bot. Porto Rico and VirginIsl. 260. 1924;Lanjouwin
Pulle, Fl. Suriname1. 1: 45. 1932; Harms, Nat. Pflanzenf. ed. 2. 16b:
296. 1935;Macbride,Fl. Peru2: 427. 1936;Standley,Fl. Costa Rica409.
1937;Standley& Steyermark,Fl. Guatemala4: 92. 1946;Le6n & Alain,
Fl. Cuba 2: 82. 1951;Hutchinson,Fam. Fl. P1.ed. 2. 1: 340. 1959;Ange-
ly, Fl. Anal. e Fitog. Sao Paulo 1: 55. 1969;Falclo, Fl. II. Cat. BALA 3.
1975.

Herbaceous,fleshy plants destituteof chlorophyll,parasitizingroots of

treesand shrubs,rarelyherbs,with yellowish-whiteto yellow, orangeto redor
browncolors. Developingby germinationon a suitablehost root a branched
Balanophoraceae 23

or unbranched, variously shaped tuber. One or more supraterranean or partly

subterranean, inflorescence bearing stems endogenously arising from a tuber.
Stems with or without leaves, unbranched. Leaves scaly, without stomata,
spirally arranged. Inflorescence once branched or apparently spadix-like, the
branches being much suppressed. Branches subtended by scaly, ? transform-
ed and reduced, caducous bracts, sometimes with sterile apical part peltately
widened. Bracts scaly, triangular or peltate with angularly ovate to hexagonal
or almost circular pelta or reduced and clavate. Flowers unisexual. Male
flowers 3-merous, with a 3-lobed perianth and a synandrium composed of 3
anthers free from each other. Synandria opening irregularly, anthers bilocular,
rarely trilocular, opening by longitudinal slits. Female flowers with a much
reduced, 2-lobed or irregularly lobed perianth, and then with ovary inferior, or
perianth apparently absent. Ovaries without definite locules, placentas and
ovules. Styles 2, alternate with the perianth lobes, or 1; stigma slightly
capitellate. Generally one few-celled embryo develops in the central tissues of
the ovary, surrounded by an endosperm and a layer of stone cells at maturity.
Fruit a small, 1 seeded achene.
A mainly tropical or subtropical family of 43 species in 18 genera
distributed throughout both hemispheres from sea level to 4000 m alt. and
from tropical evergreen forest to dry bushlands and even to deserts.
Type genus. Balanophora J. R. & G. Forst. The name is derived from the
Greek words 3coXcuvos and cpEQomeaning "acorn bearer."

Key to the Genera of Balanophoraceae in America

1. Styles 2; starch in tubers and elsewhere; flowers of both sexes embedded in filiform hairs
or anthers 2, not merged into a synandrium.
2. Flowers embedded in a layer of filiform hairs; anthers merged into a usually 3-merous
synandrium. subfam. Scybalioideae.
3. Young inflorescences covered by more or less triangular scales; stems densely covered
by the same kind of scales. tribe Scybalieae.
1. Scybalium.
3. Young inflorescences covered by hexagonal, peltate bracts; stems naked or at most
with a few much reduced, inconspicuous bracts inserted at the same level.
tribe Helosieae.
4. Inflorescences emerging from buds on elongated, rhizome-like structures, which
again appear from a tuber connected with the host root. 2. Helosis.
4. Inflorescences appear directly from the tuber connected with the host root.
3. Corynaea.
2. Flowers not embedded in hairs, on conspicuous, + elongated branches, which are sub-
tended by often early deciduous ? peltate bracts. subfam. Lophophytoideae.
5. Branches subtended by bracts with angularly-ovate pelta; apical part of female branch
not peltately enlarged. 4. Lophophytum.
5. Branches subtended by bracts with ? hexagonal or more irregularly shaped pelta; api-
cal part of female branch peltately enlarged, ? covering the flowers.
6. Male flowers solitary, each flower subtended by a peltate bract; anthers sessile.
5. Lathrophytum.
6. Male flowers in many-flowered branches, which are each subtended by a peltate or
clavate bract; pelta sometimes 4 reduced; anthers with a conspicuous filament.
6. Ombrophytum.
1. Style 1; wax in tubers and elsewhere; flowers not embedded in filiform hairs; anthers 3,
merged into a synandrium. Subfam. Balanophoroideae.
7. Langsdorffia.
24 Flora Neotropica

Balanophoraceae subfam. Scybalioideae Engler, Nat. Pflanzenf. ed. 1. 3. 1:

253. 1889 p.p. excl. Lophophyteae.

Helosidoideae Harms, Nat. Pflanzenf. ed. 2. 16b: 311. 1935; Angely, Fl. Anal. e Fitog. Sao
Paulo 1: 55. 1969; nom. illeg. superfl.

Inflorescence with much depressed, hardly discernible branches, each

subtended by a triangular or peltate bract. Flowers embedded in a dense layer
of filiform, narrowly clavate hairs. Female flowers obviously compressed.
Type genus. Scybalium Schott & Endlicher, Melet. Bot. 3. 1832.
Within Scybalioideae two tribes are recognized in this treatment viz.
Scybalieae and Helosieae. They are beautifully linked together through the
rare Andean species, Scybalium depressum, which combines Scybalieae tribal
characters with such striking Helosieae characters as 3-nucleate pollen grains
and the compressed ovaries with their edges pointing towards the petioles of
the bracts.

Tribus Scybalieae Eichler, Actes Congr. Bot. Paris 152. 1867; in Martius, Fl.
Bras. 4. 2: 7. 1869; emend. Eichler in DC., Prod. 17: 131. 1873; Engler,
Nat. Pflanzenf. ed. 1. 3. 1: 256. 1889; Angely, Fl. Anal. e Fitog. Sao
Paulo 1: 56. 1969.
Stem with triangular, scaly leaves. Inflorescences when young covered by
triangular, imbricate bracts. Pollen grains 6-polypantoporate.
Type genus. Scybalium Schott & Endlicher, Melet. Bot. 3. 1832.

1. Scybalium Schott & Endlicher, Melet. Bot. 3. 1832; Endlicher, Gen. Plant.
74. 1836; Griffith, Trans. Linn. Soc. London 20: 104. 1846; Hooker f.,
Trans. Linn. Soc. London 22: 31, 50. 1856; Eichler in Martius, Fl. Bras.
4. 2: 35. 1869; in DC., Prod. 17: 131. 1873; Hooker f. in Bentham &
Hooker f., Gen. P1. 3: 236. 1880; Engler, Nat. Pflanzenf. ed. 1. 3. 1: 256.
1889; Fawcett & Rendle, Fl. Jamaica 3: 103. 1914; Harms, Nat.
Pflanzenf. ed. 2. 16b: 318. 1935; Leon & Alain, Fl. Cuba 2: 83. 1951;
Angely, Fl. Anal. e Fitog. Sao Paulo 1: 56. 1969; Falcao, Atas Soc. Biol.
Rio de Janeiro 14: 154. 1971; Adams, Flow. PI. Jamaica 244. 1972;
Falcao, Loefgrenia 57: 1. 1973; Fl. II. Cat. BALA 18. 1975.
Phyllocoryne Hooker f., Trans. Linn. Soc. London 22: 31, 51. 1856; Grisebach, Fl. Brit. W.
Ind. 309. 1860.
Sphaerorhizon Hooker f., Trans Linn. Soc. London 22: 31, 50. 1856.

Fleshy root parasites. Stems cylindrical or obconical, appearing from

basal, swollen, starchy tubers or horizontal rhizome-like structures. Leaves
scaly, densely imbricate, triangular, more or less petiolate. Inflorescences ap-
parently not branched, depressed-discoid to ovoid or almost cylindrical, at
first covered by imbricate, scaly bracts. Bracts triangular, more or less
petiolate, peltate or not, deciduous. Each bract subtending an extremely
depressed, unisexual or bisexual branch. Flowers embedded in a dense layer of
filiform, narrowly clavate hairs. Male flowers with a tubular, 2-3 lobed
perianth. Filaments connate into a column, upwards often splitting into
separate filaments and then again joining into a synandrium longitudinally
Balanophoraceae 25

(4-)6(-9)-celled. Pollen grains spheroid to slightly ellipsoid, 6-polypanto-

pororate, medium-sized. Female flowers more or less compressed, with
perianth adnate to ovary, lobes 2, short and broad. Styles 2, filiform, stigmas
capitellate. Fruit a small, I-seeded achene.
Type species. Scybalium fungiforme Schott & Endlicher, Melet. Bot. 3, t.
2. 1832.
Distribution. Greater Antilles, Colombia and Ecuador, SE Brazil.
Schott & Endlicher (1832) described the genus Scybalium based on their
Scybalium fungiforme. Apparently with some doubt they transferred
Cynomorium jamaicense Swartz to their new genus. Hooker f. (1856) propos-
ed two genera close to Scybalium each with one species Sphaerorhizon
depressum, a new species, and Phyllocoryne jamaicensis for the transfer of
Swartz' species. Eichler (1873) united the three entities mentioned above in
Scybalium together with his new species Scybalium glaziovii. Eichler's
delimitation is followed here and leaves us with an easily defined genus
characterized by the rather stout stems and inflorescences being covered by
densely imbricate, triangular, scaly leaves and bracts, and by the ellipsoid-
spheroid, 6-polypantopororate pollen-grains. The clavate hairs on the in-
florescence and the compressed female flowers are shared in common with the
close relatives Helosis and Corynaea. The peculiar affinity of Scybalium
depressum to these two genera has been mentioned earlier. The other three
species show affinity to Lophophytoideae in their leaf morphology, particular-
ly Scybalium fungiforme, and in the arrangement of the female flowers on the
secondary axes. It should be pointed out that Hooker's generic concept would
place Scybalium glaziovii in Phyllocoryne.

Key to the Species of Scybalium

1. Bracts of inflorescence obviously peltate with narrowly angularly-ovate pelta; female flowers
compressed, arranged with edge towards petiole of bract; male flowers (2-)3-merous; heads
bisexual. 1. S. depressum.
1. Bracts of inflorescence not obviously peltate; female flowers ? compressed, arranged with
flat side facing petiole of bract; male flowers 3-merous; heads bisexual or unisexual.
2. Stem strongly obconical, terminating in a depressed, disk-like, unisexual inflorescence;
bracts of inflorescence subpeltate with flattened-canaliculate petiole 5-8 mm long and 2.5-
5 mm broad. 2. S. fungiforme.
2. Stem narrowly cylindrical, terminating in an ellipsoid to long-ellipsoid inflorescence;
bracts of inflorescence basally with very broad and short, extremely flattened-canaliculate
petiole 3 mm long and 7-16 mm wide.
3. Bracts of inflorescence arranged in ca. 6 vertical ranks. 3. S. jamaicense.
3. Bracts of inflorescence arranged in 14-20 vertical ranks. 4. S. glaziovii.

1. Scybalium depressum (Hooker f.) Eichler in de Candolle., Prod. 17: 133.

1873; Engler, Nat. Pflanzenf. ed. 1. 3. 1: 257. 1889; Harms, Nat.
Pflanzenf. ed. 2. 16b: 319. 1935. Fig 10, Fig 3 (pollen), Fig 12 (Distr.)

Sphaerorhizon depressum Hooker f., Trans. Linn. Soc. London 22: 65, t. 10. 1856.
Sphaerorhizon curvatum Hooker f., Trans. Linn. Soc. London 22: 31, 50. 1856.

Description based on wet and dried specimens. Monoecious (inflorescence

26 Flora Neotropica

a
C"
W a? y. id ,,,,,I

FIG 10. Species of Scybalium. A-E, S. depres.sum, A, after Hooker f. (1856), B, C (Holm-Niel.sen
el a! 1267), D (SSteyermrk 52634), E, (fosberg et Core 21518) original. A, habit x0.5; B, bract
subtending inflorescence branch, front view of peltate part x 3; C, the same, side view (compare
Fig 1, 4B and 4C) x 3; D, dimerous male flower with perianth segments irregularly incised at apex
x6; E, female flower x7. F-M, S. fungiforme, F-L, after Eichler (1869), M (de Lima 238)
original. F, typical habit with one male inflorescence surrounded by several female ones, all ap-
pearing from the same tuber x0.5; G, bract subtending inflorescence branch, front view x 1; H,
the same, side view x 1; 1, section of male inflorescence with two buds and the two kinds of hairs
x4; K, male flower x4.5; L, longitudinal section of male flower with ovoid-
conical body basally in the staminal tube x 4; M, female flower x 7.

bisexual) brown to rose-purple root parasites. Total length from point of con-
tact with host root (4-)6-12(-15) cm. Tuber subspherical 0.8-4(-7) cm diam.,
single or a few fused together, surface finely granular, often with a few conical
appendages 2.25 mm long and 1.5 mm diam. at base. Stem cylindrical, 1.3-8
cm long, 0.8-1 cm wide. Leaves scaly, numerous, apparently spirally arranged,
at base of stem sessile, long triangular, obtuse at apex, up to 17 x 5 mm, up-
wards becoming more or less peltate, pelta narrowly angularly-ovate. In-
florescence ovoid-obovoid, 1.8-3.0(-3.8) x 1.2-2.3(-2.7) cm, densely covered
by bracts when young. Bracts peltate with a terete petiole 0.5-1.5 mm across
Balanophoraceac 27

and 1.5 mm long, pelta narrowly angularly-ovate, 12-13 x4-5 mm, apparently
spirally arranged and densely imbricate, early deciduous and then for a short
time leaving a circular cavity in the dense layer of filiform hairs covering the
inflorescence, cavity later on closing completely. The rhombic, slightly
transversally extended area delimitated by imaginary lines connecting four
neighboring petioles represents one extremely depressed branch subtended by
the lowermost of the four bracts in question. Each branch with a slightly curv-
ed central depression under which 10-15 male flowers are embedded in the
layer of hairs, surrounded by 40-60 female flowers; strongly proterogynous.
Hairs thinly clavate, 1.4-1.5 mm long. Male flowers with a 2-3-lobed tubular
perianth, by rapid growth in the tubular part appearing above layer of hairs
during anthesis, and then with tube 1.75 x 0.4 mm and tepals ovate, 2.5 x 1.5
mm. Filaments connate into a column 2.5 x0.3 mm, widening to 0.6 mm
diam. just below synandrium. Synandrium ellipsoid, 0.9x0.75 mm,
longitudinally 4-9 celled. Pollen grains spheroid-ellipsoid, polypantopororate,
26.2 Mtm diam., with extremely thin exine 0.5-0.7 Mtm,nuclei 3. Female flowers
with perianth adnate to ovary, 2 short and broad, lip-like segments protruding
above ovary, compressed with edge generally facing petioles of bracts, 1.5 mm
long from base to tip of segments, 0.4 mm wide; styles 2, filiform, 1.5 mm
long, appearing above layer of hairs; stigmas capitellate. Fruit a small,
1-seeded achene.
Type. Purdie sn July 1846, Colombia, between Niva and Paramo de Ruiz
(holotype, K).
Distribution. In montane forests 1500-3000 m, apparently flowering all
year round. Host not known.

COLOMBIA. Ewan 16075 (US); Little 7650(US); Pittier 1531 (US); Fosberg et al 21518 (US).
ECUADOR. Acosta Solis 5837 (F); Holm-Nielsen et al 1262 (AAU); Steyermark 52634 (NY).

By an error Hooker created different names when he published this

species: ie S. curvatum which accompanied the description, and S. depressum
with the plate, both names were thus validly published. However, on the
transfer to Scybalium Eichler chose S. depressum.

2. Scybalium fungiforme Schott & Endlicher, Melet. Bot. 3, t. 2. 1832; Hooker

f., Trans. Linn. Soc. London 22: 31, 50. 1856; Eichler in Martius, Fl.
Bras. 4. 2: 36, t. 7, 8. 1869; in DC., Prod. 17: 132. 1873; Engler, Nat.
Pflanzenf. ed. 1. 3. 1: 257,f. 162. 1889; Glaziou, Mem. Soc. Bot. France
1(3): 610. 1913; Harms, Nat. Pflanzenf., ed. 2. 16b: 319, f. 160. 1935;
Angely, Fl. Anal. e Fitog. Sao Paulo 1: 56. 1969; Falcao, Atas Soc. Biol.
Rio de Janeiro 14: 154. 1971. Fig 10, Fig 3 (pollen), Fig 12 (distr.).

Description based on dry material and on the excellent description by

Schott & Endlicher. Monoecious, dark brown parasites with unisexual in-
florescences, ususally 1 male centrally and 4-10 female ones from each tuber.
Tuber subspherical, more or less lobed, 5-7 cm diam., surface verrucose.
Stems appearing from lobes of tuber, 5-10 cm long, obconical and at base 1.5
cm diam. increasing upwards to 4-7 cm diam. Leaves scaly, densely imbricate,
28 Flora Neotropica

apparently spirally arranged, in lower part of stem lanceolate, subentire,

becoming rhomboid to ligulate above, 9 x 4.5 mm, in uppermost part of stem
broad triangular with margin lacerate. Inflorescence depressed into a slightly
convex, disciform structure 4-7(-9) cm diam., first covered by imbricate, scaly,
apparently spirally arranged bracts. Bracts subpeltate with petiole flattened-
canaliculate, 5-8 mm long and 2.5-5 mm wide, pelta subcircular to rhombic,
6-7.5 mm long and 6-9 mm wide, border fringed except in lower subtriangular
part, early deciduous and then leaving a horseshoe-shaped scar in the dense
layer of filiform, clavate hairs covering the inflorescence; scars soon closing by
the hairs being pushed by growing flowers. The area immediately centripetal to
(ie above) each bract represents one extremely depressed branch subtended by
that bract. Hairs in male inflorescences of two kinds, thinly clavate hairs
1.6-2.6 mm long and very densely arranged, and long subulate hairs 6.5-8 mm
long and more loosely arranged (abortive carpels ?); in female inflorescences
only the shorter, clavate hairs are found. Male flowers at first buried in the
layer of clavate hairs, later growing to 5.8-6.5 mm, with a lower tubular part
and a tripartite perianth, segments ovate, concave, 1.3x0.8 mm. Filaments
connate into a tubular column 4-4.5 x 0.25-0.5 mm. Synandrium ellipsoid to
subspherical, 0.8-1.2 x0.8-1 mm, composed of 3 anthers, each with 2 cells.
Rudimentary ovary (?) protruding as a short ovoid-conical structure about 1
mm long into lower part of perianth-tube. Pollen grains spheroid,
6-pantopororate, 54 ~m diam., exine 2.1 /im, nuclei 2. Female flowers with
perianth adnate to ovary and two short and broad lip-like segments protruding
above ovary. Ovary slightly compressed, elliptic, 1 mm by 0.5-0.6 mm; styles
2, 3.5-4 mm long, appearing above layer of hairs; stigmas capitellate. Fruit a
small, 1-seeded achene.
Type. Schott sn December 1820, Brazil, Rio de Janeiro, Sierra d'Estrella
(holotype, W).
Distribution. In dense evergreen forests up to 1500 m alt. Records from
June-August, October and December. Host not known.

BRAZIL. Glaziou 4070a (P); Goias: Anderson et al 7412 (NY); Minas Gerais: Regnell sn
(BR); Lemos sn (SP); Sao Paulo: Eiten, Eiten et al 3094 (K); Lima 238 (SP); Pickel 3423 (R); Mello
sn (K); Gehrt sn (SP).

3. Scybalium jamaicense (Swartz) Schott & Endlicher, Melet. Bot. 12. 1832;
Eichler in DC., Prod. 17: 133. 1873; Engler, Nat. Pflanzenf. ed. 1. 3. 1:
257. 1889; Urban, Symb. Ant. 4: 210. 1905; Fawcett & Rendle, Fl.
Jamaica 3: 104, pl. 5. 1914; Urban, Symb. Ant. 8: 190. 1920; Harms, Nat.
Pflanzenf. ed. 2. 16b: 319. 1935; Le6n & Alain, Fl. Cuba 2: 83, f. 31.
1951; Adams, Flow. PI. Jamaica 244. 1972.
Fig 11, Fig 3 (pollen), Fig 12 (distr.).

Cynomorium jamaicense Swartz, Nov. Gen. & Sp. P1. 12. 1788; Fl. Ind., Occ. 1: 1797.
Helosis jamaicensis (Swartz) Richard, Mem. Mus. Hist. Nat. 8: 432. 1822.
Phyllocoryne jamaicensis (Swartz) Hooker f., Trans. Linn. Soc. London 22: 31, 51, t. 11.
1856; Grisebach, Fl. Brit. W. Ind. 309. 1860; Mem. Acad. Amer. Scient. & Artium, N.
Ser. 8: 191. 1860; Cat. PI. Cub. 118. 1866.
Cynomorium coccineum auct. non Linn.; Descourtils, Fl. Med. Antill. 2: 113, t. 96. 1822.
Balanophoraceae 29

Description based on wet and dry specimens. Dioecious (inflorescence

unisexual), red-orange to pink or dark red root parasites. Total length from
point of contact with host root (4-)9-11(-23) cm. Tuber subspherical, 1-5 cm
diam., giving out short, cylindrical, horizontal rhizome-like structures about 1
cm diam. Stem appearing at intervals from rhizome-like organs or sometimes
apparently directly from the tuber, cylindrical, (2-)5-8(-18) cm long and
0.7-1.5 cm wide. Leaves scaly, numerous in 6 vertical ranks, first densely im-
bricate, almost triangular, acute (6-)7-9(-11) mm long and (5-)7-10 mm wide at
base, erect. Inflorescence ellipsoid to cylindrical-ellipsoid, 2-7x1-2.5 cm,
densely covered by imbricate, 6-ranked bracts when young. Bracts transversely
triangular, acute, 6-12 x 7-16 mm at base, inserted by a broad-canaliculate 3
mm long petiole, which is transversally extended and attached along the entire
lower margin of the limb, twice bent at obtuse angles about 130?-150?, early
deciduous and thus leaving a twice bent scar in the dense layer of filiform hairs
covering the inflorescence. The hexagonal, transversally extended, area
delimitated by the scars of four neighboring bracts represents one extremely
depressed branch subtended by the lowermost bract. Hairs thinly clavate, 2.4
mm long. Male flowers 20-30 on each branch, at first embedded in layer of
hairs, later growing to about 5 mm with a lower, tubular part about 2.5 mm
and tepals ovate, concave, 2.5 x 1.2 mm. filaments fused into a column 6-7
mm long, tubular, inserted at base of perianth-tube, below synandrium split
into 3 free filaments. Synandrium ovoid-ellipsoid, composed of 3 anthers each
with 2 cells, dehiscing above insertion of filaments, 1.1 x 1.0 mm. Rudimen-
tary ovary (?) protruding as a short, conical structure into lower part of
perianth-tube. Pollen grains spheroid, 6-pantopororate, 40-44 [m diam., exine
2 /m, nuclei 2. Female flowers with perianth adnate to ovary, 2 short and
broad, lip-like segments protruding above ovary. Ovary 1.3-1.4 mm long,
compressed, 0.4 mm wide, with flat side generally facing insertion of bracts;
styles 2, 1.7 mm long, appearing above layer of hairs; stigmas capitellate, 0.2
mm across. Fruit a small, 1-seeded achene.
Type. Swartz sn, Jamaica (lectotype here selected, UPS; isolectotype, S).
Distribution. In forests in dense shade, alt (200-)300-700(-1400) m.
Flowering all year round, 45% of known collections February-April. Host
plant recorded: Cassia emarginata L. (Leguminosae).

CUBA. Oriente: Clement sn (NY); Ekman 3348 (K, NY, S); 5113 (S); 9037 (NY, S); Morton &
Acuna 3615 (UC); Shafer 8858 (NY); Taylor 219 (NY); Wright 546 (BR, GOET, K, MO, NY, P,
UC). JAMAICA. Alexander sn (GOET, K); Anderson & Sternberg 3117 (MO); Britton & Hollick
2780 (NY); Howard & Proctor 13400 (BM, LE, NY, S, U); Kramer 1780 (U); Lloyd sn (P); Maxon
9137 (S); Maxon & Killip 559 (NY); Morley & Whitefoord 638 (MO); Morris sn (K); Orcutt 2844
(UC); 5706 (K, UC); Philipson 1253 (BM); Proctor 9341 (NY); 21465 (NY); Purdie sn (K); Ridley
sn (K); Schrink sn (MO); Webster et al 4981 (BM); 8352 (S); Wullschligel 1300 (GOET, M); Yun-
cker 17553 (S); 18466 (BM, S). HAITI. Ekman 2653 (S); 3877 (S); 10581 (S); Leonard 5763 (NY).
SANTO DOMINGO. Bertero sn (MO); Eggers 1620 (C, K, L, M); Ekman 14285 (K, S); Liogier
14463 (NY); Tuerckheim 700 (BR); 2657 (BM, BR, K, L, M, MO, NY, S). PUERTO RICO.
Sintenis 4238 (B, BM, BR, GOET, K, L, LE, M, MO, NY, P, S).

4. Scybalium glaziovii Eichler in de Candolle, Prod. 17: 133. 1873; Engler,

Nat. Pflanzenf. ed. 1. 3. 1: 257. 1889; Glaziou, Mem. Soc. Bot. France
1(3): 610. 1913; Harms, Nat. Pflanzenf. ed. 2. 16b: 319. 1935; Brade,
30 Flora Neotropica

'7~~~~~~~~~~~~~

-. F~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~J

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

-~
CC ~~ ~ ~

N ~ 3i

~~~~h ~ ~~
N.~~~~~~~~~~~~~~~~r?.?
E
LT~~~Sa`
FI .Seiso SyaimadCrnaa -,S aaces,A ,atrHokrf I86,B
C Egr n 87,E(lchlseimna )oignl ,hbi,bat o nlrseneaese
xO5 B ongiforsecesil oeedb rat x.;C,ml ifoesec wt ud p

FIG 11I.Speciesof Scybaliumand Corynaea.A-E, S. jamaicense,A, D, afterHookerf. (1856),B,

C (Eggerssn 1887),E (alcoholspecimenat K) original.A, habit, bractsof inflorescenceareshed
x 0.5; B, young inflorescencestill coveredby bracts x 0.5; C, male inflorescencewith budsap-
Balanophoraceae 31

Arch. Jard. Bot. Rio de Janeiro13: 65. 1953;Falcao, Atas Soc. Biol. Rio
de Janeiro 14: 154. 1971;Fl. I1. Cat. BALA 22, f. 6. 1975.
Fig 11, Fig 3 (pollen), Fig 12 (distr.).

Descriptionbased on wet and dry specimens.Monoeciousor dioecious

root parasites(inflorescencebisexualor unisexual).Total lengthfrom point of
contactwith host root 15-20cm. Tubersubspherical,2.5-5 cm highand 2.5-7.5
cm across, surfaceslightlytessellate.Stems up to 10 from one tuberor com-
plex of tubers,cylindrical,7-11x 1-1.5cm. Leavesscaly, numerous,apparent-
ly whorled and arrangedin 8-12 vertical ranks, densely imbricate,almost
triangular,acute, 10-14x 3.5-6 mm, erect. Inflorescencesubsphericalto ellip-
soid, 2.8-5 x 2.5-3.5 cm, denselycoveredby imbricate,(10-)14-20ranked,sca-
ly bractswhenyoung. Bractstransverselytriangular,acute, marginscrenateto
fringed, 13-19x 9-14 mm, insertedby a broad-canaliculate 3 mm long petiole,
whichis transversallyextendedand attachedalong the entirelowermarginof
the limb, twice bent at obtuse angles about 120?-140?,early deciduousand
thus leavinga twice bent scar in the dense layerof filiform hairscoveringthe
inflorescence.The hexagonal,transversallyextendedarea delimitatedby the
scars of four neighboringbracts representsone extremelydepressedbranch
subtendedby the lowermostbract.Hairsthinlyclavate, 1.5 mm long. Mostin-
florescencesin the materialstudiedpurelyfemale with only 2-3 rudimentary
(?) male flowers in the centre(top) of each branch.One inflorescencewith a
few femalebranchesbasallyand purelymale branchesabove. Male flowersas
in S. jamaicensewith a lower tubularpart crownedby 3 tepals, the filaments
forminga columnsplittingbelowthe synandriuminto 3 free filaments.Pollen
grains spheroid, 6-pantopororate,44.5 ,/m diam., exine 1.5 ,tm. Female
flowers with perianthadnate to ovary, 2 short and broad, lip-likesegments
protrudingabove ovary. Ovary compressed, 1.2 mm long and 0.3-0.4 mm
wide with flat side generallyfacing insertionof bracts;styles 2, 1.9 mm long,
appearingabove layer of hairs; stigmas capitellate, 0.2 mm across. Fruit a
small, 1-seededachene.
Types. Glaziou 4070b and 4700, Brazil, Rio de Janeiro, Alto Macahe,
novo Friburgo,alt. 1400m (mixedcollection, lectotype, P; isolectotype,C).
Distribution.In forests (?), alt. 1400-2000m, season not known. Host
plants recorded:Myrsinaceaeand Melastomataceae.
BRAZIL. Rio de Janeiro. Brade 18909 (RB 74289); Dungs sn (HB); Lima & Brade sn (RB
24795); Peckolt sn 1884 (B); Smith 1777 (S, GH).

Tribus Helosieae Schott & Endlicher,Melet. Bot. 11. 1832 p.p. guoad gen.
Helosis; Endlicher, Gen. P1. 74. 1836 p.p. quoad gen. Helosis.

pearing above layer of hairs x0.5; D, male flower x5; E, female flower x7. F-K, S. glaziovii
(Glaziou 4070b, 4700) original. F, habit, inflorescence still covered by bracts x0.5; G, in-
florescence with bracts partly removed, each branch visible due to deep fissures in the layer of
hairs x 1; H, bract subtending inflorescence branch, front view x 3; 1, the same, side view x 3; K,
female flower x 7. L-O, Corynaea crassa, L, N after Hooker f. (1856), M, O (G6moezsn) original.
L, habit of large specimen x0.5; M, detail of another specimen x0.5; N, male flower of var
crassa x 5; 0, male flower of var sprucei x 4.5.
 32 Flora Neotropica

~:. ,f BI?'
9 0T o Is
W 9f

^ -4-i ---.-./- ^.';oc^^ ^^,^ ^ . 4 '

... .- - ..... ... . .. .. .. . . - -

'-N, '" ^ C ^v^

t\\ -- Lu_,a,>
t,-,'1- ,
! '"i

i; 2t ' .
tX.z

-- --. -. .. -. + _ - -. .-. t -

.- -1- -
-- ----- '- T:-4op.co -

FIG 12. Distribution of Scybaliurn. S. depressum,4; S. f/ungiJorme, *; S. jamaicense, A; S.

gla:.iovii,&.

Helosideae Hooker f. [Trans. Linn. Soc. London 22: 1856 nom. nud. stat. incert.] ex
Eichler, Actes Congr. Bot. Paris 152. 1867 p.p. quoad gen. Helosis, Corynaea, Rhopaloc-
nemis; in Martius, Fl. Bras. 4. 2: 5. 1869 p.p. quoad gen. op. cit.; emend. Eichler in DC.,
Prod. 17: 134. 1873; Engler, Nat. Pflanzenf. ed. 1. 3. 1: 258. 1889.

Stem without scaly leaves. inflorescences when young covered by hex-

agonal, peltate, valvate bracts. Pollen grains 3-colpate.
Type genus. Helosis Richard, Mem. Mus. Hist. Nat. 8: 416, 430, 432.
1822.
American genera: Corynaea and Helosis.
Obviously Helosis and Corynaea are closely related to each other and to
Balanophoraceae 33

the additional members of Helosieae, ie Ditepalanthus (Madagascar),

Rhopalocnemis (Himalaya, Vietnam, W Malesia) and Exorhopala (Malay
peninsula) all of them sharing the tribal and subfamiliar characters mentioned
above. However, Helosis and Corynaea are morphologically well distinguish-
ed from each other by the structure of the synandrium and the underground
parts as well as by the scaly to ring-like involucrum found only in the stems of
Helosis. In their common area of distribution along the Andes from Costa
Rica to Bolivia they are remarkably altitudinally separated (Fig 13) with
Helosis mainly below 2000 m alt. and Corynaea mainly above 2000 m alt. Fig
13 shows a slight tendency for the demarcation line between the two species to
ascend to higher altitudes in both directions from the equatorial region,
although more collections from Peru and Ecuador are desirable to settle this.

2. Helosis L. C. Richard, Mem. Mus. Hist. Nat. 8: 416, 430, 432. 1822, nom.
cons.; Schott et Endlicher, Melet. Bot. 11. 1832; Martius, Nov. Gen.
Spec. Plant. 3: 184. 1832; Endlicher, Gen. Plant. 74. 1836; Griffith,
Trans. Linn. Soc. London 20: 104. 1846; Hooker f., Trans. Linn. Soc.
London 22: 31, 55. 1856; Grisebach, Fl. Brit. W. Ind. 309. 1860; Eichler
in Martius, Fl. Bras. 4. 2: 21. 1869; in DC., Prod. 17: 134. 1873; Hooker
f. in Bentham et Hooker f., Gen. P1. 3: 237. 1880; Engler, Nat. Pflanzenf.
ed. 1. 3. 1: 258. 1889; Duss, Ann. Inst. Bot.-Geol. Marseille 3: 325. 1897;
Lanjouw in Pulle, Fl. Suriname 1. 1: 45. 1932; Harms, Nat. Pflanzenf.
ed. 2. 16b: 319. 1935; Macbride, Fl. Peru 2: 430. 1936; Standley, Fl.
Costa Rica 410. 1937; Standley & Steyermark, Fl. Guatemala 4: 92. 1946;
Le6n & Alain, Fl. Cuba 2: 84. 1951; Lemee, Fl. Guy. Franc. 1: 545. 1955;
Nevling, Ann. Missouri Bot. Gard. 47: 304. 1960; Falcao, Rodriguesia
25(37): 135. 1966; Angely, Fl. Anal. e Fitog. Sao Paulo 1: 56. 1969;
Falcao, Atas Soc. Biol. Rio de Janeiro 14: 151. 1971; Fl. II. Cat. BALA
27. 1975.

Caldasia Mutis ex Caldas in Seman, Nuevo Reyno Granada 1810(2): 26. 1810, not seen,
nom. illeg.
Latraeophila Leandro do Sacramento ex A. St.-Hilaire, Ann. Sci. Nat. Bot. Ser. 2. 7: 32.
1837, nom. nud.

Fleshy root parasites. Subterranean system formed by a central, starchy

tuber at point of contact with host root; from the central tuber a number of
horizontal, rhizome-like structures spread in all directions. Supra-terraneous
inflorescence-bearing stems appearing from buds on the rhizome-like organs,
leafless. Bud scales 2-6, free or often joined to a ? oblique ring-like mark,
mostly inserted at the same level close to the base of the stem, sometimes at
higher levels. Inflorescence ellipsoid or ovoid, at first covered by hexagonal
bracts. Bracts peltate, early deciduous. Flowers embedded in a dense layer of
filiform hairs. Male flower with a tubular, 3-lobed perianth. Stamens 3, con-
nate; synandrium 9-locular with locules longitudinally arranged. Pollen grains
spheroid, 3-colpate, medium sized, 3-nucleate. Female flowers compressed
with perianth adnate to ovary, segments 2, inconspicuous. Styles 2. Fruit a
small, I-seeded achene.
34 Flora Neotropica

m alt. A m alt.

A A
3000 . 3000
A A

A
A

2000 * 2000
A 0o
* O
0
0 0
0 0 0
A 0
0? ?o
*?.0 *
1000 0 1000
% %
*

S lat. 20 15 10 5 0 5 10 15 20 N lat.
A Corynaea crassa
o Helosis cayennensis var mexicana
* Helosis cayennensis var cayennensis

FIG 13. Altitudinal distribution of Helosis and Corynaea along the Andes.

Type species. Helosis guyanensis L. C. Richard, Mem. Mus. Hist. Nat. 8:

416, 430, 432. 1822, nom. illeg. = Helosis cayennensis (Swartz) Sprengel.
Distribution. Tropical rain forests and montane forests at lower altitudes
from Mexico through Central America to tropical South America.
The distinction at specific level between Helosis cayennensis and H. mex-
icana is abandonned here for various reasons. In the rich material now
available from the entire area of distribution there is an almost continuous
variation in the one and only distinguishing character, the level at which the in-
volucrum is inserted. In 215 individuals of 90 well preserved specimens the
ratio, R, between the length of the stem and the distance: base of stem to in-
volucrum, was calculated. The values were transferred to 10 classes, each class
corresponding to 1/10 the length of the stem. The distribution of the material,
calculated in percent, over the 10 classes is shown in Fig 14. Around class 4
there is a distinct gap from 2.41 to 3.14 in the R-values represented. I have
chosen per definition to discriminate between var cayennensis and var mex-
Balanophoraceae 35

icana at the R-value 2.8, in other words specimens with R-values >2.8 are
referred to var cayennensis, those with R-values <2.8 to var mexicana. The
position of the type of var mexicana with R = 1.27 is indicated in Fig 14. Thus
defined var mexicana is represented with classes 7-10 in Mexico, 5, 7 and 9 in
Central America, 6 in Cuba, 5 and 7-9 in the Lesser Antilles, 7 in Colombia, 9
in Ecuador, and 5 and 7 in SE Brazil. It is apparently confined to more or less
hilly country. Short-stemmed forms with proportionately large inflorescences
are found throughout the distribution area. Such forms were the basis of
Helosis brasiliensis Schott & Endl., an entity I find impossible to distinguish at
any taxonomic level.

1. Helosis cayennensis (Swartz) Sprengel, Syst. Veg. 3: 765. 1826; Sandwith,

Kew Bull. 1931: 58; Lanjouw in Pulle, Fl. Surinamense 1. 1: 46. 1932;
Harms, Nat. Pflanzenf. ed. 2. 16b: 321. 1935; Macbride, Fl. Peru 2: 430.
1936; Lemee, Fl. Guy. Franc. 1: 545. 1955; Falcao, Fl. II. Cat. BALA 32.
1975. Fig 15, Fig 4 (pollen), Fig 16, 17 (distr.).

Cynomorium cayennense Swartz, Nov. Gen. & Sp. PI. 12. 1788; Fl. Ind. Occ. 1: 13.
1797; ("cayanense").

Class stem R
(levelof scales) base to scales
10
9
8 -------type of var mexicana-. 1.

6
2
2.5
4 2.8
3
5
2
10

10 20 30 40 50% of specimens
FIG 14. Helosis cayennensis. Frequency classes corresponding to position of scales on stem (see
text).
36 Flora Neotropica

Helosis guyanensis Richard, Mem. Mus. Hist. Nat. 8: 416, 432, t. 20. 1822 nom. illeg.
superfl. ("guyannensis"); Mart., Nov. Gen. Spec. Plant. 3: 184, t. 298, 300. 1832; Schott
& Endlicher, Melet. Bot. 12. 1832; Hooker f., Trans. Linn. Soc. London 22: 31, 57, t. 16.
1856; Grisebach, Fl. Brit. W. Ind. 309. 1860; Eichler, Bot. Zeitung (Berlin) 26: t. 9, f. 19,
20. 1868; in Martius, Fl. Bras. 4.2: 23, t. 4-6. 1869; in DC., Prod. 17: 136. 1873; Engler,
Nat. Pflanzenf. ed. 1. 3. 1: 258. 1889; Duss, Ann. Inst. Bot.-Geol. Colon. Marseille 3:
326. 1897; Pulle, Enum. P1. Surinam 158. 1906; Huber, PI. Duck. 341. 1908; Glaziou,
Mem. Soc. Bot. France 1(3): 610. 1913; Le6n & Alain, Fl. Cuba 2: 84. 1951; Angely, Fl.
Anal. e Fitog. Sao Paulo 1: 56. 1969. Type. Richard sn, French Guiana, Cayenne (P).
Helosis brasiliensis Schott & Endlicher, Melet Bot. 12. 1832 ("brasileensis"); Eichler in DC.,
Prod. 17: 135. 1873; Engler, Nat. Pflanzenf. ed. 1. 3. 1: 258,f. 164. 1889; Harms, Nat.
Pflanzenf., ed. 2. 16b: 321,f. 161, 162. 1935; Falcao, Rodriguesia 25(37): 136. 1966; Atas
Soc. Biol. Rio de Janeiro 14: 151. 1971; Martins f., Leandra 2: 98. 1972; Falcao, Fl. I1.
Cat. BALA 28. 1975. Type. Schott sn, Brazil, Mt. Serra d'Estrella (Not located)

wF C B

FIG15. Helosis cayennensis. A, after Eichler (1869), B, C, after Richard (1822), D, after Hooker
f. (1856). A, habit of var cayennensis x0.5; B, male flower basally surrounded by clavate hairs
x 7; C, female flower surrounded by clavate hairs x 12; D, habit of var mexicana x 0.5.
Balanophoraceae 37

Caldasia cayennensis (Sw.) Mutis ex Steudel, Nom. ed. 2. 1: 255. 1840; Kuntze, Rev. Gen.
2: 590. 1891.
Helosis guyanensis forma brasiliensis (Schott & Endlicher) Eichler in Martius, Fl. Bras. 4. 2:
23, t. 5, f. 1. 1869.
Caldasia brasiliensis (Schott & Endlicher) Kuntze, Rev. Gen. 2: 590. 1891.
Helosis mexicana auct. non Liebmann: Standley & Steyermark, Fl. Guatemala 4: 93. 1946
p.p.; Nevling, Ann. Missouri Bot. Gard. 47: 304. 1960 p.p. quoad Kuntze 74 (= 1926) and
Seibert 469.

Description based on wet and dried specimens. Fleshy orange-red to

brown-red root parasites. Subterranean system formed by a subspherical,
starchy, central tuber, 4-5 x 3-4 cm, at point of contact with the host root;
from the central tuber a number of horizontal, rhizome-like structures spread
in the soil in all directions, apparently forming new tubers at contact with
possible host roots. Supraterranean, inflorescence-bearing stems appear from
buds on the rhizome-like organs, (1-)5-10(-31) cm long and (0.2-)0.4-0.6(-1.1)
cm diam. Bud scales 2-6, free or often joined into a ring-like mark, mostly in-
serted at the same level close to the base of the stem, sometimes inserted at
higher levels. Inflorescence ellipsoid or ovoid (0.7-)2-4(-7.2) cm long and
(0.6-)1-2(-3.4) cm wide, covered by the flattened or conical, marginally coher-
ing, hexagonal peltas of peltate, petiolate bracts when young. Bracts with pelta
3.5-4.5 mm diam. and 2.2-2.3 mm thick, flattened or with a low, conical knob
in the centre, petiole 2.2.-2.4 mm long, 0.3-0.5 mm wide below and 1-1.2 mm
wide in upper part, early deciduous. Each bract subtends an extremely depress-
ed, hardly discernible branch delimitated by imaginary lines between the
petioles of four neighboring bracts. Branch bisexual and strongly pro-
terogynous. Flowers embedded in a dense layer of filiform, narrowly clavate
hairs, which are 1.5-2.3 mm long. Male flowers with perianth tubular below
and 3-lobed in upper part, segments ligulate, 2 x 1 mm. Stamens 3, filaments
united into a tubular column inserted in the perianth tube; free tubular part of
column 1.5-2 mm long and 0.4-0.6 mm wide, in upper part split into 3
filaments 0.4-0.7 mm long and 0.2-0.3 mm wide, which carries the synan-
drium. Anthers connate into a 9-locular synandrium with the locules
longitudinally arranged; synandrium spheroidal, 0.9-1.1 mm diam. A short,
conical structure 0.5 mm long protrudes from the receptacle into the basal part
of the perianth tube (rudimentary ovary ?). Female flower with perianth ad-
nate to ovary, 2 short and broad ligulate segments protruding above ovary,
compressed with edge generally facing petioles of bracts, 1.2 mm long and
0.4-0.5 mm wide. Styles 2, filiform, 1.2-1.5 mm long, appearing above layer of
hairs, caducous; stigma capitellate. Fruit a small, 1-seeded achene.

Key to the Varieties of Helosis cayennensis

1. Stem basally with 2-6 minute, triangular scales. a. var cayennensis.

1. Stem in middle or upper part with a more or less oblique, slightly swollen, ringlike mark.
b. var mexicana.
38 Flora Neotropica

la. Helosis cayennensis (Swartz) Sprengel var cayennensis.

Stem basally with an involucre of 2-6 minute, triangular scales; the ratio
between the length of the stem and the distance from base of stem to involucre
not less than 2.8.
Type. Richard sn, French Guiana, Cayenne between 1784 and 1789 (lec-
totype here selected, C; isolectotype, P).
Distribution. Tropical rain forests and montane forests in damp places,
alt. 0-1500(-2000) m. Flowering specimens collected all year round. Hosts
recorded: Boehmeria sp. (Urticacaceae), Inga sp. (Leguminosae).

MEXICO. Morelos: Miranda 1483 (MEXU); Tabasco: Sarakhan 1985 (MEXU); Matuda
3558 (MEXU); Chiapas: Breedlove 22479 (MO); Miranda 7530 (MEXU). GUATEMALA. Peten:
Hedger 74 (BM); Steyermark 46131 (F, MO, NY); Alta Verapaz: Tuerckheim 8576 (K, NY);
Izabal: Steyermark 39584 (F). BRITISH HONDURAS. Belize: Dwyer 11696 (MO); Muchaca:
Schipp 1129 (BM, F, K, MO, NY, S). HONDURAS. Atlantida: Standley 54815 (F). COSTA
RICA. Guanacasta: Brenes 12638 (F); Tilaran: Kupper 1549 (M); Heredia: Maas 1334 (U); Car-
tago: Maas 1178 (C, U); Golfo Dulce: Pittier 9900 (BR). PANAMA. Chiriqui: Wilburet al 15454
(MO); Cocle: Seibert 469 (K, MO); Canal Zone: Kuntze 1926 (NY); Kennedy & Wilder3101 (MO);
Darien: Duke 13621 (MO); Gentry & Mori 14018 (MO); Stern et al 737 (LE). COLOMBIA.
Boioassu: Black et al 46241 (TAN); Bolivar: Curran sn (US); 334 (US); Santander: Haught 2010
(US); Choc6: Fosberg et al 21520 (US); Tolima: Barriga 8395 (COL, US); Meta: Giovanni 36
(COL); El Valle Cauca: Alston 7819 (BM); Killip 8354 (GH, NY, US); Vaupes: Pinto & Sastre 918
(COL, P); Caqueta: Castaneda 4054 (COL); Amazonas: Barriga 14700 (COL); Jaramillo 2451
(COL); Plowman et al 2345 (GH); Schultes 6197 (GH); 6323 (COL); 6657 (COL); 6829 (US); 8117
(GH); 12091a (GH). VENEZUELA. Merida: Bernardi 3347 (NY); Fendler sn (GH); Amazonas:
Maguire et al 31606 (F, GH, K, NY, P, RB, U, US); 31607 (NY, U); Aragua: Robyns et al sn (BR);
Steyermark 89838 (US); Bolivar: Steyermark 86840 (NY); 88983 (NY, US); 89372 (US); Wurdack
& Monachino 39757 (F, GH, K, NY, R, RB, US); Delta Amacuro: Rusby & Squires 105 (BM, F,
GH, M, MO, NY, US). TRINIDAD. Arena forests: Broadway 6434 (BM, S); 6568 (BM, S);
Crueger sn (K); Aripo: Herb. Trin. 15416 (K); Mt. Tamana: Purdie sn (K). GUYANA. Brit. Gui.
For. Dept. G. 79 (K); F 3023 (K); Cowan et Soderstrom 2005 (NY); Jenman 1852 (K); Maguire et
al 23524 (F, GH, K, MO, NY, U, US); Marlyn 269 (K); Sandwith 620 (K); Ward 8629 (K).
SURINAME. Cowan & Lindeman 39116 (NY); Daniels & Jonker 733 (U); 1134 (U); 1277 (U);
Florschitz & Maas 2545 (U); 2604 (GH, U); 2863 (U); 2942 (U); Jonker- Verhoef & Jonker 443
(NY, U); Kegel 607 (GOET); Kramer & Hekking 2478 (U); Maas 11068 (U); Maguire 40733 (NY,
U); 40794 (F, NY, U); Prance et al 55899 (U); Schulz 10287 (U); Splitgerber 728 (L); Stahel 803
(U); 4659 (U); Teunissen 12951 (C); Wessels Boer 734 (F, U, UC); Wullschlagel sn (BR); 843
(GOET). FRENCH GUIANA. Broadway 799 (GH, NY, US); Granville 72 (CAY, P); 2164
(CAY); Halle 636 (P); Maas 2180 (C, U); Oldeman 1141 (CAY); 1199 (CA Y); Sastre 1635 (P);
Schnell 12239 (P). ECUADOR. Guayas: Haught 2881 (US); Azuay: Lehmann 401 (K); Morona
-Santiago: Holm-Nielsen et al 4155 (AAU); Sparre 19070 (S). PERU. Loreto: Asplund 14226 (S);
Croat 18824 (MO); Ducke 7595 (MG); Ferreyra3401 (US); Killip 27622 (US); Simpson & Schunke
660 (F, NY); Wurdack 2112 (US); Amazonas: Berlin 404 (MO); 457 (MO); Junin: Ruiz sn (FI);
Cuzco: Plowman 4934 (C, CUZCO, F, GH). BOLIVIA. Pando: Prance et al 5804 (NY); 8513
(NY). BRAZIL. Gwynne Vaughan 55 (K); Miers sn (BM, K); Riedel sn (LE); Snethlage 231 (B);
Rondonia: Prance et al 8765 (F, GH, MG, NY, R, S, U); Roraima: Pires et al 14521 (IAN); Prance
et al 4457 (NY); 9370 (NY); 10499 (U); 10819 (C, NY); 13565 (MG, NY, U); Amapi: Black 498423
(IAN); Egler & Irwin 46697 (NY); Luetzelburg 20143 (R); 20173 (R); 20212 (R); 21245 (M, R);
21679 (M); 21866 (M); 21891 (M); Maguire et al 47006 (NY, U, US); 47091 (IAN, K, NY, SP, U);
Amazonas: Baldwin 3302 (US); Black et al 483416 (IAN); Cavalcante 3205 (MG); Chagas 3122
(INPA); 3553 (IAN, INPA, MO); 6309 (IAN, INPA); Ducke 6875 (MG); Ernani 6424 (INPA);
Fereira 58221 (IAN); Martius 2643 (M); Prance et al 3589 (NY, US); 8080 (MG, C); 10200 (F, MO,
NY, R, S, U); 14391 (NY, U); 15787 (C, F, MG, NY, S, U); 16309 (F, GH, M, MG, NY, S, U);
24212 (C); 24410 (C); Rodrigues 8347 (INPA); Silva 689 (INPA); Spruce 877 (BM, FI, LE, M);
1129 (GH, K); Ule 6069 (B); Para: Austin & Cavalcante 4137 (MO); Black 47957 (IAN); 472039
Balanophoraceae 39

.. -.- .. ...-.- - Tc
---

sTsf.! 1il9r^ .e,x ^ _ S

-- 1- . - '> - t--

~(IAN)~; 482388 (IAN);l 482961 (IAN); 498584 (IAN); Black et al 1410 (IAN); Ca

FIG 16. Distribution of Helosis cayennensis var cavennensis.

(IAN); 482388 (IAN); 482961 (IAN); 498584 (IAN); Black et al 1410 (IAN); Cavalcante 2261
(MG); Ducke 3462 (MG); 3561 (MG); 3632 (MG); Guedes 2298 (MG); Lima 531437 (IAN);
531526 (IAN); Luetzelburg 21085 (R); 21259 (M, R); Pires 1239 (IAN); Pires & Silva 4511 (IAN);
Prance et al 25483 (C, NY); Sastre 1518 (P); Silva 1241 (MG, SP); Spruce 151 (K); 574 (GH, K);
Acre: Prance et al 2764 (NY); 7833 (MG, NY, U); Paraiba (?): Luetzelburg sn (M); Guanabara:
Araujo et al 204 (RB); Rio de Janeiro: Araujo et al 523 (RB); Glaziou 4969a (BR, C, P); Lutz &
Sigueira sn (R); Miers sn (BM); Occhioni 981 (RB); Sick 783 (HB); Sao Paulo: Kuhlmann 4613
(SP); Mathias 5673 (SP); ParanA: Hatschbach 11287 (US); 20981 (C); Santa Catarina: Lourteig
2320 (P); Moller sn (B); Rio Grande do Sul: Leite 720 (NY); Malme sn (S).

The typification of Helosis cayennensis (Sw.) Spr. involves a few pro-

blems. In a posthumous work L. Richard (1822) stated that in Cayenne he gave
von Rohr some material in liquid of a parasitic plant later known as
 40 Flora Neotropica

Cynomorium cayennense Sw. This event must have taken place during
Richard's visit to Guiana and the West Indies 1784-1789 more precisely in 1784
or 1785 when von Rohr visited Cayenne. von Rohr was understood to bring
the material to Banks in London, which apparently he did, for Swartz (1788)
published Cynomorium cayennense and clearly indicated that the description
was drawn on the basis of material studied in London on his way back to
Sweden where he arrived in 1789. No type material has been traced at BM and
it must have been lost. However, von Rohr used to send plant material from
his extensive journeys to Vahl in Copenhagen. As a matter of fact there is at C
a specimen from Vahl's herbarium with "Cynomorium cajanense" and
"Richard" in Vahl's handwriting. I consider this a duplicate of the material
studied by Swartz and have chosen it as lectotype. A Richard specimen at P
may be another duplicate, and I have called it an isolectotype. The specimen at
P must be the type for the superfluous name Helosis guyanensis Richard.

lb. Helosis cayennensis (Swartz) Sprengel var mexicana (Liebmann) B.

Hansen, Bot. Tidsskr. 72: 188. 1978.

Helosis mexicana Liebmann, Forhandl. Skand. Naturf. 4. Mode 1844: 181. 1847; Hooker
f., Trans. Linn. Soc. London 22: 31, 59, t. 15, 16. 1856; Grisebach, Cat. PI. Cub. 118.
1866; Eichler in DC., Prod. 17: 136. 1873; Engler, Nat. Pflanzenf. ed. 1. 3. 1: 258. 1889;
Standley, Fl. Yucatan 250. 1930; Fl. Lancet. Hond. 178. 1931; Harms, Nat. Pflanzenf.
ed. 2. 16b: 321. 1935; Standley & Record, For. & Fl. Brit. Hond. 123. 1936; Standley, Fl.
Costa Rica 410. 1937; Standley & Steyermark, Fl. Guatemala 4: 92. 1946 p.p.; Nevling,
Ann. Missouri Bot. Gard. 47: 304,f. 87. 1960 p.p. quoad ill.
Helosis guyanensis var andicola Hooker f., Trans. Linn. Soc. London 22: 57. 1856. Type.
Goudot 140, Colombia (lectotype here selected, K; isolectotype, FI).
Helosis guyanensis forma andicola (Hooker f.) Eichler in Martius, F. Bras. 4. 2: 23. 1869.
Helosis mexicana var andicola (Hooker f.) Eichler in DC., Prod. 17: 136. 1873.
Caldasia mexicana (Liebmann) Kuntze, Rev. Gen. 2: 590. 1891.

Stem in middle or upper part with an involucre represented by a more or

less oblique, slightly swollen, ring-like mark; the ratio between the length of
the stem and the distance from base of stem to involucre less than 2.8.
Type. Liebmann sn, Mexico, Veracruz, Mirador, alt. 1000 m, 1842; sn,
Mexico, Oaxaca, Tepilongo, alt. 1500 m, 1842 (mixed collection, holotype, C).
Distribution, Tropical rain forests and montane forests in damp places,
alt. 500-1500(-1800) m. Flowering specimens collected all year round. Hosts
recorded: Achras zapota L. (Sapotaceae), Ficus sp. (Moraceae).

CUBA. Arroyo veintecenco: Wright2636 (BM, GOET, K, LE, MO, NY, P, S). MARTINI-
QUE. d'Alleizette 6363 (L); Duss 2106 (NY); Hahn 400 (BM). SANTA LUCIA. Proctor 17764
(BM); Ramage sn (K). MEXICO. Matuda 423 (MEXU, MO, NY); 4167 (MEXU, NY); San Luis
Potosi: Pringle 5050 (MEXU); Veracruz: Beaman 6185 (U); Bourgeau 3030 (FI, K, LE); Dressier
& Jones 38 (MEXU, NY); Linden 146 (BR, FI, K); Purpus 2750 (BM, F, MO, NY); 5896 (BM,
MO, NY); 7691 (MO, NY); Chiapas: Peldes sn (MEXU). GUATEMALA. Huehuetenango:
Steyermark 51848 (F, NY); Quezaltenango: Walker449 (BM); Alta Verapaz: Tuerckheim 1044 (K,
NY, P); Suchitep6quez: Steyermark 46801 (F). BRITISH HONDURAS. Lundell 6414 (F, MEXU,
NY, S); Schipp 719 (F). NICARAGUA. Hawkes, Hjerting & Lester 2152 (C, K). COSTA RICA.
Cartago: Lent 788 (F); 1282 (F, MO); Wilbur & Stone 10309 (F); Alajuela: Lent 1658 (F); San
Jose: Skutch 2669 (K, MO, S). PANAMA. Chirique: Croat & Porter 16158 (MO). COLOMBIA.
Balanophoraceae 41

- r'- I
Tg

--!t-f
-.
t
t*^ + ?*--

_ --_- "- -1-- ---- -- Tiod-.a

_ _ _ _ _ _ _ _ I
Soo0 000 500

i ^
^h

.-- ...-.4--... . -
-1-\.-.J.i.- . . ....
_ ____ L
FIG 17. Distribution of Helosis cayennensis var mexicana.

El Valle Cauca: Welden sn (C); Melgar: Purdie sn (K). ECUADOR. Los Rios: Eliasson 13 (GB).
BRAZIL. Rio de Janeiro: Pires sn (R); Santa Catarina: Reitz & Klein 7328 (UC, US); 17353 (US);
Ule 748 (US).

3. Corynaea Hooker f., Trans. Linn. Soc. London 22: 31, 54. 1856 emend.
B. Hansen; Eichler in DC., Prod. 17: 137. 1873; Hooker f. in Bentham &
Hooker f., Gen. PI. 3: 237. 1880; Engler, Nat. Pflanzenf. ed. 1. 3. 1: 258.
1889; Harms, Nat. Pflanzenf. ed. 2. 16b: 322. 1935; Macbride, Fl. Peru 2:
431. 1936; Standley, Fl. Costa Rica 410. 1937; Nevling, Ann. Missouri
Bot. Gard. 47: 305. 1960.

Itoasia Kuntze, Rev. Gen. 2: 590. 1891.

Helosis auct. non Richard: Baker f., J. Bot. 28: 161. 1890.
42 Flora Neotropica

Fleshy root parasites. Underground tuber somewhat irregularly lobed,

starchy. Stem leafless, short or elongated. Inflorescence subspherical to ellip-
soid or obovoid, when young covered by hexagonal bracts. Bracts peltate, ear-
ly deciduous. Flowers embedded in a dense layer of filiform hairs. Male
flowers with a tubular 3-lobed or irregularly crenate perianth. Stamens 3, con-
nate; synandrium 6-locular with locules longitudinally arranged. Pollen grains
spheroid, 3-colpate, medium sized, 3-nucleate. Female flowers compressed
with perianth adnate to ovary, segments 2, inconspicuous. Styles 2. Fruit a
small, 1-seeded achene.
Type species. Corynaea crassa Hooker f., Trans. Linn. Soc. London 22:
31, 54, t. 13. 1856, lectotype here selected; hereafter paratypes: Corynaea
sphaerica Hooker f. l.c.: 55, t. 14, and Corynaea purdiei Hooker f. l.c.: 55.
Distribution. Montane forests at higher altitudes from Costa Rica to
Bolivia and in NW Venezuela.
J. D. Hooker (1856) described the genus Corynaea to accommodate three
species, each based on one or two collections and all having a tubular, funnel
shaped, irregularly crenate perianth without obvious lobes. Eichler (1873) add-
ed a fourth species characterized by an obviously 3-lobed perianth. Forty
specimens were available for the present treatment. A wide and continuous
variation was found in the length of stem as well as the shape and size of
inflorescence. These same characters were used by Hooker in distinguishing
his species. Investigations of a large number of specimens of Balanophora
have shown that such characters are most unreliable in this group of parasites
(Hansen 1972). This is apparently the case in Corynaea too, and I have been
forced to unite Hooker's species into one. Furthermore I have reduced
Eichler's C. sprucei to a variety of C. crassa. The only reliable distinguishing
character is the 3-lobed perianth, found in C. crassa var sprucei, but even in C.
crassa var crassa specimens may be found with 3 deeper incisions in the crenate
perianth. Finally it should be mentioned that purely unisexual inflorescences
are found quite often ie in 12 of the 40 specimens studied. It is not possible at
present to refer purely female inflorescences to varietal level if they do not oc-
cur together with male ones from one tuber.

1. Corynaea crassa Hooker f., Trans. Linn. Soc. London 22: 31, 54, t. 13.
1856; Eichler in DC., Prod. 17: 137. 1873; Engler, Nat. Pflanzenf. ed. 1.
3. 1: 259,f. 165F. 1889; Harms, Nat. Pflanzenf. ed. 2. 16b: 322,f. 163F.
1935; Standley, Fl. Costa Rica 40. 1937; Nevling; Ann. Missouri Bot.
Gard. 47: 306,f. 88. 1960. Fig 11, Fig 5 (pollen), Fig 18 (distr.).

Corynaea purdiei Hooker f., Trans. Linn. Soc. London 22: 31, 55. 1856; Eichler in DC.,
Prod. 17: 138. 1873; Engler, Nat. Pflanzenf. ed. 1. 3. 1: 259. 1889; Harms, Nat.
Pflanzenf. ed. 2. 16b: 322. 1935; Macbride, Fl. Peru 2: 431. 1936; Steyermark, Fieldiana
28: 898. 1957. Type. Purdie sn, Colombia and Weddell sn, Peru (not extant, neither at K
nor P).
Corynaea sphaerica Hooker f., Trans. Linn. Soc. London 22: 31, 55, t. 14. 1856; Eichler
in DC., Prod. 17: 137. 1873; Engler, Nat. Pflanzenf. ed 1. 3. 1: 259. 1889; Harms, Nat.
Pflanzenf. ed. 2. 16b: 322. 1935. Type Purdie sn, Colombia (holotype K).
Itoasia crassa (Hooker f.) Kuntze, Rev. Gen. 2: 590. 1891.
Itoasia purdiei (Hooker f.) Kuntze, I.c.
Itoasia sphaerica (Hooker f.) Kuntze, I.c.
Balanophoraceae 43

' ~ ,?I r s

001

-- - -} ~: _
' ~ '

4-

'- !
--v ^ ^^
^^^ -r

TM
--- .i . .
J' -/ ^.*l^ >^:Df, ____
- -
^ -?
?------?-??
---i--(------?-i-- .....------:,;
-^ -\ ^- ,--
_ri o;
--STopic at

FIG 18. Distribution of Corynaea crassa.

Description based on wet and dried specimens. Fleshy yellowish-brown to

deep purple root parasites. Tuber subspherical to ellipsoid, (2.3-)4-6 cm long
and (2.1-)4-8(-10) cm wide, often irregularly lobed and bearing short, cylindri-
cal processes. One or more inflorescence-bearing stems appear from each
tuber, breaking through the outer tissues of the tuber, thus forming an
inconspicuous volva or ring-like mark at the base of the fully developed stem.
Stem leafless, short or elongated, cylindrical, (0.2-)3-7(-10.5) cm long and
(0.3-)0.7-1.6(-2.9) cm wide, often slightly longitudinally furrowed in upper
part. Inflorescences apparently not branched, subspherical to ellipsoid or obo-
void, when young covered by the flattened or conical, marginally cohering
hexagonal peltas of peltate, petiolate bracts; (1.0-)3-7(-11) cm long and (1.2-)
44 Flora Neotropica

2-4.5(-8.5) cm wide without bracts. Bracts with pelta 4-8 mm diam. and 1.5-3
mm thick, flattened or with a low conical knob in the centre, petiole 3 mm
long, 0.5-1 mm wide below and 1.7-2 mm wide in upper part, early deciduous.
Each bract subtends an extremely depressed, hardly discernible branch delimi-
tated by imaginary lines between the petioles of four neighboring bracts.
Branch unisexual or bisexual and then strongly proterogynous. Flowers
embedded in a dense layer of filiform, narrowly clavate hairs, which are 1.8-2
mm long. Male flowers with perianth tubular below and funnelshaped, irregu-
larly crenate to obviously 3-lobed above, 3.5-5 mm long and up to 1 mm wide,
lobes ligulate 1.8 x 0.8 mm. Stamens 3, filaments united into a tubular col-
umn, which again is united with the lower third of the perianth tube; column
2.5-6 mm long and up to 0.3 mm wide. Anthers connate into a 6-locular synan-
drium with the locules longitudinally arranged, 0.8 x 1 mm. A short, conical
structure 0.8 mm long protruding from the receptacle into the base of the
staminal tube might constitute a rudimentary ovary. Female flowers with peri-
anth adnate to ovary, 2 short and broad, lip-like segments protruding above
ovary, compressed with edge generally facing petioles of bracts, 1-1.2 mm long
from base to tip of perianth segments and 0.5-0.6 mm wide, Styles 2, filiform,
1-1.2 mm long, appearing above layer of hairs, caducous; stigma capitellate.
Fruit a small, 1-seeded achene.

Key to the Varieties of Corynaea crassa

1. Perianthof maleflowerswithoutlobes, funnelshaped,irregularlycrenateor sometimeswith
3 deeperincisions, a. varcrassa.
1. Perianthof maleflowersobviously3-lobed. b. varsprucei.

la. Corynaea crassa Hooker f. var crassa

Perianth of male flowers without lobes, funnelshaped, irregularly crenate

or sometimes with 3 deeper incisions.
Type. Purdie sn Colombia near Sta Fe de BogotA (holotype, K).
Distribution. Montane forests, alt. 2000-3000 m. Flowering specimens
collected all year round. Host not recorded.
COSTA RICA. Heredia: Volcan de Barba, Wendland sn (P). COLOMBIA. Choc6: Core
331a (US); Cundinamarca: Forero 1051 (COL); El Rosal, Hirschel sn (COL); Sabaneta, Hirschel
sn (COL); Huertas y Camargo 6336 (COL); Schneider 612 (S). ECUADOR. Tungurahua,
Asplund 10010 (S).

lb. Corynaea crassa Hooker f. var sprucei (Eichler) B. Hansen, Bot. Tidsskr.
72:188. 1978.
Corynaea sprucei Eichler in DC., Prod. 17: 137. 1873; Engler, Nat. Pflanzenf. ed. 1. 3. 1:
259. 1889; Harms, Nat Pflanzenf. ed. 2. 16b: 322. 1935.
Helosis whymperi Baker f., J. Bot. 28: 161, pi. 297, f. A-E. 1890 synon. nov. Type.
Whympersn, Ecuador (holotype, BM).
Itoasia sprucei (Eichler) Kuntze, Rev. Gen. 2: 590. 1891.

Perianth of male flowers obviously 3-lobed.

Balanophoraceae 45

Type. Spruce 5186, Ecuador (lectotype K); Trianasn, Colombia (P).

Distribution. In montane forests, alt. (1250-)1800-3000(-3600) m. Flow-
ering specimens collected all year round. Hosts recorded, Eupatorium angu-
lare B. L. Robinson (Compositae) and "on bamboo roots."
COSTA RICA. Heredia: Volcan de Barba, Burger & Stolze 6060 (F, MO); Gollmer sn (B);
Sierra la Talamanca, Williams & Gibson 28189 (F); Cartago: G6mez 786722 (C). PANAMA.
Chiriqui: Volcan de Chiriqui, Woodson & Schery 470 (MO). COLOMBIA. Goudot sn (P); Meta:
Rio Manzanares valley, Grant 10029 (US); Narifio: Rio de Potreros, Ewan 16544 (US).
ECUADOR. Pichincha: Madison 2356 (C); Sparre 14338 (S); 16947 (S); Napo: Holm-Nielsen et al
6910 (AAU); Azuay: Camp 4295 (C, NY); Harling & Andersson 13209 (GB). PERU. Bongara:
Wurdack 952 (US); Carabaya: Weddell 4786 (P); Ayacucho: Plowman 4688 (C, GH). BOLIVIA.
La Paz, Rusby 256 (NY).

The following material consists of juvenile or female specimens and can-

not at present be referred to varietal level.
COLOMBIA. Choc6: Core 332a (US); Cundinamarca: Forero 1049 (COL); Grant & Fosberg
9311 (US); Grant 9701 (US); Meta: Fosberg 20923 (US); Cauca: Cuatrecasas 19121 (GH); Huila:
Little 7663 (US); Narifo: Cuatrecasas 11851 (COL); Fosberg 21151 (US). VENEZUELA. Lara:
Barclay et al 10325 (US); Steyermark 55247 (F, US); Santiago Zamora: Acosta Solis 5070 (F).

Balanophoraceae subfam. Lophophytoideae Harms, Nat. Pflanzenf. ed. 2.

16b: 311. 1935; Angely, Fl. Anal. e Fitog. SAo Paulo 1: 56. 1969.
Scybalioideae Engler, Nat. Pflanzenf. ed. 1. 3. 1: 253. 1889. p.p. quoad Lophophyteae.

Type genus. Lophophytum Schott & Endlicher, Melet. Bot. 1. 1832.

4. Lophophytum Schott & Endlicher, Melet. Bot. 1. 1832; Endlicher, Gen.

Plant. 73. 1836; Hooker f., Trans. Linn. Soc. London 22: 30, 47. 1856;
Eichler, Bot. Zeitung (Berlin) 26: 551. 1868; in Martius, Fl. Bras. 4. 2: 45.
1869; in DC., Prod. 17: 127. 1873; Hooker f. in Bentham & Hooker f.,
Gen. PI. 3: 238. 1880; Engler, Nat. Pflanzenf. ed. 1. 3. 1: 255. 1889;
Harms, Nat. Pflanzenf. ed. 2. 16b: 324. 1935; Macbride, Fl. Peru 2: 428.
1936; Falcao, Rodriguesia 25(37): 136. 1966; Angely, Fl. Anal. e Fitog.
Sao Paulo 1: 56. 1969; Falcao, Atas Soc. Biol. Rio de Janeiro 14: 153.
1971; Fl. II. Cat. BALA. 11. 1975.
Archimedea Leandro do Sacramento ex A. St.-Hilaire, Ann. Sci. Nat. Bot. Ser. 2. 7: 32.
1837.
Lepidophytum Hooker f. in Lindley, Veg. Kingd. ed. 3: 90. 1853 nom. nud.; in
Bentham & Hooker f., Gen. P1. 3: 238. 1880 pro. syn. "Lepidophyton. "

Fleshy, extremely starchy root parasites. Tuber more or less depressed or

globose .to cylindrical elongated; surface coarsely tessellately cracked, scaly or
not in upper part. Stem with sterile lower part extremely short or almost absent
without a sheath at base, covered by triangular, scaly leaves. Inflorescence
once branched, unisexual or bisexual with female branches below and male
branches above, when young covered by angularly-ovate bracts. The transition
in shape and size from scales on the tuber through scaly leaves on the stem to
scaly bracts on the inflorescence is remarkably smooth. Bracts deciduous,
46 Flora Neotropica

subpeltate, pelta angularly-ovate, each subtending a branch. Male flowers

with an inconspicuous, 2(-1)-lobed perianth. Stamens 2, free; filaments short,
filiform, anthers basifixed, linear-oblong, longitudinally 4-celled, dehiscing
lengthwise. Pollen grains prolate, tricolporate, medium-sized. Female flowers
sessile, densely arranged and covering the branch completely, prismatic with
+ truncate top part, with or without an inconspicuous, 2-lobed perianth.
Styles 2, inserted in a small cavity, filiform; stigma capitellate. Fruit a small,
1-seeded achene.
Type species. Lophophytum mirabile Schott & Endlicher, Melet. Bot. 1,
t. 1. 1832.
Distribution. Colombia, Peru, Bolivia, N Argentina, Paraguay, Brazil
(Acre and SE).
Four species have been described. Of these I consider three as distinct, ie
L. leandri, L. weddellii and L. mirabile. Lophophytum bolivianum is dis-
tinguished from L. mirabile only by the shape of the anther. It is treated here
as a subspecies of L. mirabile, because it is allopatric.

Key to the Species of Lophophytum

1. Femaleflowerseach subtendedby a bract,the swollenapex of whichprotrudesabove the
layerof flowers;some of the lowerflowersin the lowermalebrancheseachsubtendedby a
rudimentarybract;male flowerwith two perianthsegments,the lowerone large,the upper
one small;monoecious. 1. L. leandri.
1. Femaleandmaleflowersnot subtendedby bracts;monoeciousor dioecious.
2. Rimof ovarywith2 conspicuousmedianmamillae(perianthsegments)aboveandbelowa
centralcavity;male flowerswith filamentsinsertedbetween2 median,smoothmamillae
(perianthsegments);dioecious. 2. L. weddellii.
2. Rim of ovaryregularlytruncate,withoutmamillae;male flowerswith lower,largeperi-
anthsegmentonly;monoecious. 3. L. mirabile.

1. Lophophytum leandri Eichler in Martius, Fl. Bras. 4. 2: 59, t. 15. 1869; in

DC., Prod. 17: 129. 1873; Engler, Nat. Pflanzenfam. Ed. 1.3. 1: 255,f.
161D. 1889; Harms, Nat. Pflanzenf. ed. 2. 16b: 326, f. 164D. 1935;
Burkart, Darwiniana 9. 1: 170. 1949; Sleumer, Bot. Jahrb. Syst. 76: 271.
1954; FalcRo, Rodriguesia 25(37): 136. 1966; Atas Soc. Biol. Rio de
Janeiro 14: 154. 1971; Fl. II. Cat. BALA 15. 1975.
Fig 19, Fig 6 (pollen), Fig 21 (distr.).
ArchimedeapyramidalisLeandrodo Sacramentoex A. St.-Hilaire,Ann. Sci. Nat. Bot. Ser.
2. 7: 32. 1837p.p. quoadspec., excl. descr.
Lophophytum mirabile auct. non Schott & Endlicher: Weddell, Ann. Sci. Nat.
Bot. S6r.3. 14: 185,f. 31-33. 1850;Hookerf., Trans.Linn. Soc. London22: 30, 48. 1856
p.p. quoadspec. Leandrodo Sacramento;Platzmann,Leipz.I11.Zeit. 1865n.v.

Description based on dry material. Fleshy, brownish to maroon root

parasites. Tuber subterranean, very large, 1.5-5 x 4.2-17 cm, starchy; surface
coarsely tessellately cracked into polygonate warts up to 2 cm diam., these
with radial cracks. 1-4 inflorescence-bearing stems appear from upper part of
tuber. Inflorescence 10-24 cm long, 4-7 cm wide; lower female part 1.5-5.5 cm,
upper male part 6-19 cm. Branches each subtended by an early caducous, scaly
Balanophoraceae 47

bract. Bractssubpeltatewith petiole 1-1.3cm long, 0.1-0.4 cm wide, flattened,

and pelta angularly-ovate,1.5-3.5 cm long, 1.2-1.6 cm wide. Male branches
0.7-1.5 cm long and 0.9-1.4 cm widewith flowersexpanded.Each flowercon-
sists of 2 median perianthsegmentsbetweenwhich 2 stamensappear;lower
segmentmuch largerthan upperone; filament 1.9-2 mm long, anthersbasi-
fixed, 2.7-4 mm long and 1.2 mm wide;some of the lowerflowersin the lower-
most male brancheseach subtendedby a rudimentarybract similarto the
bractssubtendingthe femaleflowers. Pollen grainsspheroid(-prolate),tricol-
porate,P 27, 8 /m, E 24,3 Am,nuclei2. Femalebranchesincl. of flowers1-1.8
cm long and 0.7-1.2 cm wide, denselycoveredby flowers, apicallyrounded.
Each flower subtendedby a clavatebract3-3.4 mm long, 0.1-0.2 mm wide in
lowerpartand 0.6-1 mm acrossin the abruptlywidenedtop part.Flowerswith
perianthadnateto ovary, obconical-prismatic,3 mm long and 0.9-1 mm wide
in truncatetop part, perianthlobes as 2 median, inconspicuousmamillae;
styles 2, 1.3 mm long and 0.1 mm wide with capitellatestigma0.2 mm wide,
appearingfrom 2 centralcavities, diverginglaterally.Fruit a small, 1-seeded
achene.
Type. Syntypes:Leandrodo Sacramentosn, Rio de Janeiroand Platz-
mann sn, Bahia Paranagua,not located. The name is typified by Eichler's
illustrationin Mart., Fl. Bras. 4. 2: t. 15. 1869, which was based on Platz-
mann'sdrawingof a livingspecimen.
Distribution.Mixedtropicalforest in SE Braziland NE Argentina.Most-
ly flowering July-September(-January).Hosts recorded: Anadenanthera
macrocarpa (Benth.) Brenan, Apuleia sp., Piptadenia rigida Benth. (all
Leguminosae).

BRAZIL. Hubbard sn (K); Moller sn (B); Schiffner & Wettstein sn (M, W). ParanA:Hatsch-
bach 9842 (US); 19606 (C). Santa Catarina: Mueller 340 (K); Smith & Klein 7287 (US). ARGEN-
TINA. Missiones: Grondosa et al 1709 (SI); Krapovickas et al 15509 (C, F, MO, P); Marunak 633
(AK); Schwabe sn (COL). Corrientes: Krapovickas et al 28596 (C, MO).

Leandrodo Sacramento'scollectionsfrom Rio de Janeiro"nearthe river

Guapiacuand at Tijuca" constitutethe type materialof Archimedeapyrami-
dalis Saint-Hilaire(1837).As alreadymentioned,this materialcould not be lo-
cated for the presentmonograph.However,it was studiedby Weddell(1850),
indirectlyby J. D. Hooker (1856), and by Eichler(1869), all of them paying
specialattentionto the bractsubtendingeach female flower, a uniquefeature
in the family. Eichler(1869)maintainedthat the type materialconsistedof two
discordantelements, one belongingto Lophophytummirabile,the other de-
scribed by Eichler (l.c.) as a new species, Lophophytumleandri. It is quite
clearthat Saint-Hilaireoverlookedthe bracts,whendescribinghis Archimedea
pyramidalis.The descriptionfits Lophophytummirabileprecisely,and Ar-
chimedeapyramidalisthus becomesa taxonomicsynonymundcrL. mirabile.
The reasonwhy Saint-Hilairedid not realizethat his taxon was at leastin part,
conspecificwith L. mirabileSchott& Endl. is, I think, that theseauthorspub-
lished in the rare work, MeletemataBotanica, which was unknownto Saint-
Hilaire. Only 50-60 copies were printed, and they were not for sale. I have
48 Flora Neotropica

1 ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
k.(i~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
H L
D~~~~~~~~~~~~~~~~~~~~~~t~;~?:~

FIG 19. Species of Lophophytum. A-G, L. leandri (Krapovickas et al 15509) original. A, habit,
tuber were attached to host rot at scar to the left x 0.5; B, male brach, basal end to the right x 3;
C, portion of male branch with distal end upwards, anthers removed but filaments left in natural
position, in the center one male flower with its small upper perianth segment and its large lower
one, two filaments inbetween and the whole flower exceptionally subtended by a clubshaped bract
(compare E-G) x 9; D, two open anthers x 8; E, female branch, basal end to the right, the wid-
of bracts
ened distal ends of bracts subtending female flowers almost concealing these xx 3; F, female branch
female flowers
 Balanophoraceae 49

marked the approximate locality of Leandro do Sacramento's collection on

the distribution map for L. leandri. However, the species has to my knowledge
never been collected again in this area.

2. Lophophytum weddellii Hooker f., Trans. Linn. Soc. London 2: 30, 49, t.
9. 1856 emend. B. Hansen; Eichler in DC., Prod. 17: 129. 1873; Engler,
Nat. Pflanzenf. ed. 1. 3. 1: 255. 1889; Harms, Nat. Pflanzenf. ed. 2. 16b:
326. 1935. Fig 20, Fig 6 (pollen), Fig. 21 (distr.).

Description based on dry material and spirit material. Fleshy to almost

woody, dark-brown root parasites. Tubers subterranean, very large, in clumps
up to 60 cm diam., starchy; surface coarsely cracked into irregularly shaped
warts up to 2 cm across. Stem appearing from upper, scaly part of tuber.
Scales apparently arranged in 14-16 vertical ranks, triangular, 0.5 x 0.4 cm in-
creasing to 3.2 x 1.7 cm at transition from tuber to inflorescence-bearing
stem, where the scales become spirally arranged. Sterile part of stem very short
or absent. Inflorescence unisexual, 5-15 cm long and 5-7 cm wide including the
bracts. Branches each subtended by a persistent (?), scaly bract. Bracts subpel-
tate with petiole 1-1.8 cm long, 0.2-0.4 cm wide, and pelta angularly-ovate,
3.5-5 cm long, 1.6-2.7 cm wide. Male branches about 1 cm long. Each flower
consists of 2 median perianth segments of equal size between which 2 stamens
appear; filaments 1 mm long, anthers basifixed 3 mm long and 0.8 mm wide.
Pollen grains spheroid-subprolate, tricolporate, P 31.9 zm, E 25. 7 /zm, outer
layer of wall granular, loosening irregularly (artefact caused by acetolysis ?).
Female branches incl. of flowers 1.5 cm long and 1 cm wide, densely covered
by flowers, apically rounded. Flowers with perianth adnate to ovary, obconi-
cal-prismatic, 3 mm long and 1.5 mm wide above, perianth lobes present as 2,
median, conspicuous mamillae about 0.3 mm high and 0.8 mm wide; styles 2,
1 mm long and 0.1 mm wide with capitellate stigma 0.15 mm wide, appearing
from 2 central cavities, diverging laterally. Fruit a small, 1-seeded achene.
Type. Purdie sn, Colombia, not located. The name is typified by
Hooker's description and illustration in Trans. Linn. Soc. London 22: 30, 49,
t. 9. 1856.
Distribution. Humid tropical forests, alt. 500-1000 m, in Colombia and
Peru. The three flowering specimens studied were collected in April, October
and December. No host recorded.
COLOMBIA. Sander & Co. sn (K). PERU. San Martin: Plowman 5940 (C, GH); Schunke
5675 (F, S, U).

3. Lophophytum mirabile Schott & Endlicher, Melet. Bot. 1, t. 1. 1832:

seen from below x3; G, female flower and its subtending bract x 15. H-M, L. mirabile subsp
bolivianum, H, K-M (Ule sn 1911) I (Weddell 3631) original. H, portion of male branch with distal
end upwards, anthers and most of the filaments lost, upper perianth segment completely reduced
so that one flower is represented by the large lower segment and two stamens (compare C) x 9; I,
two stamens, anthers open x 8; K, portion of female branch, distal end upwards, styles lost x 9;
L, female flower, side view x 12; M, same seen from above, two styles have been inserted in center
of cavity x 12.
50 Flora Neotropica

jF EF H

:,,~~~~~~~~~~~~~~~~~~~~~~~~U~;

'_:. ..
.'-~.

vc/'
FI2. ohohyu wdceli.,aferHokr . 156, ,C Plwan590ad-
(Sc~~~~~~
~~~~~~hune57)oiia.A ai 05 ,batsbedn nlrsec rnh rn iwo
pett atx ;C h am,sd iw(cmaeFg1,1 n ) ;D
aebrnhbaled

donad;E
rnh
asledt
fml
h ih t x;F,par
ffml rnh aa n
downard
x1; G,femle lowe, sde iew,x15 H,the ame dital artx15

FIG 20. Lophophytum weddel/ii. A, after Hooker f. (1856), B, C (Plowman 5940) and D-H
(Schunke 5675) original. A, habit x 0.5; B, bract subtending inflorescence branch, front view of
peltate part x 1; C, the same, side view (compare Fig 1, 1B and IC) x 1; D, male branch, basal end
downwards x 7; E, female branch, basal end to the right, x 3; F, part of female branch, basal end
downwards x 10; G, female flower, side view, x 15; H, the same, distal part x 15.
Balanophoraceae 51

Hooker f., Trans. Linn. Soc. London 22: 30, 48. 1856 p.p. quoad spec.
Schott; Eichler, Bot. Zeitung (Berlin) 26: 551, t. 9, f. 16-18. 1868; in Mar-
tius, Fl. Bras. 4. 2: 46, t. 9-14. 1869; in DC., Prod. 17: 128. 1873; Engler,
Nat. Pflanzenf. ed. 1. 3. 1: 255, f. 161A-C, E-K. 1889; Glaziou, Mem.
Soc. Bot. France 1(3): 611. 1913; Hauman & Irigoyen, Anales Mus. Nac.
Hist. Nat. Buenos Aires 32: 58. 1923; Harms, Nat. Pflanzenf. ed. 2. 16b:
325,f. 164 A-C, E-K, 1935; Falcao, Rodriguesia 25(37): 137. 1966; Ange-
ly, Fl. Anal. e Fitog. Sao Paulo 1: 56. 1969; Falcao, Atas Soc. Biol. Rio
de Janeiro 14: 154. 1971; Martins f., Leandra 2: 99. 1972; Falcao, Fl. II.
Cat. BALA 12. 1975. Fig 19, Fig 7 (pollen), Fig 21 (distr.).
Archimedea pvramidalis Leandro do Sacramento ex A. St.-Hilaire, Ann. Sci. Nat. Bot. Ser.
2. 7: 32. 1837 p.p. quoad spec., incl. descr. Type. Leandro do Sacramento sn., Rio de
Janeiro, Brazil, not located.

i \f;D taS AL
E

'o
4x ;:
Y_
/I LT~~~~~~~~o
20~

FIG 21. Distribution of Lophophytum. L. leandri, ; L. weddellii,*: L. mirabile subsp

mirabile, *;
mirabile, mirabile subsp
e; L. mirabile bolivianum, "",.
subsp bolivianum,
52 FloraNeotropica

Descriptionbased on dry materialand spirit material.Fleshy, brownish

root parasites.Tubers subterranean,3-20 cm long and 4-13 cm across, sub-
sphericalto subcylindrical,starchy;surfacecoarselycrackedinto irregularly
shapedwarts. Inflorescence-bearing stem appearingfrom upper,scaly partof
tuber, 12-30cm long, lower sterilepart very short or absent, coveredby tri-
angularscaly leaves. Inflorescencebisexualwith female branchesbelow and
male ones above, each branchsubtendedby a scaly bract. Bractsubpeltate,
with petiole 0.3-0.5 cm long and 0.1-0.2 cm wide, and pelta angularly-ovate,
1.5 cm long and 1 cm wide. Male branches0.5-1.5 cm long and 1-2 cm wide
with flowers expanded, conspicuouslylongitudinallyfurrowedby pressure
from the anthers, only lower, large perianth segment present, upper one
apparentlycompletelyreduced;stamens2, filaments0.9-1.3 mm long, anthers
basifixed,3-4.5 mm long and 1.2-1.8mmwide. Pollengrainsspheroid-subpro-
late (-perprolate),tricolporate,P 32-36l/m, E 20-25.5 tim,2-nucleate,in some
specimensheteromorphicwith a small proportionof hexa-colporate,tetra-
hedralgrainstogetherwith normalones withinthe sameanther.Femalebran-
ches includingthe flowers0.7-2 cm long and 0.5-1.5 cm wide, denselycovered
by flowers, apically rounded. Flowers with perianth adnate to ovary,
obconical-prismatic,3.3 mm long and 1.8 mm wide at truncatetop part,
segmentsinconspicuousand hardlyvisible;styles2, 0.6 mm long and 0.1 mm
wide, with capitellate stigma 0.15 mm diam., appearing from 2 central
cavities,diverginglaterally.Fruita small, 1-seededachene.

Key to the Subspeciesof Lophophytummirabile

1. Stamens with thecae of the same length. 3a. subsp mirabile.
1. Stamens with thecae unequal, anterior one 3/4 the length of posterior one, in at least some of
the anthers. 3b. subsp bolivianum.

3a. Lophophytummirabilesubspmirabile

Stamenswith thecaeof the samelength.

Type. Schottsn, Brazil,nearSebastianopol,1820,not located. The name
is typifiedby Schott& Endl., Melet.Bot. 1, t. 1. 1832.
Distribution. Humid tropical forests up to 500 m alt. in SE Brazil.
Floweringspecimensmostly collectedin July-October.Hosts recorded:Inga
sp., (Leguminosae).
BRAZIL. Bahia: Curran 22 (GH, S, US). Minas Gerais: Barreto 7431 (F); Sucre 10218 (RB).
Rio de Janeiro: Emmerich sn, Tingua (R); Glaziou sn, Serra dos Orgaos (BR); Netto sn, 1869 (P);
Occhioni 982, 983 (RB); Pabst 5416 (HB); Peckolt sn, Canta Gallo 1862 (BR); Planchon sn, 1864
(K). Santa Catharina: Glaziou 306 (BR).

3b. Lophophytum mirabile subsp bolivianum (Weddell) B. Hansen, comb.

nov.
Lophophytum bolivianum Weddell, Ann. Sci. Nat. Bot. S6r. 3. 14: 185, t. 10, f. 29, 30.
1850; Hooker f., Trans. Linn. Soc. London 22: 30, 48. 1856; Eichler in DC., Prod. 17:
129. 1873; Engler, Nat. Pflanzenf. ed. 1. 3. 1: 255. 1889; Harms, Nat. Pflanzenf. ed. 2.
16b: 326. 1935; Macbride, Fl. Peru 2: 428. 1936; Burkart, Darwiniana 9. 1: 172. 1949;
Sleumer, Bot. Jahrb. Syst. 76: 272. 1954.
Balanophoraceae 53

..r'

,-I

FIG 22. Lathrophytum

LathrophYlum peckoltii. A-H, Peckolt sn 1886 after Hansen (1976), I1 and K (Pereira
5645) original. A, habit of young specimen, tuber lost, inflorescence covered by peltate parts of
bracts x 1; B, bract subtending female branch, side view x 6; C, the same, front view of pelta x 6;
D, bract subtending male flower, from below x 6; E, male flower, side view x 6; F, the same,
from above x6; G, female branch, side view x6; H, female flower, side view x 17.5; 1, habit of
flowering specimen with host root, tuber, volva lobes, lower female part of inflorescence showing
peltas of branches, upper male part with anthers opened, some 30 flowers have lost the anthers
x 1; K, four male flowers with anthers open, three anthers lost, seen from distal end x 2.5
54 Flora Neotropica

Stamens with one theca conspicuously shorter than the other, at least in
some of the anthers.

Type: Weddell 3631, Bolivia, Prov. de Cordillera, fl. November (lec-

totype, P; isolectotype, K).
Distribution. Evergreen forests up to 700 m alt. in Brazil (Acre), Bolivia,
Paraguay, and Argentina. Flowering (August-)December-February(-April).
Hosts recorded: Enterolobium sp., Piptadenia sp., and Pithecolobium sp. (all
Leguminosae).
BRAZIL. Amazonas: Ule sn, Alto Acre (B). BOLIVIA. Pflanz 8, Villamontes, Sotos (S).
PARAGUAY. Balansa 2717, Paraguari (K, P). ARGENTINA. Salta: Golbach sn, Oran (LIL);
Krapovickas 1687 (SI); Lourteig 690 (LIL, NY); Venturi 6612 (US).

5. Lathrophytum Eichler, Bot. Zeitung (Berlin) 26: 550. 1868; in Martius, Fl.
Bras. 4. 2: 64. 1869; in DC., Prod. 17: 130. 1873; Hooker f. in Bentham &
Hooker f., Gen. P1. 3: 239. 1880; Engler, Nat. Pflanzenf, ed. 1. 3. 1: 255.
1889; Harms, Nat. Pflanzenf. ed. 2. 16b: 328. 1935; Falcao, Rodriguesia
25(37): 139. 1966; Atas Soc. Biol. Rio de Janeiro 14: 153. 1971; Fl. I1.
Cat. BALA 24. 1975; B. Hansen, Bot. Tidsskr. 71: 77. 1976.
Lathrophytium Swart, Ind. Nom. Gen. 1965, orth. mut.

Fleshy parasites forming subterranean tubers on the roots of trees. Tubers

ellipsoid to subspherical, with a rough, warty surface, containing starch
grains. Stem and inflorescence breaking through a volva. Volva continuous
with surface of tuber, adnate to lower part of stem, bursting into 2-3 lobes ap-
pressed to stem and inflorescence. Stem leafless, glabrous or with a few peltate
scales above and rudimentary branches. Inflorescence spadix-like, bisexual,
when young covered by numerous, marginally connivent, rhombic to irregu-
larly polygonate top parts of peltate, early caducous bracts, during anthesis
completely destitute of bracts. Male-flowers in upper part of inflorescence,
without a perianth. Stamens 2, filaments absent, anthers 2-locular, each locule
2-locellate. Female-flowers densely arrangedon slender branches, which termi-
nate in a peltate top part, perianth absent, styles 2. Fruit a small, 1-seeded
achene.
Type species. Lathrophytum peckoltii Eichler, Bot. Zeitung (Berlin) 26:
550, t. 9, f. 1-10, 12-15. 1868.
Distribution. Brazil (Prov. of Rio de Janeiro).

1. Lathrophytum peckoltii Eichler, Bot. Zeitung (Berlin) 26: 550, t. 9, f. 1-10,

12-15. 1868; in Martius, Fl. Bras. 4. 2: 64, t. 16. III,f. 1-14. 1869; in DC.,
Prod. 17: 131. 1873; Engler, Nat. Pflazenf. ed. 1. 3. 1: 256. 1889; Harms,
Nat. Pflanzenf. ed. 2. 16b: 328. 1935; Falcio, Rodriguesia 25(37): 139.
1966; Atas Soc. Biol. Rio de Janeiro 14: 153. 1971; Fl. II. Cat. BALA 25.
1975; B. Hansen, Bot. Tidsskr. 71: 78,f. 1-2. 1976.
Fig 22, Fig 8 (pollen), Fig 23 (distr.).
Balanophoraceae 55

Fleshy parasites forming tubers on the roots of trees. Total length up to 16

cm. Tuber up to 4.5 cm long and 4.6 cm wide, ellipsoid to subspherical, with a
rough surface caused by numerous warts about 0.5 mm diam., with a few
larger warts 1.5 mm diam. Volva adnate to stem in lowermost 1-3.5 cm, com-
pletely enveloping young inflorescences, surface as on tuber except for larger
warts, later bursting into a few long-persistent obtuse lobes; 2.5 cm long and
2.4 cm wide at base. Stem above insertion of volva-lobes 1.6-3.2 cm long, 1.6-2
cm diam., cylindrical. Inflorescences when young ellipsoid, 4.3-6.2 cm long,
3.3-4.0 cm diam., later becoming 8-10 cm long and 1.4-2 cm wide incl. flowers,
lower female part 0.9-3 cm long with 30-50 branches, upper male part 1.4-7 cm
long with about 200 flowers. Bracts in lower part of inflorescence up to 7.1
mm long with a narrow petiole 0.7-1.2 mm wide and a fleshy pelta, which is
rhombic or irregular-polygonate, about 4.5 mm diam., with irregularly fringed
border, upwards on the inflorescence decreasing in size but getting a relatively
thicker petiole, each bract subtending either one female branch or one male
flower. Male flower without a perianth, inserted on an obtuse, conical axis
about 3.5-4 mm wide and 3.5 mm long. Stamens 2, filaments absent; anthers
inserted laterally about 1 mm apart a little below the blunt top of axis, length
1.3 mm, lateral width 1.6 mm, median width 1.1 mm, dehiscing laterally by
two longitudinal slits, 2-celled and 4-locellate. Longitudinal axis of the anther
in its natural position horizontal with one slit facing upwards and one down-
wards, later on deflexed with walls of loculi completely recurved. Pollen grains
spheroidal to suboblate, tricolporate, nuclei 2. Female branch 6-8 mm long
and 1 mm wide with top part extended to a subcircular pelta 4-5 mm diam.
Female flowers 50-100 densely arranged on narrow part of branch, more or
less hidden by the pelta, without a perianth, about 1-1.2 mm long and 1 mm
wide, truncate, with 2 (rarely 3) styles inserted at top. Style 0.3 mm long and
0.1 mm wide with slightly swollen, stigmatic top part. Fruit a small, 1-seeded
achene.
Type. Peckolt sn, Brazil, near Canta Gallo, Rio de Janeiro, 1867
(holotype, BR).
Distribution. In dense forests, parasitizing roots of trees. Hosts recorded:
Senefeldera sp., (Euphorbiaceae), Myristica bicuhyba Schott (Myristicaceae).
Flowering Oct.-Nov.
BRAZIL. Rio de Janeiro: Peckolt sn, 1886 (B, C, M); Glaziou sn, 1886 (K); Pereira 5645
(RB).

6. Ombrophytum Poeppig [Leipz. Literaturz. 1833: 1874. 1833] ex Endlicher,

Gen. Plant. 73. 1836; Poeppig & Endlicher, Nov. Gen. 2: 40, t. 155. 1838;
Hooker f., Trans. Linn. Soc. London 22: 30, 49. 1856; Eichler, Bot.
Zeitung (Berlin) 26: 550, t. 9, f. 11. 1868; in Martius, Fl. Bras. 4. 2: 61.
1869; in DC., Prod. 17: 129. 1873; Hooker f. in Bentham & Hooker f.,
Gen. PI. 3: 239. 1880; Engler, Nat. Pflanzenf. ed. 1. 3. 1: 255. 1889;
Harms, Nat. Pflanzenf. ed. 2. 16b: 326. 1935; Macbride, Fl. Peru 2: 428.
1936; Falcao, Fl. 11.Cat. BALA 34. 1975.
56 Flora Neotropica

Juelia Asplund, Svensk Bot. Tidskr. 22: 273, J: 1-3, t. 3. 1928; Harms, Nat. Pflanzenf.
ed. 2. 16h: 328. 1935; Macbride, Fl. Peru 2: 429. 1936; Pfister, Bol. Soc. Biol. Concepcion
22: 15. 1947; Sleumer, Bot. Jahrb. Syst. 76: 272, t. 16-18. 1954.

Fleshy, extremely starchy root parasites. Tuber spheroid to more or less

depressed ellipsoid; surface finely verrucose. Stem with a variously shaped, 2-
3 lobed volva or sheath at base. Inflorescence once branched, unisexual and
then female or bisexual with female branches below and male branches above.
Bracts subtending the branches variously peltate or reduced and clavate, cadu-
cous. Branches up to 20 mm long, cylindrical with a peltately widened apical
part. Male flowers apparently without a perianth. Stamens 2, free; filaments
short, filiform; anthers basifixed, elliptic, 4-celled, dehiscing lengthwise. Pol-
len grains spheroid to prolate, tricolporate to pentadecacolpate or quite irregu-
lar, rarely tetraloxocolporate or hexacolporate and tetrahedral, medium to
large sized. Female flowers sessile, densely arranged on lower, narrow part of
branch, prismatic due to mutual pressure, with + truncate apical part and ap-
parently no perianth. Styles 2, inserted apically in a small cavity, filiform;
stigmas capitellate. Fruit a small, 1-seeded achene.
Type species. Ombrophytum peruvianum Poepp. & Endl., Nov. Gen. 2:
40, t. 155. 1838.
Distribution. Ecuador, Peru, Brazil (W Amazonas, Acre) Bolivia, Chile,
Argentina, Galapagos Islands.
Until recently two names had been validly published within Ombrophy-
turn, ie 0. peruvianum and 0. zamioides. These species are considered conspe-
cific here because of obvious similarities in the overall habit and in the struc-
ture of the branches of first order, as expressed in drawings of the now lost
type-material. Efforts have been made to obtain material from the type locali-
ty for the neotypification of 0. peruvianum, but so far without success.
Another species collected in Brazil (Acre) by Ule 1911 was known under the
nomen nudum: 0. ulei. Recently it was proved, that Poeppig quite obviously
collected this species at the same time as his 0. peruvianum around 1830.
Based on Poeppig's observation it was named 0. violaceum Hansen (1977). A
third species, 0. microlepis, is described here based on material collected by
Ule 1911 and later by Schultes 1946 and Schunke 1970 and supported by a col-
our slide by F. Halle 1974. A fourth species, 0. subterraneum, is established
here by the transfer of Juelia subterranea Asplund to Ombrophytum. In the
original generic description of Ombrophyum Poeppig & Endlicher wrote
"ovaria . . . bilocularia." In the specific description, however, they stated
"ovaria . . . unilocularia, uniovulata." Their illustration shows a bilocular
ovary, but the legend runs "7) Idem (ie the ovary) horizontaliter sectum, (er-
rore Chaleographi biloculare)." Asplund found in his material from Bolivia
unilocular ovaries with one ovule and described a new genus, Juelia with the
species J. subterranea. However, I do not follow Asplund and subsequent
authors in paying so much attention to what is obviously the "engraver's er-
ror." Even if this minor difference was a reality, the morphological similarities
between the Juelia-entitites and the three species of Ombrophytum mentioned
above are so striking, that a transfer of Juelia is fully justified. When assisting
Adsersen (1976) in the identification of his Galapagos material I anticipated
Balanophoraceae 57

i ac 70 so so
.5

av

' '
~ S .'*' 5
SINU$SOIALPROJECTION
'-; ... ...; . ^_ !""""''^) '*) / '"**Y/-'^" V1*
0

in
~ I 2 . sriuiono
ahrpyu ipcoti, ] O
mropyu vi laeu,
recr , o

literature, and Ombrophytum microlepis,.

Intensive collecting in southern Peru, in Bolivia and Argentina may throw

light on this problem.It is highlyprobablethat the 0. subterraneumformsare
apomictic,some of them havingirregular,non-functionalpollen, otherscom-
pletelylackingmaturepollen grains.This mightalso explaintheiradaptability
to a wide varietyof habitatsfrom evergreenforest on the Equator(Galapagos)
to Andinepunasabout 27?Slat. (Argentina).

Key to the Speciesof Ombrophytum

1. Male branches with 4-8 opposite decussate flowers; bracts hexangular, violet, obviously im-
bricate. 1. 0. violaceum.
58 Flora Neotropica

1. Male branches with 10-50 alternate flowers; bracts not hexangular, not obviously imbricate,
sometimes inconspicuous.
2. Bracts inconspicuous, almost clavate with filiform petiole and a short, obtriangular
blade; female part of inflorescence usually 1/4-1/3 the length of male part. 2. O. microlepiN.
2. Bracts conspicuous, peltate with large, variously shaped peltas; female part of inflores-
ence 1/2 the length or more of male part.
3. Bracts with almost terete petiole and ? regularly crenate pelta; female part of inflores-
cence 1/2 the length of or equal to male part. 3. 0. peruvianum.
3. Bracts with a flattened petiole and an irregularlyincised, mostly oblique pelta bearing a
subulate or biforked process in the centre; female part of inflorescence mostly much
longer than male part or inflorescence entirely female. 4. 0. euhierruneum.

1. Ombrophytum violaceum B. Hansen, Bot. Tidsskr. 71: 231,J: 1-3. 1977.

Fig 24, Fig 8, 9 (pollen), Fig 23 (distr.).
OmbNrophyvln ulei Harms, Nat. Pflanzenf. ed. 2. 16bh:328, ./. 165 A-D. 1935, s. descr.
latino; Macbride, Fl. Peru 2: 429. 1936.
Otnhrophvtumln peruviunul/ auct. non Poeppig & Endlicher: Wurdack, Gard. .1. N.Y. Bot.
(lard. 14: 145. 1964.

Description based on dry material and color slide.Violet and yellowish,

monoecious plants (inflorescences bisexual). Total length up to 16 cm. Tuber
spheroidal, 1.8-3.5 cm diam., by a constriction passing into a volva, surface of
both yellowish light-brown, irregularly verrucose by small warts 0.5-1 mm
across. Volva 3-6 cm long, narrowly funnel shaped, violet inside, bursting ir-
regularly upwards into 2 or 3 lobes. Inflorescence 4.5-13 cm long, lower female
part 1.5-6 cm long, upper male part 3-7 cm long, up to 2.5 cm across inclusive
of flowers. Inflorescence branches each subtended by a caducous peltate bract
with petiole 3-5 mm long and 0.5-1 mm across, terete or slightly flattened,
peltas red-brown to violet, flat, thin, more or less hexagonal, densely imbri-
cate, each pelta with lower 1/3 free and upper 2/3 covered by the lower parts
of the peltas of three neighboring bracts inserted at higher levels. Male bran-
ches composed of 4-8 decussately arranged flowers, 1.5-4 mm long, 1-3 mm
wide, with a distinct, flattened, circular apical part 0.5-1.5 mm diam. Each
male flower appearing on the branch as a reduced axis with two shallow pits
laterally on either side of a more or less conspicuous mamilla, one stamen at
each pit; perianth absent. Stamen with filament 0.25-0.5 mm long; anther
1-1.25 x 0.5-1 mm. Axis of inflorescence between male branches densely
mamillate. Pollen grains spheroid, tricolporate, P 26.1 /xm, E 24.2 /m, nuclei
2. Female branches 4-6 mm long, 1-1.5 mm wide at fertile part, peltate apical
part 2-4 mm diam. Female flowers densely arranged on narrow part of branch,
without a perianth. Ovary slightly obconical, 1.2-1.5 mm by 1 mm; styles 2,
0.5 mm long, diverging from a shallow pit at the truncate apical part of the
ovary; stigmas capitellate.
Type. Wurdack 2360, Peru, Dpt. de Loreto, Prov. de Alto Amazonas,
Cerros Campanquiz at Pongo de Manseriche, 19-21 Oct. 1962 (holotype, US;
isotype, NY).
Distribution. Tropical rain forest at low altitudes (200-500) in Ecuador,
Peru and Brazil (Amazonas). Flowering October to December. Parasitizing
roots of trees; host not known.
Balanophoraceae 59

I! E
I

i 1

,,/

?
" , .' -.' G"

'?r :L' ~ '

FIG 24. Species of Ombrophyfum. A-E, 0. violaceurn, A-C (Wurdack 2360) and D, E (Mexia
6337a) after Hansen (1977). A, habit, upper part of inflorescence lost, most female branches lost
x 1; B, peltate bract subtending branch of inflorescence, side view x 6; C, the same, front view of
pelta (compare Fig 1, 2B and 2C) x 6; D, male branch, basal end to the right x 6; E, the same,
seen from distal end x 6. F-K, 0. microlepis (Ule sn 1911) original. F, clavate bract subtending
branch of inflorescence, side view x 18; G, the same, from above x 18; H, the same, front view
x 18; I, male branch, basal end to the right x 4; K, the same, seen from distal end x 4.
60 FloraNeotropica

ECUADOR.Napo: Sparre13049(S). PERU. Loreto:Mexia6337a(F, UC). BRAZIL.Ama-

zonas:Ulesn, Nov. 1911(B).

2. OmbrophytummicrolepisB. Hansen,sp. nov.

Fig 24, 25, Fig 8 (pollen),Fig 23 (distr.).
Ombrophytum peruvianumauct. non Poeppig& Endlicher:Harms,Nat. Pflanzenf.ed. 2.
16b:328,f. 165P-Y. 1935.

Ab Ombrophytoperuvianoet Ombrophytoviolaceodiffertbracteolisax-
ium florescentiumlateraliumvalde diminutis,laminis bracteolarumtriangu-
laribus, non peltatis. Ab Ombrophytoviolaceo porro distinctumquod flores
in ramis lateralibusmasculinisplures quam octo sunt, plerumqueusque ad
viginti,et floribusalternantibus,numquamcruciatimoppositis.
Descriptionbased on alcohol material, dried material and color slide.
Whitishto light brownor flesh coloured,monoeciousplants(inflorescencebi-
sexual). Total lengthup to 20 cm. Tuberspheroidal,3-4 cm diam., by a con-
strictionpassinginto a volva, surfaceof both yellowishlight-brown,irregular-
ly verrucoseby smallwarts0.5-1 mm across. Volva up to 6 cm long, narrowly
funnelshaped,violet inside, burstingirregularlyupwards.Inflorescencewith
lower, sterilepart up to 6 cm long, intermediarfemale part up to 4 cm long,
and uppermale partup to 11 cm long, 2.5-3 cm acrossinclusiveof flowers.At
least some of the branchesof the inflorescencesubtendedby an inconspicuous
bract. Bract3-4 mm long with a filiformpetioleabout 3 mm long and 0.3 mm
wide, blade obtriangular,0.7-1 mm long, 1.5 mm wide, and 0.5 mm thick.
Male branches3-7 mm long, 2-3 mm wide, with a peltate, transverselyelliptic
to subcircular,slightlycrenateapicalpart about 4 mm wide. Male flowersup
to 20 in one branch, without a perianth.Stamens2, alternatelyinsertedin
fissureson the branch;filament1-1.2mm long; anther1-2mm long, 0.5-1 mm
wide. Axis of inflorescencebetweenmale branchesdenselymamillate.Pollen
grainsspheroid,tricolporate,P 22.1 Atm,E 22.2 t4m,nuclei2. Femalebranches
about 4 mm long, with peltateapicalpart as in male branches,narrowlower
partdenselycoveredby 50-80flowers.Femaleflowerswhitish,withouta peri-
anth. Ovary about 1 mm long and 0.5-1 mm wide; styles 2, 0.5 mm long,
divergingfrom a shallowpit at the truncateapical part of the ovary;stigmas
capitellate.
Type. Ule sn, Brazil,Acre, Alto Acre, Seringal,S. Francisco,Sept. 1911
(holotype,B).
Distribution.Humid tropicallowland forests in Peru and Brazil(Acre).
Floweringspecimensfrom April, September,Octoberand November. Host
recorded:a bignoniaceousclimber.
PERU. San Martin:Schunke3943 (F). Loreto:Schultes8615 (IAN). BRAZIL.Acre:Hall'
sn, Oct. 1974(photoonly).

3. OmbrophytumperuvianumPoeppig& Endlicher,Nov. Gen. 2: 40, t. 155.

1838;Hooker f., Trans. Linn. Soc. London 22: 30, 50. 1856;Eichlerin
Martius, Fl. Bras. 4. 2: 62, t. 16. 1869; in DC., Prod. 17: 130. 1873;
Engler,Nat. Pflanzenf.ed. 1. 3. 1: 255. 1889;Harms,Nat. Pflanzenf.ed.
Balanophoraceae 61

i
s,~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~

_ ,~~~~~~~~~~~~~~r

. .
:' ...
,. 't r fn , "'

.2r tX^
Fi ;4?r
K-~"' .l'
:~~~~~~~~~?.
' = <__~~~~~~~~
s' ?

:".~i[~ ;e. .. ..
Wi2E
t
r,?
'.~;?..
'liY ' ,,

.,-? .
,:'%.-

~
~~~~~~~~~~~~~.,~..!t
i~~.: ~~~~~~~~~~~~~~~~~~~~~~~~~
,..

;~~~~~~~~~~~
~.~g.
?~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
,. ::,
..::
?~t.,:
i~~~~~~~~~ij"?tl~~~~~~~~~~~~~~~~~~~~~~
:..::~~~ ','::, t,,.~',..:....~:::,.:.~
.??..:i??

FI 5 mrpyu
icreis,i phot.w F. Hal,rai,Are,
194 Ot
, po. . Hall, 17.<
. .
Brail . . .cre,.. . t.
FIG 25. Ombrophytum microlepis:..~,
62 Flora Neotropica

2. 16b: 328. 1935; Macbride, Fl. Peru 2: 429. 1936; Falcao, Fl. II. Cat.
BALA 6. 1975. Fig 26, Fig 9 (pollen), Fig 27 (distr.).
Omhirophylumlzamioides Weddell, Ann. Sci. Nat. Bot. Ser. 3. 14: 184, t. 10. 1850; Eichler,
Bot. Zeitung (Berlin) 26: 551, t. 9, f. 11. 1868; in Martius, Fl. Bras. 4. 2: 63, t. 16. 1869; in
DC., Prod. 17: 130. 1873; Engler, Nat. Pflanzenf. ed. 1. 3. 1: 255. 1889; Harms, Nat.
Pflanzenf. ed. 2. 16b: 328. 1935; Macbride, Fl. Peru 2: 429. 1936; Falcao, Fl. II. Cat.
BALA 38. 1975. Type. W'eddellsn, Peru, not extant; the name is typified by Weddell I.e.:
t. 10. 1850.

Description based on dry material, illustrations, and color slide. Whitish

to light flesh coloured, monoecius plants (inflorescence bisexual). Total length
up to 35 cm. Tuber spheroidal to depressed ellipsoidal, up to 13 cm wide.
Volva 5-10 cm long, irregularly 2-3 lobed, enveloping lower sterile part of in-
florescence. Inflorescence with lower sterile part 7-13 cm long and 3-7 cm
diam., intermediar female part 5-7 cm long, and upper male part 8-15 cm long.
Inflorescence branches each subtended by a caducous, peltate bract with a
terete or slightly flattened petiole 6-7 mm long and a conspicuously crenate,
subcircular pelta 7-8 mm diam. Male branches 5-10 mm long with 10-50
flowers alternately inserted in fissures, and a peltately widened, slightly cren-
ate apical part about 5-6 mm diam. Flowers without a perianth. Stamens 2,
filaments about 1 mm long, anthers about 2 mm long and 1 mm across. Pollen
grains spheroid to prolate, tricolporate, P 28 Asm, E 18-27 ztm. Female
branches 8-20 mm long, with 40-100 flowers densely arranged in the narrow,
terete or slightly flattened lower part; apical part peltately enlarged, pelta 6-8
mm diam., irregularly crenate-dentate often somewhat undulate and viscose.
Female flowers without a perianth, obconical, prismatic due to mutual pres-
sure. Ovary 1.5 mm long and up to 0.75 mm wide; styles 2, appearing from a
shallow pit at the truncate apical part of the ovary.
Type. Poeppig sn, "in sylvis densis Peruviae subandinae ad Cuchero" ca.
1830, studied at W by Eichler (1873), but not located there by Asplund (1928)
nor later. The name may be typified by the description and illustration in
Poepp. & Endl., Nov. Gen. 2: 40, t. 155. 1838. Attention should be drawn also
to the details illustrated by Eichl. in Mart., Fl. Bras. 4. 2: t. 16, f. 1. 1-4. 1869.
Distribution. In dense tropical rain forest up to 500 m alt. in Ecuador,
Peru and Brazil (Acre). Flowering September to November. Parasitizing roots
of trees; host not known.
&ECUADOR.Napo: Harling et al 7026 (GB); Madison et al 5450 (C). PERU. Weddell sn,
Prov. de las Cordilleras (illustration only). San Martin: Poeppig sn, Cuchero c. 1830 (illustration
only). BRAZIL. Acre: Halle sn Oct. 1974 (photo only); Prance et al 7664 (INPA).

4. Ombrophytum subterraneum (Asplund) B. Hansen comb. nov.

Fig 28, Fig 9 (pollen), Fig 27 (distr.).
Juelia subterranea Asplund, Svensk Bot. Tidskr. 22: 274, J: 1-3, t. 3. 1928; Harms, Nat.
Planzenf. ed. 2. 16b: 329. 1935; Macbride, Fl. Peru 2: 430. 1936; Pfister, Bol. Soc. Biol.
Concepci6n 22: 15. 1947.
FIG 26. Ombrophytum peruvianum. A, phot. F. Halle, Brazil, Acre, Oct. 1974; B, illustration of
type, after Poepp. et Endl. (1838); C, the same, female branch; D and E, female and male
branches drawn from type material at Vienna, after Eichler (1869).
Balanophoraceae 63

t~~~~~~~~~~~~~~~~~~~~~~~~~~~~~J. K -,
i
I"

it~~~~i

I~~~~~~~~~~~~~~~~~~~~~

/~~~~~~:/

'. ~'.3/. ? ? '

,.. 4'-

a
'-"
64 Flora Neotropica

Juelia lilloana Sleumer, Bot. Jahrb. Syst. 76: 276, f. 1-3, t. 18. 1954. Type. Sleumer 3037
Argentina, Catamarca, Mesada de las Rosas, 1600 m. alt., 10.2. 1952 (holotype, LIL).
Jielia meyeri Sleumer, Bot. Jahrb. Syst. 76: 276, t. 16, 18. 1954. Type. Meier 17236,
Argentina, Tucuman, Cruz Alta, Los Alderetes cerca del Rio Sali, 430 m. alt., 20.5. 1949
(holotype, LIL).
01mbrophvtum peruvianum auct. non Poeppig & Endlicher: Spegazzini, Anales
Soc. Ci. Argent. 77: 150. 1914; Hauman & Irigoyen, Anales Mus. Nac. Hist. Nat. Buenos
Aires 32: 59. 1923; Adsersen, Bot. Not. 129: 113. 1976 quoad sp. Galapag.

Description based on dry material, illustrations, and color slides. Whit-

ish to rose coloured plants with bisexual or female inflorescences. Total length
(10-)15(-26) cm. Tuber depressed ellipsoidal, (2-)4(-5) x (3-)6(-7.5) cm, yel-
lowish, verrucose. Volva (1-)3(-5) cm long, irregularly 2-3-lobed, enveloping
lower part of inflorescence. Inflorescence cylindrical, clavate or almost
spheroidal, with lower, sterile part (0.5-)5(-10) x (2-)3(-4) cm, intermediar fe-
male part (3-)7(-15) x (2.5-)5(-7) cm, and upper male part (1-)3.5(-7) x
(3.5-)5(-7.5) cm, subterranean or partly supraterranean when flowering. In-
florescence branches each subtended by a caducous, peltate bract with a +
flattened petiole 5-18 mm long, 1-3 mm wide at base, up to 5 mm wide in upper
part, pelta irregularly angular to crenate, developed obliquely downwards,
with a subulate to bifurcate or irregularly incised dent protruding upwards
from its lower part. Axis of inflorescence between male branches conspicuous-
ly mamillate. Male branches 3-8 mm long, 1.5-4 mm wide, with about 50
flowers alternately inserted in fissures, and a peltately, below obliquely widen-
ed apical part which is 2-11 mm wide with -? thickened and undulate margin.
Flowers without a perianth. Stamens 2, filaments 0.6-2 mm long, anthers vio-
let, 1.4-2.3 mm long and 0.6-1.5 mm wide. Pollen grains spheroid, tricolpor-
ate, P 28-32 /m, E 28-30 /tm, or irregular, sometimes pentadecacolpate, up to
60 Am diam. and probably non-functional, in the Galapagos population com-
pletely absent despite of normally developed, longitudinally dehiscing anthers.
Female branches (3-)11(-22) cm long, flattened and 1-4 x 5-8 mm wide, apical
part ? obliquely peltately widened, 2-4 x 5-8 mm, pelta with swollen ?i
undulate margin. Female flowers about 100, densely arranged in narrow part
of branches, without a perianth. Ovary obconical, prismatic due to mutual
pressure, 1-2 mm long and 0.5-1 mm wide in upper part; styles 2, 0.5-0.75 mm
long, appearing from a shallow pit at the truncate apical part of the ovary.
Type. Asplund 2568, Bolivia, Pacajes, Rio Caranguilla 10 km S of
Ulloma, alt. 3850 m, 21.2. 1921 (lectotype, S; isotype, UPS); Asplund 3873,
ibid., 4.5.1921 (paratype, S, UPS).
Distribution. Andine mountain zone in dry bushland (puna) up to 4000 m
alt in Bolivia and N Argentina, occasionally in evergreen and deciduous moun-
tain forests 1000-2000 m alt., rarely down to 430 m alt. in Argentina (Catamar-
ca, Tucuman), Galapagos populations in evergreen mountain forst at 600-650
m alt. Flowering (November-)February-April(-June) in the Andes, April-
December in Galapagos. Parasitizing roots of trees, shrubs and herbs. Hosts
recorded: Baccaris grisebachii Hieron., B. petiolata DC., Eupatorium bu-
pleurifolium DC., Heterothalamus spartioides Hook. & Arn., Lepidophyllum
quadrangulare (Meyen) Benth. & Hook., Scalesia peduculata Hook. f., Tes-
saria absinthioides DC., Viguieriamollis Gris. (Compositae), Dioscorea mega-
Balanophoraceae 65

0"

-.- --- -.-

-- _

FIG 27. Distribution of OmbrophvYtunlperuvianunm,* and Omnbrophylumsubterraneum, *;

open symbols are localities of specimens cited in literature, not available for the present study.

lantha Gris. (Dioscoreaceae), Medicago sativa L. (Leguminosae), Nicotiana

glauca Grah. (Solanaceae).
GALAPAGOS. Isla Santa Cruz: Adsersen et al 243 (C), 244 (C), 1323 (C); Coppois sn, 9.9.
1974 (C); van der Werjf 597 (U). BOLIVIA. Pacajes: Asplund 6312 (US); Shepard 246 (GH, UC,
US). Cochabamba: Parker et al 37b (NY). Potosi: Troll 99 (M). CHILE. Antofagasta: Martin 486
(SI); PJister sn, 1946 (S). ARGENTINA. Jujuy: Cabezas SI 20263 (SI), SI 20341 (SI); Sleutner
3331 (LIL, SI). Catamarca: Sleumer3054 (LIL); Viirsoosn, 29.4. 1955 (K, L), sn, April 1956 (L).

Sleumer (1954) recognized three species, 1. Juelia subterranea, at high

altitudes, which is completely subterranean when flowering, with a sweetish
taste, much collected and consumed by the local inhabitants, 2. Juelia meyeri
at low altitudes also completely subterranean when flowering, with a bitter
taste, and 3. Juelia lilloana at intermediate altitudes, which is partly suprater-
66 Flora Neotropica

LI

,,N
"

:A

FIG 28. Ombrophytum sut;errar.um (Adsersenet al 243) original.A, habit x 0.5; B, bractsub-
tendinginflorescencebranch,side view x ; C, the same, front view x 3; D, two bractsfusedto-
getherx 3; E, the same, frontview x 3; F, malebranchseenfromdistalend in naturalpositionon
mamillatesurfaceof the mainaxis x 4; G, the sameremovedfrommainaxis, sideview x 4; H, the
sameseen fromproximalend x 4; I, K, stamensx 10;L, femalebranchin side view, distalend to
theleft x 4; M, the sameseenfromdistalend x 4; N, femaleflowerfromthe side x 15.
Balanophoraceae 67

ranean when flowering, with an astringent taste. I consider all these forms as
belonging in one variable species together with the Galapagos populations.
The morphology of the subtending bracts, of the male and female branches
and particularly their peltately widened apical part is strikingly uniform
throughout the entire distribution area. The variation is mostly found in the
shape and colour of the inflorescences and in the distribution of the sexes,
some individuals being entirely female, others bisexual, even within the same
population (Galapagos). These characters have proved to be most unreliable
for taxonomic purposes in other genera of Balanophoraceae also, e.g. within
Balanophora (Hansen 1972).

Balanophoraceae subfam. Balanophoroideae Engler, Nat. Pflanzenf. ed. 1. 3.

1: 261. 1889; Harms, Nat. Pflanzenf. ed. 2. 16b: 311. 1935; Angely, Fl.
Anal. e Fitog. Sao Paulo 1: 57. 1969.

Inflorescences not branched. Style 1. The plants containing a waxy con-

stituent in all organs, especially in the tubers, never starchy.
Type genus. Balanophora J. R. & G. Forst., Char. Gen. P1. 99, t. 50.
1776.

7. Langsdorffia Martius in Eschwege, J. Bras. 2: 179. 1818; Nov. Gen. Spec.

Plant. 3: 182. 1832; Schott & Endlicher, Melet. Bot. 12. 1832; Endlicher,
Gen. Plant. 74. 1836; Klotzsch, Linnaea 20: 460. 1847; Hooker f., Trans.
Linn. Soc. London 22: 29, t. 2. 1856; Eichler in Martius, Fl. Bras. 4. 2: 9.
1869; in DC., Prod. 17: 140. 1873; Hooker f., in Bentham & Hooker f.,
Gen. PI. 3: 236. 1880; Engler, Nat. Pflanzenf. ed. 1. 3. 1: 262. 1889;
Harms, Nat. Pflanzenf. ed. 2. 16b: 335. 1935; Macbride, Fl. Peru 2: 430.
1936; Standley, Fl. Costa Rica 410. 1937; Nevling, Ann. Missouri Bot.
Gard. 47: 307. 1960; Falcao, Rodriguesia 25(37): 133. 1966; Angely, Fl.
Anal. e Fitog. Sao Paulo 1: 57. 1969; Falcao, Atas Soc. Biol. Rio de
Janeiro 14: 152. 1971; Loefgrenia 57: 3. 1973; Fl. II. Cat. BALA 5. 1975;
Hansen in Fl. Mal. Ser. 1. 7: 804. 1976.
Senftenbergia Klotzsch & Karsten ex Klotzsch, Linnaea 30: 460. 1847. pro syn.
Thonningia auct. non Vahl: Liebmann, Forhandl. Skand. Naturf. 4. Moede 1884: 180. 1847;
Fawcett, Trans. Linn. Soc. Ser. 2. 2: 239. 1886.

Fleshy root parasites. Tuber rhizome-like, often more or less horizontal

and elongated, cylindrical, often branched, slightly swollen at point of contact
with host root. Inflorescence-bearing stems breaking through buds on the
tuber, leaving a lobed volva basally. Stem short or elongated, with numerous
spirally arranged, triangular lanceolate, scaly leaves. Inflorescences unisexual,
cylindrical in male specimens and hemispherical in female specimens, un-
branched. Male flowers pedicellate, trimerous. Perianth segments 3. Stamens
connate into a synandrium. Anthers 3, horseshoe-shaped with the bend up-
wards. Pollen grains spheroid, (3-)4(-5)-porate, medium-sized, 2-nucleate. Fe-
male flowers linear-prismatic, ebracteate. Perianth narrowly tubular; style 1,
exserted. Ovary apparently without a cavity. Fruit a small, 1-seeded achene.
Type species. Langsdorffia hypogaea Mart. in Eschw., J. Bras. 2: 179, t.
5. 1818.
68 Flora Neotropica

Distribution. Tropical rain forests and mixed forests from Mexico

through Central America to tropical South America.

1. Langsdorffia hypogaea Martius in Eschwege, J. Bras. 2: 179, t. 5. 1818;

Nov. Gen. Spec. Plant. 3: 182, t. 298. 299. 1832; Hooker f., Trans. Linn.
Soc. London 22: 29, 39, t. 2. 1856; Eichler in Mart., Fl. Bras. 4. 2: 10, t.
1-3. 1869; Warming, Vidensk. Meddel. Dansk Naturhist. Foren. Koben-
havn 1872: 131; Eichler in DC., Prod. 17: 140. 1873; Engler, Nat. Pflan-
zenf. ed. 1. 3. 1: 263,f. 167. 1889; Glaziou, Mem. Soc. Bot. France 1(3):
610. 1913. Harms, Nat. Pflanzenf. ed. 2. 16b: 336, f. 156b, 169. 1935;
Macbride, Fl. Peru 2: 430. 1936; Standley, Fl. Costa Rica 410. 1937; Nev-
ling, Ann. Missouri Bot. Gard. 47: 307, f. 89. 1960; Falcao, Rodriguesia
25(37): 134. 1966; Angely, Fl. Anal e Fitog. Sao Paulo 1: 57. 1969; Fal-
cao, Atas Soc. Biol. Rio de Janeiro 14: 153. 1971; Martins f., Leandra 2:
99. 1972; Falcao, Fl. II. Cat. BALA 6. 1975.
Fig 29, Fig 7 (pollen), Fig 30 (distr.).
Langsdorffia janeirensis Richard, Mem. Mus. Hist. Nat. 8: 412, t. 19. 1822; Schott &
Endlicher,Melet. Bot. 12. 1832;Harms, Nat. Pflanzenf. ed. 2. 16b: 336. 1935. nom.
superfl.illeg.
Thonningiajaneirensis (Richard)Liebmann,Forhandl.Skand.Naturf.4. Moede1844:180.
1847.nom superfl.illeg.
ThonningiamexicanaLiebmann,Forhandl.Skand.Naturf.4. Moede1844:180. 1847.Type
Liebmannsn, Mexico,Tepilongo,Tonagnia,Oajaca,alt. 3-4000ped. (holotype,C).
LangsdorffiamoritzianaKlotzsch& Karsten,Linnaea20: 461.1847;Harms, Nat. Pflan-
zenf. ed. 2. 16b:336. 1935.Type. Syntypes:Moritz1619,Venezuela,ColoniaTovar1865
(K, BM);Karsten22, ibid. (P).
LangsdorffiarubiginosaWeddell,Ann. Sci. Nat. Bot. S6r. 3. 14: 187, pi. 11, f. 48-51.
1850; Hooker f., Trans. Linn. Soc. London 22: 29, 42, t. 2. 1856. Type. Typified by the
protologuein Wedd.l.c.

Description based on wet and dry specimens. Fleshy root parasites with
yellowish to wine red colors. Tuber in male specimens mostly horizontal and
much elongated, in female specimens shorter and ascending, 0.7-1.9 cm diam.,
more or less pubescent. Inflorescence-bearing stems basally with a 5-8-lobed
volva. Total length from volva inclusive of inflorescence in males 3.1-13 cm, in
females 2-14 cm. Stems with numerous spirally arranged, triangular-lanceo-
late, scaly leaves 0.4-1.0 x 1.6-3.5 cm, margin ciliate. Inflorescences unisexu-
al, not branched; male ones cylindrical, 1.8-3.2 x 2.5-5.0 cm, with flowers ex-
panded; female ones hemispherical, 1.7-5.8 cm wide and 1.4-3.5 cm long.
Male flowers bracteate with a pedicel 6-10 x 1-1.3 mm. Bracts each reduced to
2 conical bodies with a slightly widened tip. Perianth segments 3, valvate,
cucullate, 1.8-1.9 x 1.2-1.5 mm. Stamens connate into a synandrium. Anthers

FIG29. Langsdorffiahypogaea.A-E (Schunke9617),F-I (Reinhardtsn) original.A, habitof two

malespecimensx 1; B, longitudinalsectionof malespecimenx 1; C, viewof maleinflorescence,
leavesremovedtogetherwith someof the maleflowers,sectionsof pedicelshatched,notethatthe
bractsarebiforkedexceptfor a verynarrowtransitionalzone immediatelyabovethe upperleaves
x 2; D, maleflower,side view x 3; E, the same,anthersopen x 3; F, malespecimenof elongated
form x 1; G, female plant with two inflorescencesx 1; H, longitudinalsectionof femaleplant
x 1; I, sectionof femaleinflorescenceshowingunistylarfemaleflowers x 15.
Balanophoraceae 69

,1
/1 1'
', . . . .' .

....
...
' ......

v/ \/'i yv"-- i~~~~~~.'N<- A

'L: ~ _i~ c~

F~~~~~~~~~~~

E `

D~~~~~~~~~

\l ^'"---.
'^*-- -..^----"^~~~~~~~~~~~~
A ?
,-'~ ~ -~-

'"K ^^>"J /^"'^-.

&F-" I .^^y ...
70 Flora Neotropica

- ^ *---- - I-
^ g~ -t^^ -- - _
-
...
- * -I 5- - .-.. - -

' ~
*> = :,.. ; t
._,,_ _- . . . . . . /
_ _

'
I - - --' 'l C* 1 '
o_
,. Y' , . 0

.---i--------
~~~~~~~~~~1T'opIc
I
- ....- .... -
/
- . .--:----jM---
* ^r
j - of C

FIG 30. Distribution of LangsdorJfia hypogaea.

3, connate into a head 1-1.7 x 1 mm, each anther 2-locular, horseshoe-

shaped with the bend upwards. Female flowers linear-prismatic, ebracteate,
densely arranged but free from each other, 1.5 x 0.25 mm. Perianth tubular,
irregularly lobed. Style exserted, 0.5-0.6 x 0.1 mm. Ovary apparently without
a cavity. Fruit a small, 1-seeded achene.
Type. Langsdorffsn, Brazil, N of Rio de Janeiro, Mandiocca, Serra de
Estrella, 1817 (lectotype, LE; isolectotype, H).
Distribution. In montane forests and Campos Cerrados (400-)800-2800
(-3100) m; apparently flowering all year round. Host plants recorded:
Geonoma sp., Iriartea sp. etc. (Arecaceae), Mimosa sp. (Leguminosae), Byr-
sonima sp. (Malpighiaceae), Trichilia sp. (Meliaceae), Ficus sp. (Moraceae).
Balanophoraceae 71

MEXICO. Oaxaca: Liebmann sn (C). COSTA RICA. Heredia: Burger & Stolze 6066 (F,
MO); G6mez 4034 (K). PANAMA. Chiriqui: Allen 4773 (BR, C, F, K, MO, P, S, U, UC); Darien:
Duke 5320 (MO); Gentry & Mori 14019 (MO). COLOMBIA. Karsten sn (P); Purdie sn (K);
Ocanae: Purdie sn (K); Chocb: Core 330a (US); Boyaca: Lawrance 251 (F, NY, US); Cundina-
marca: Cuatrecasas 5429 (US); Jaramillo 2660 (COL); Meta: Grant 10030 (US); El Valle Cauca:
Cuatrecasas et al 6263 (US); Sneidern 1263, 1264 (S); Agustin: Little 7684 (US); Huila: Little 8991
(US); Caqueta: Little 8214 (US). VENEZUELA. Cornelio sn, Avila (US); Merida: Bernardi 969
(NY); Fendler 1743 (BR, F, GOET, K, MO, NY, P, US); Amazonas: Maguire et al 28054 (K, NY,
P, RB, US); 28057 (NY, US); Aragua: Steyermark 89852 (F, K, US); Bolivar: Maguire et al 33832
(F, NY); 33841 (F, IAN, NY, U); 33927 (F, NY); Moore et al 9709 (UC); Stevermark 75657 (NY);
4Wurdack 34062 (NY); 34063 (NY); 34073 (F, MO, NY, UC). GUYANA. Schomburgk sn (BM).
ECUADOR. Imbabura: DrewtE-51 (US); Azuay: Camp 4418 (NY, US); Santiago Zamora: Acosta
Solis 5039 (F); Stevermark 53515 (NY). PERU. San Martin: Schunke 9617 (C); Chachapoyas:
Hutchinson & H'right5551 (US); Huanuco: Wolfe sn (F). BOLIVIA. Mapiri: Buchtien 1641 (US)
Chuquisaca: Orhigny 1188 (BR, P). BRAZIL. Para: Bamps 5198 (BR); Oliveira 145 (US); 196
(IAN); 278 (IAN); 446 (IAN); Silva 2661 (MG); Ceara: Benjamin 9284 (RB); Ducke 1674 (MA);
Ule 9025 (K); Pernambuco: Barreto 8799 (F); Reitz & Klein 7156 (L, M, UC, US); Mato Grosso:
Anderson et al 9754 (C, NY); Argent 6706 (U); Hoehne 3874, 3875, 3876 (R); Hunt & Ramos 6027
(MO); Irwin et al 16560 (GH, NY); 16848 (GH, NY, S); Malmnesn (S); Rojas 3836 (US); Goias:
Duarte 10281 (RB); Heringer 8859 (HB, NY, US); Rizzo 4706 (RB); Schwarzmayersn (SP); Bahia:
Duarte 9568 (RB); Minas Gerais: Anderson et al 8419 (NY); Claussen 1410, 1411 (P); Heringer &
Britoni 6974 (HB); Lund sn (BR); Macedo 2337 (MO, NY, SP, US); Mosen 4404 (C, S); Regnell sn
(BR); Reinhardt sn (C); St. Hilaire sn (P); Warmingsn (C); Espirito Santo: Luetzelburg sn (M);
Sao Paulo: Hashitnoto 150 (SP); Hoehne 27517 (GH, K, NY, S, SP, US); Loefgren 2513 (SP);
Sucre 3052 (RB); Rio de Janeiro: Baumbart sn (HB); Langsdorff sn (H, LE); Martius sn (M);
Miers sn (BM, K); Pabst 7301 (HB); Schott sn (W); Parana: Hatschbach 20771 (C).

ACKNOWLEDGMENTS

I would like to thank the directors and curators of the herbaria consulted
or from which material was borrowed. The herbaria are AAU, AK, B, BBS,
BM, BR, C, CAY, COL, F, FI, GB, GH, GOET, GZU, H, HB, IAN, INPA,
K, L, LE, LIL, M, MEXU, MG, MO, NY, P, R, RB, S, SI, SP, U, UC, US,
W. Photographs and colour slides were kindly put at my disposal by H. Adser-
sen, W. C. Burger, T. B. Croat, A. Gentry, L. D. G6mez, F. Halle, N. Halle,
G. Harling, J. P. Hjerting, L. Holm-Nielsen, L. Nevling, T. Plowman, B.
Sparre, R. T. Thorne, and J. J. Wurdack; I am most grateful for this invalu-
able cooperation.
For help and advice I am much indebted to K. Engell, who checked the
embryological and anatomical information, A. Fox Maule, who prepared the
latin diagnosis and G. Prance, who carefully read the manuscript and cor-
rected the language.
I am especially grateful to the artist, Mrs. Victoria Friis, who prepared the
original line drawings in Figs 10, 11, 15, 19, 20, 22, 24, 28 and 29, mostly from
dried or alcohol-preserved material, and in a few cases by copying selected
illustrations, the source of which is always cited. Illustrations from Hansen
1976a and 1977 are by Bent Johnsen.
SEM-investigations were carried out with a Cambridge instrument by J.
Fuglsang Nielsen at the Institute of Historical Geology and Palaeontology,
University of Copenhagen. LM-investigations by means of a Leitz Dialux-
72 FloraNeotropica

Orthomatfrom the DanishNaturalScienceResearchCouncil,who also made

fundsavailablefor the artist.
 Balanophoraceae 73

LITERATURE CITED
Adsersen, H. 1976. Ombrophytum peruvianurn (Balanophoraceae) found in the Galapagos Is-
lands. Bot. Not. 129: 113-117.
Asplund, E. 1928. Eine neue Balanophoraceen-gattung aus Bolivien. Svensk Bot. Tidskr. 22: 261-
277, fig. 1-3, t. 3.
Baker, E. G. 1890. New plants from the Andes. J. Bot. 28: 161-162, pl. 297.
Beccari, 0. 1869. Illustrazione di nuove specie di piante bornensi. Balanophoreae. Nuovo Giorn.
Bot. Ital. 1: 65-84, t. 2-4.
Blume, C. L. 1828. Flora Javae 1-2: 1-24, pl. 1-6. Bruxelles.
Browne, P. 1756. The civil and natural history of Jamaica 1-503, t. 1-49, London.
Dahlgren, R. 1975. A system of classification of the angiosperms to be used to demonstrate the
distribution of characters. Bot. Not. 128: 119-147,fig. 1-2.
Eichler, A. W. 1867. Sur la structure de la fleur femelle de quelques Balanophorees. Act. Congr.
Int. Bot. Paris 137-155, t. 1-2.
. 1869. Balanophoreae. In Martius, Fl. Bras. 4.2: 1-74, t. 1-16.
. 1873. Balanophoraceae. In DC., Prod. 17: 117-150.
Endlicher, S. L. 1836. Rhizantheae. In Gen. Plant. 72.
Engler, A. 1889. Balanophoraceae. In Engler & Prantl, Die natiirlichen Pflanzenfamilien ed. 1. 3.
1: 243-263,fig. 156-167.
Erdtman, G. 1952. Pollen morphology and plant taxonomy, angiosperms 1-539. Stockholm.
Fagerlind, F. 1938a. Bau und Entwicklung der floralen Organe von Helosis cayennensis. Svensk
Bot. Tidskr. 32(2): 139-159.
. 1938b. Ditepalanthus, eine neue Balanophoraceen-gattung aus Madagaskar. Ark.
Bot. 29A. 7: 1-15,fig. 1-5.
1945a. Bildung und Entwicklung des Embryosacks bei sexuellen und agamo
spermischen Balanophora-arten. Svensk Bot. Tidskr. 39: 65-82.
. 1945b. Blute und Blutenstand der Gattung Balanophora. Bot. Not. 1945(4): 330-
350.
. 1945c. Bau der floralen Organe bei der Gattung Langsdorffia. Svensk Bot. Tidskr.
39: 197-210.
. 1948. Bau und Entwicklung der vegetativen Organe von Balanophora. Kongl.
Svenska Vetenskapsakad. Handl. 25(3): 1-72.
Falcio, W. F. de A. 1975. Balanoforaceas. In Fl. I1. Cat. BALA 1-43.
Geesink, R. 1972. A new species of Langsdorffia from New Guinea (Balanophoraceae). Acta Bot.
Neerl. 21: 102-106,fig. 1.
Govindappa, D. A. & G. R. Shivamurthy. 1975. The pollination mechanism in Balanophora
abbreviata Blume. Ann. Bot. 39: 977-978, pi. 1.
_ 1976. "Seed" germination in Balanophora abbreviata. Phytomorphology 26: 135-
138,fig. 1-2.
Griffith, W. 1845. On the root-parasites referred to Rhizantheae. Trans. Linn. Soc. London 19:
303-347, t. 34-39.
Hansen, B. 1972. The genus Balanophora J. R. & G. Forster. A taxonomic monograph. Dansk
Bot. Ark. 28. 1: 1-188, fig. 1-44, t. 1-6, pl. 1-8.
. 1976a. Lathrophytum peckoltii Eichl. emend. B. Hansen (Balanophoraceae).
Bot. Tidsskr. 71: 75-79,fig. 1-2.
. 1976b. Pollen and stigma conditions in the Balanophoraceae s. lat. Bot. Not.
129: 341-345,fig. 1, 1. 1.
. 1977. Ombrophytum violaceum B. Hansen sp. nov. (Balanophoraceae). Bot.
Tidsskr. 71: 231-236,fig. 1-3.
. 1978. Changes in rank within Helosis and Corynaea (Balanophoraceae). Bot.
Tidsskr. 72: 188.
& K. Engell. 1978. Inflorescences in Balanophoroideae, Lophophytoideae and Scy-
balioideae (Balanophoraceae). Bot. Tidsskr. 72: 177-187, fig. 1-8.
Harms, H. 1935. Balanophoraceae. In Engler & Prantl, Die nattirliche Pflanzenfamilien ed. 2.
16b: 296-339, fig. 154-170.
Heinricher, E. 1907. Beitrage zur Kenntnis der Gattung Balanophora. Sitzungsber. Kaiserl. Akad.
Wiss., Math.-Naturwiss. Cl., Abt. 1. llb(l): 439-465.
74 Flora Neotropica

Hooker, J. D. 1856. On the structure and affinities of Balanophoreae. Trans Linn. Soc. London
22: 1-68,pl. 1-16.
. 1880. Balanophoraceae. In Benth. & Hook. f., Gen. Plant. 3: 232-239. London.
Howard, R. A. 1959. The Balanophoraceae in the Caribbean flora. Rhodora 61: 79-81.
Junghuhn, F. 1841. Uber Javan'sche Balanophoreen. Nova Acta Phys.-Med. Acad. Caes. Leop.-
Carol. Nat. Cur. 18(Suppl. 1): 202-228.
Klotzsch, J. F. 1847. Balanophoreae C. Richard, Bartling, Schott, Endlicher, R. Brown. Linnaea
20:460-461.
Knuth, P. 1904. Balanophoraceae. In Handbuch der Bliitenbiologie 3. 1: 260-265. Leipzig.
Kuijt, J. 1969. The biology of parasitic flowering plants 1-246. Berkeley & Los Angeles.
Lanjouw, J. 1932. Balanophoraceae. In A. Pulle, Fl. Suriname 1. 1: 45-46. Amsterdam
Lemee, A. 1955. Balanophoracees. In Fl. Guy. Franc. 1: 545. Paris.
Le6n, H. & H. Alain. 1951. Balanoforaceas. In Fl. Cuba 2: 82-84,fig. 31. La Habana.
Liebmann, F. M. 1847. Over nye rodparasiter henhorende til Balanophorernes, Cytineernes, Oro-
banchineernes og Monotropieernes familier. Forhandl. Skand. Naturf. 4. Moede 1844: 177-
187.
Lotsy, J. P. 1901. Rhopalocnemis phalloides Jungh. a morphological-systematical study. Ann.
Jard. Bot. Buitenzorg 17: 73-101, pi. 3-14.
Macbride, J. F. 1936. Balanophoraceae. In Fl. Peru 2: 427-431. Chicago.
Mangenot, G. 1947. Recherches sur l'organisation d'une Balanophoracee: Thonningia coccinea
Vahl. Rev. Gen. Bot. 54: 201-244, 271-294,fig. 1-39.
Martius, C. F. P. 1818. Uber eine neue Brasilianische Pflanzengattung in W. E. de Eschwege.
J. Bras. 2: 178-191, t. 5.
. 1832. Balanophoraceae. In Nov. Gen. Spec. Plant. 3: 181-188, pi. 298-300.
Munchen.
Moore, L. B. 1940. The structure and life-history of the root parasite Dactylanthus taylorii Hook.
f. New Zealand J. Sci. Technol. 21: 206B-224B.
Nevling, L. 1. Jr. 1960. Balanophoraceae. In Fl. Panama. Ann. Missouri Bot. Gard. 47: 303-308,
fig. 87-89.
Poeppig, E. F. 1833. Ombrophytum. In Leipz. Litteraturz. 1833: 1874.
& S. L. Endlicher. 1838. Ombrophytum peruvianum. In Nov. Gen. Spec. Plant.
2: 40-41, t. 155.
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31-33.
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Standley, P. C. 1930. Balanophoraceae. In Fl. Yucatan 250. Chicago.
1931. Balanophoraceae. In Fl. Lanc. Valley Honduras 178. Chicago.
1937. Balanophoraceae. In Fl. Costa Rica 409-410. Chicago.
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Steenis, C. G. G. J. van. 1932. Some remarks on the genus Rhopalocnemis Junghuhn. Handel. 6.
Nederl.-Ind. Natuurwet. Congr. 1931: 464-473.
Steyermark, J. A. 1957. Balanophoraceae. In Contr. Flora Venezuela. Fieldiana 28: 898.
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. 1797. Cynomorium. In Fl. Ind. Occ. 11-14. Erlangen.
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Rich. Dissertation Zurich 1-54. Freiburg.
Balanophoraceae 75

Weddell, H. A. 1850. Considerations sur l'organe reproducteur femelle des Balanophor6es et des
Rafflesiac6es. Ann. Sci. Nat. Bot. S6r. 3. 14: 166-187.
Zweifel, R. 1939. Cytologisch-embryologische Untersuchungen an Balanophora abbreviata Blume
und Balanophora indica Wall. Vierteljahrsschr. Naturf. Ges. Zurich 84: 245-306, fig. 1-16, t.
1-4.

NUMERICAL LIST OF TAXA

1. Scybalium 2. L. weddellii Hook.f.

1. S. depressum(Hook.f.) Eichl. 3. a. L. mirabile Schott & Endl. subsp
2. S. fungiformeSchott&Endl. mirabile
3. S. jamaicense(Sw.) Schott&Endl. 3. b. L. mirabile Schott & Endl. subsp
4. S. glazioviiEichl. bolivianum (Wedd.) B. Hansen

2. Helosis
1. a. H. cayennensis(Sw.) Spr.varcayen- 5. Lathrophytum
nensis 1. L. peckoltii Eichl.
1. b. H. cayennensis(Sw.) Spr.varmexi-
cana(Liebm.)B. Hansen 6. Ombrophytum
1. 0. violaceum B. Hansen
3. Corynaea 2. 0. microlepis B. Hansen
1. a. C. crassaHook.f. varcrassa 3. 0. peruvianum Poepp. & Endl.
1. b. C. crassaHook.f. var sprucei(Eichl.) 4. 0. subterraneum (Asplund) B. Hansen
B. Hansen

4. Lophophytum 7. Langsdorffia
1. L. leandriEichl. 1. L. hypogaea Mart.
76
LIST OF EXSICCATAE
Acosta Solis, M., 5039(7-1); 5070 (3-1);
5837 (1-1)
Adsersen, H. et al., 243 (6-4); 244 (6-4);
1323 (6-4)
Alexander, R. C., sn (1-3)
Alleizette, C. d', 6363 (2-lb)
Allen, P. H., 4773 (7-1)
Alston, A. H. G., 7819 (2-la)
Anderson, W. R. et al, 3117 (1-3); 7412 (1-2)
8419 (7-1); 9754 (7-1)
Araujo, D. et al, 204 (2-la); 523 (2-la)
Argent, G. C. G., 6706 (7-1)
Asplund, E., 2568 (6-4); 3873 (6-4);
6312 (6-4); 10010 (3-la); 14226 (2-la)
Austin, et al, 4137 (2-la)
Balansa, B., 2717 (4-3b)
Baldwin, J. T., 3302 (2-la)
Bamps, P., 5198 (7-1)
Barclay, H. G. et al, 10325 (3-1)
Barreto, NI., 7431 (4-3a); 8799 (7-1)
Barriga, H. G., 8395 (2-la); 14700 (2-la)
Baumbart, 1., sn (7-1)
Beaman, 6185 (2-lb)
Benjamin, D. S., 9284 (7-1)
Berlin, B., 404 (2-la); 457 (2-la)
Bernardi, A. L., 969 (7-1); 3347 (2-la)
Bertero, C. G. L, sn (1-3)
Black, G. A. et al, 957 (2-la); 1410 (2-la);
2039 (2-la); 2388 (2-la); 2961 (2-1a);
3416 (2-la); 8423 (2-la); 8584 (2-la)
Bourgeau, E., 3030 (2-lb)
Brade, A. C., 18909 (1-4)
Breedlove, D. E., 22479 (2-la)
Brenes, A. M., 12638 (2-1a)
British Guiana For. Dept., 6170 (2-la);
6353 (2-1a)
Britton, N. L. et al, 2780 (1-3)
Broadway, W. E., 799 (2-la); 6434 (2-la);
6568 (2-la)
Buchtien, O., 1641 (7-1)
Burger, W. C. et al, 6060 (3-1b); 6066 (7-1)
Cabezas, V., SI 20263 (6-4); SI 20341 (6-4)
Camp, W. H., 4295 (3-lb); 4418 (7-1)
Castafeda, R. R., 4054 (2-la)
Cavalcante, P. B., 2261 (2-la); 3205 (2-la)
Chagas, J., INPA 3122 (2-la); INPA
3553 (2-la); INPA 6309 (2-la)
Claussen, P., 1410 (7-1); 1411 (7-1)
Clement, A. C. T., sn (1-3)
Coppois, G., sn (6-4)
Core, E. L., 330a (7-1); 331a (3-la);
332a (3-1)
Cornelio, sn (7-1)
Cowan, R. S. et al, 2005 (2-la); 39116 (2-la)
Flora Neotropica
Croat, T. B., 18824 (2-la)
Croat, T. B. et al, 16158 (2-lb)
Crueger, H., sn (2-la)
Cuatrecasas, J., 5429 (7-1); 6263 (7-1);
11851 (3-1); 19121 (3-1)
Curran, H. M., sn (2-la); 22 (4-3a);
334 (2-la); 414 (2-la)
Daniels, A. G. H. et al, 733 (2-la);
1134 (2-la); 1277 (2-la)
Dressier, R. L. et al, 38 (2-lb)
Drew, W. B., E-51 (7-1)
Duarte, A. P., 9568 (7-1); 10281 (7-1)
Ducke, A., 1674 (7-1); 3462 (2-la);
3561 (2-la); 3632 (2-la); 6875 (2-la);
7595 (2-la)
Duke, J. A., 5320 (7-1); 13621 (2-la)
Dungs, F., sn (1-4)
Duss, A., 2106 (2-lb)
Dwyer, J. D., 11696 (2-la)
Eggers, H. von, 1620 (1-3)
Egler, W. A. et al, 46697 (2-la)
Eiten, G. et al., 3094 (1-2); 3173 (1-2)
Ekman, E. L., 2653 (1-3); 3348 (1-3);
3877(1-3); 5113 (1-3); 9037 (1-3);
10581 (1-3); 14285 (1-3)
Eliasson, U., 13 (2-lb)
Emmerich, M., sn (4-3a)
Ernani, sn (2-1a)
Ewan, J. A., 16075 (1-1); 16544 (3-lb)
Fendler, A., sn (2-la); 1743 (7-1); 1744 (7-1)
Fereira, 221 (2-la)
Ferreyra, R., 3401 (2-la)
Florschutz, P. A. et al, 2545 (2-la);
2604 (2-la); 2863 (2-la); 2942 (2-la)
Forero, E., 1049 (3-1); 1051 (3-la)
Fosberg, F. R., 20923 (3-1); 21151 (3-1);
21518 (1-1); 21520 (2-la)
Gehrt, A., sn (1-2)
Gentry, A. et al, 14018 (2-1a); 14019 (7-1)
Giovanni, F., 36 (2-la)
Glaziou, A. F. M., sn (4-3a); sn (5-1);
N. 306 (4-3a); 4070a (1-2); 4070b (1-4);
4700 (1-4); 4969a (2-la)
Golbach, R., sn (4-3b)
Gollmer, J., sn (3-lb)
G6mez, L. D., 4034 (7-1); 786722 (3-lb)
Goudot, J., sn (3-lb); 140 (2-lb)
Grant, M. L. et al, 9311 (3-1); 9701 (3-1);
10029 (3-lb); 10030 (7-1)
Granville, J. J. de, C 72 (2-la); 2164 (2-la)
Grondona, E. et al, 1709(4-1)
Guedes, T., 2298 (2-la)
Balanophoraceae
Gwynne-Vaughan, D. T., 55 (2-la)
Hahn, L, 400 (2-1b)
Hall, F., sn phot. (6-3); sn phot. (6-2);
636 (2-la); 1100 (2-la)
Harling, G. et al., 7026 (6-3); 13209 (3-lb)
Hashimoto, G., 150 (7-1)
Hatschbach, G., 9842 (4-1); 11287 (2-1a);
19606 (4-1); 20771 (7-1); 20981 (2-1a)
Haught, 0., 2010 (2-1a); 2881 (2-1a)
Hawkes, J. G. et al, 2152 (2-lb)
Hedger, 74 (2-1a)
Herb. Trinidad, 15416 (2-1a)
Heringer, E. P., 8859 (7-1)
Heringer, E. P. et al, 6974 (7-1)
Hirschel, O., sn (3-la); sn (3-la)
Hoehne, F. C., 3874 (7-1); 3875 (7-1);
3876 (7-1); 27517 (7-1)
Holm-Nielsen, L. et al., 1262 (1-1);
4155 (2-la); 6910 (3-lb)
Howard, R. A., 13400 (1-3)
Hubbard, T., sn (4-1)
Huertas, G. et al, 6336 (3-la)
Hunt, D. R. et al, 6027 (7-1)
Hutchinson, P. C. et al, 5551 (7-1)
Irwin, H. S. et al, 16560 (7-1); 16848 (7-1)
Jaramillo, R., 2451 (2-la); 2660 (7-1)
Jenman, G. S., 1852 (2-1a)
Jonker-Verhoef, A. M. E. et al, 443 (2-la)
Karsten, G. K. W. H., sn (7-1)
Kegel, H. A. H., 607 (2-la)
Kennedy, H. et al, 3101 (2-1a)
Killip, E. P. et al, 8354 (2-la); 27622 (2-la)
Kramer, K. U., 1780 (1-3)
Kramer, K. U. et al, 2478 (2-la)
Krapovickas, A., 1687 (4-3b)
Krapovickas, A. et al, 15509 (4-1);
28596 (4-1)
Kuhlmann, J. G., 4613 (2-1a)
Kuntze, C. E. 0., 1926 (2-1a)
Kupper, W., 1549 (2-1a)
Langsdorff, G. H. von, sn (7-1)
Lawrance, A. E., 251 (7-1)
Lehmann, F. C., K 401 (2-la)
Leite, J. E., 720 (2-la)
Lemos, A., sn (1-2)
Lent, R. W., 788 (2-lb); 1282 (2-lb);
1658 (2-lb)
Leonard, E. C., 5763 (1-3)
Liebmann, F. M., sn (2-lb); sn (2-lb);
sn (7-1)
Lima, F. I., 238 (1-2); 1437 (2-la);
1526 (2-la)
Lima, F. I. et al, 1914 (1-4)
77
Linden, J. J., 146 (2-lb)
Liogier, A., 14463 (1-3)
Little, E. L., 7650 (1-1); 7663 (3-1);
7684 (7-1); 8214 (7-1); 8991 (7-1)
Lloyd, C. G., sn (1-3)
Lourteig, A., 690 (4-3b); 2320 (2-la)
Luetzelburg, P. von, sn (7-1); sn (2-1a);
20143 (2-la); 20173 (2-la); 20174 (2-la);
20212 (2-la); 21085 (2-la); 21245 (2-la);
21259 (2-la); 21679 (2-1a); 21866 (2-la);
21891 (2-la)
Lund, P. W., sn (7-1)
Lundell, C. L., 6414 (2-lb)
Lutz, B, et al, sn (2-la)
Lofgren, A., 2513 (7-1)
Maas, P. J. M., 1178 (2-la); 1334 (2-la);
2180 (2-la); 11068 (2-la)
Macedo, A., 2337 (7-1)
Madison, M. T., 2356 (3-lb)
Madison, M. T. et al, 5450 (6-3)
Maguire, B. et al, 23524 (2-la); 28054 (7-1);
28057 (7-1); 31606 (2-la); 31607 (2-la);
33832 (7-1); 33841 (7-1); 33927 (7-1);
40733 (2-la); 40794 (2-la); 47006 (2-1a);
47091 (2-la)
Malme, G. O. A., sn (2-la); sn (7-1)
Marlyn, 269 (2-la)
Martin, J., 486 (6-4)
Martius, K. F. P. von, sn (7-1); 2643 (2-la)
Marunak, V., 633 (4-1)
Mathias, M., 5673 (2-la)
Matuda, E., 423 (2-lb); 3558 (2-la);
4167 (2-lb)
Maxon, W. R., 9137 (1-3)
Maxon, W. R. et al, 559 (1-3)
Mello, J. C. de, sn (1-2)
Mexia, Y., 6337a (6-1)
Miers, J., sn (7-1); sn (2-la); sn (2-la)
Miranda, F., 1483 (2-la); 7530 (2-la)
Moore, S., et al, 9709 (7-1)
Moritz, J. W. K., 1619 (7-1)
Morley, B. D. et al, 638 (1-3)
Morris, D., sn (1-3)
Morton, C. V. et al, 3615 (1-3)
Mosen, C. W. H., 4404 (7-1)
Mueller, J., 340 (4-1)
Moller, F. A. G. J., sn (4-1); sn (2-1a)
Netto, L. do S. M., sn (4-3a)
Occhioni, 981 (2-la); 982 (4-3a); 983 (4-3a)
Oldeman, R. A. A., 1141 (2-1a); 1199 (2-la)
Oliveira, E., 145 (7-1); 196 (7-1); 278 (7-1);
446 (7-1)
Orbigny, A., 1188 (7-1)
Orcutt, 2844 (1-3); 5706 (1-3)
78
Pabst, G. F. J., 5416 (4-3a); 7301 (7-1)
Parker, K. et al, 37b (6-4)
Peckolt, Th., sn (4-3a); sn (5-1); sn (1-4);
sn (5-1)
Pelaes, sn (2-lb)
Pereira, E., 5645 (5-1)
Pfister, A., sn (6-4)
Pflanz, K., 8 (4-3b)
Philipson, W. R., 1253 (1-3); 1254 (1-3)
Pickel, B., 3423 (1-2)
Pinto, P. et al, 918 (2-la)
Pires, F. D. de A., sn (2-lb)
Pires, J. M. et al, 1239 (2-1a); 4511 (2-1a);
14521 (2-la)
Pittier, H., 1531 (1-1); 9900 (2-1a)
Planchon, J. E., sn (4-3a)
Plowman, T., 4688 (3-lb); 5940 (4-2)
Plowman, T. et al, 2345 (2-la); 4934 (2-la)
Poeppig, E., sn ill. (6-3)
Prance, G. T. et al, 2764 (2-la); 3589 (2-1a:
4457 (2-la); 5804 (2-la); 7664 (6-3);
7833 (2-1a); 8080 (2-1a); 8513 (2-1a);
8765 (2-la); 9370 (2-la); 10200 (2-1a);
10499 (2-1a); 10819 (2-1a); 13565 (2-la:
14391 (2-la); 15787 (2-la); 16309 (2-la;
24212 (2-la); 24410 (2-la); 25483 (2-la;
55899 (2-la)
Pringle, C. G. , 5050 (2-lb)
Proctor, G. R., 9341 (1-3); 17764 (2-1b);
21465 (1-3)
Purdie, W., sn (1-3); sn (1-1); sn (7-1);
sn (7-1); sn (2-lb); sn (3-la); sn (3-la);
sn (2-la)
Purpus, C. A., 2750 (2-lb); 5896 (2-lb);
7691 (2-lb)
Ramage, G. A., sn (2-lb)
Regnell, A. F., sn (1-2); sn (7-1)
Reinhardt, J. Th., sn (7-1)
Reitz, R. et al, 7156 (7-1); 7328 (2-lb);
17353 (2-lb)
Richard, L. C., sn (2-la)
Ridley, H. N., sn (1-3)
Riedel, L., sn (2-1a)
Rizzo, 4706 (7-1)
Robyns, W. et al, sn (2-la)
Rodrigues, W. A., 8347 (2-la)
Rojas, 3836 (7-1)
Ruiz, H., sn (2-la)
Rusby, H. H., 256 (3-lb)
Rusby, H. H. et al, 105 (2-la)
Sander & Co., sn (4-2)
Sandwith, N. Y., 620 (2-la)
Sarakhan, J., 1985 (2-la)
Castre, C., 1518 ( --a); 1635 (2-la)
Schiffner, et al, 698 (4-1)
Schipp, W. A., 719 (2-lb); 1129 (2-la)
Flora Neotropica
Schneider, M., 612 (3-1a)
Schnell, 12239 (2-1a)
Schomburgk, M. R., sn (7-1)
Schott, H. W., sn (7-1); sn (1-2)
Schrink, sn (1-3)
Schultes, R. E., 6197 (2-la); 6323 (2-la);
6657 (2-1a); 6829 (2-la); 8117 (2-la);
8615 (6-2); 12091a (2-la)
Schulz, J. P. 10287 (2-la)
Schunke, J., 3943 (6-2); 5675 (4-2); 9617 (7-1)
Schwabe, W., sn (4-1)
Schwarzmayer, S., sn (7-1)
Seibert, R. J., 469 (2-la)
Shafer, J. A., 8858 (1-3)
Shepard, R. S., 246 (6-4)
Sick, H., B 783 (2-la)
Silva, M., 1241 (2-la); 2661 (7-1)
Silva, M. F., 689 (2-la)
Simpson, D. R. et al, 660 (2-la)
Sintenis, P., 4238 (1-3)
Skutch, A. F., 2669 (2-lb)
Sleumer, H., 3054 (6-4); 3331 (6-4)
Smith, L. B., 1777 (1-4)
Smith, L. B. et al, 7287 (4-1)
Sneidern, K. von, 1263 (7-1); 1264 (7-1)
Snethlage, E. H., 231 (2-la)
Sparre, B., 13049 (6-1); 14338 (3-lb);
16947 (3-lb); 19070 (2-la)
Splitgerber, F. L., 728 (2-1a)
Spruce, R., 151 (2-la); 574 (2-la); 877 (2-la);
1129 (2-la); 5186 (3-lb)
Stahel, G., 803 (2-la); 4659 (2-la)
Standley, P. C., 54815 (2-la)
Stern, W. L. et al, 737 (2-la)
Steyermark, J. A., 39584 (2-1a); 46131 (2-1a);
46801 (2-lb); 51848 (2-1b); 52634 (1-1);
53515 (7-1); 55247 (3-1); 75657 (7-1);
86840 (2-1a); 88983 (2-1a); 89372 (2-1a);
89838 (2-la); 89852 (7-1)
St.-Hilaire, A., sn (7-1)
Sucre, D., 3052 (7-1); 10218 (4-3a)
Swartz, O. P., sn (1-3)
Taylor, N., 219 (1-3)
Teunissen, P. A., 12951 (2-1a)
Triana, J. J., sn (3-lb)
Troll, K., 99 (6-4)
Tuerckheim, 700 (1-3); 1044 (2-lb);
2657 (1-3); 8576 (2-la)
Ule, E., sn (6-1); sn (6-2); sn (4-3b);
948 (2-lb); 6069 (2-la); 9025 (7-1)
Venturi, 6612 (4-3b)
Viirsoo, sn (6-4); sn (6-4)
Walker, J. W., 449 (2-lb)
Ward, 8629 (2-la)
Balanophoraceae
Warming, E., sn (7-1)
Webster, G. L. et al, 4981 (1-3); 8352 (1-3)
Weddell, H. A., 3631 (4-3b); sn (6-3);
4786 (3-lb)
Welden, A. L., sn (2-lb)
Wendland, J. C., sn (3-la)
Werff, H. van der, 597 (6-4)
Wessels Boer, J. G., 734 (2-la)
Whymper, E., sn (3-lb)
Wilbur, R. L. et al, 10309 (2-lb);
15454 (2-1a)
79
Williams, L. O. et al, 28189 (3-lb)
Wolfe, F., sn (7-1)
Woodson, R. E. et al, 470 (3-lb)
Wright, C., 546 (1-3); 2636 (2-lb)
Wullschlagel, sn (2-1a); 1300 (1-3)
Wurdack, J. J., 952 (3-lb); 2112 (2-la);
2360 (6-1); 34062 (7-1); 34063 (7-1);
34073 (7-1)
Wurdack, J. J. et al, 39757 (2-1a)
Yuncker, T. G., 17553 (1-3); 18466 (1-3)
80 Flora Neotropica

INDEX OF SCIENTIFIC NAMES

Archimedea45 subterranea 62
pyramidalis46, 51 Langsdorsfia 67
Balanophoroideae (subfam.)67 hypogaea 68
Caldasia33 janeirensis 68
brasiliensis37 moritziana 68
cayennensis37 rubiginosa 68
mexicana40 Lathrophytium 54
Corynaea41 Lathrophytum 54
crassa42 peckoltii 54
varcrassa44 Latraeophila 33
varsprucei44 Lepidophytum 45
purdiei42 Lophophytoideae (subfam.) 45
sphaerica42 Lophophytum 45
sprucei44 bolivianum 52
Cynomorium leandri 46
cayennense35 mirabile 49
coccineumauct. 28 subsp bolivianum 52
jamaicense28 subsp mirabile 52
Ditepalanthus33 mirabile auct. 46
Exorhopala33 weddellii 49
Helosideae(tribus)32 Ombrophytum 55
Helosidoideae(subfam.)24 microlepis 60
Helosieae(tribus)31 peruvianum 60
Helosis33 peruvianum auct. 58, 60, 64
Helosisauct.41 subterraneum 62
brasiliensis36 ulei 58
cayennensis35 violaceum 58
varcayennensis38 zamioides 62
varmexicana40 Phyllocoryne 24
guyanensis36 jamaicensis 28
formaandicola40 Rhopalocnemis 33
varandicola40 Scybalieae (tribus) 24
formabrasiliensis37 Scybalioideae (subfam.) 24, 45
jamaicensis28 Scybalium 24
mexicana40 depressum 25
varandicola40 fungiforme 27
mexicanaauct. 37 glaziovii 29
whymperi44 jamaicense 28
Itoasia41 Senftenbergia 67
crassa42 Sphaerorhizon 24
purdiei42 curvatum 25
sphaerica42 depressum 25
sprucei44 Thonningia
Juelia56 Thonningia auct. 67
lilloana64 janeirensis 68
meyeri64 mexicana 68