Revision Geonoma Phytotaxa 2011

Phytotaxa 17: 1271
(2011)
www.mapress.com/ phytotaxa /
Copyright 2011 Magnolia Press
Monograph
ISSN 1179-3155 (print edition)
PHYTOTAXA
ISSN 1179-3163 (online edition)
PHYTOTAXA
17
A revision of Geonoma (Arecaceae)

ANDREW HENDERSON
The New York Botanical Garden, Bronx, NY 104585126, U.S.A.
Email: ahenderson@nybg.org
Magnolia Press
Auckland, New Zealand
Accepted by W. Baker: 2 Nov. 2010; published: 18 Feb. 2011
ANDREW HENDERSON
A revision of Geonoma (Arecaceae)
(Phytotaxa 17)
271 pp.; 30 cm.
18 Feb. 2011
ISBN 978-1-86977-647-3 (paperback)
ISBN 978-1-86977-648-0 (Online edition)
FIRST PUBLISHED IN 2011 BY

Magnolia Press
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2011 Magnolia Press

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ISSN 1179-3155
(Print edition)
ISSN 1179-3163
(Online edition)
Phytotaxa 17 2011 Magnolia Press
HENDERSON
Table of contents
Abstract ................................................................................................................................................................................ 3
Introduction .......................................................................................................................................................................... 3
Materials and Methods ......................................................................................................................................................... 4
Phylogeny............................................................................................................................................................................. 9
Distribution ........................................................................................................................................................................ 14
Infraspecific Variation ........................................................................................................................................................ 15
Morphology........................................................................................................................................................................ 23
Taxonomic Treatment......................................................................................................................................................... 29
Acknowledgements .......................................................................................................................................................... 162
References ........................................................................................................................................................................ 163
Appendix I. Qualitative VariablesCharacters and Traits ................................................................................................ 168
Appendix II. Quantitative Variables................................................................................................................................. 171
Appendix III. Excluded Names........................................................................................................................................ 172
Appendix IV. Plates of Type Images ................................................................................................................................ 177
Appendix V. Numerical List of Taxa and Specimens Examined ..................................................................................... 249
Abstract
A taxonomic revision and phylogeny of the neotropical palm genus Geonoma based on morphological data and
morphometric methods was carried out. 4990 herbarium specimens were scored for 44 qualitative variables and 27
quantitative variables. Qualitative variables were divided into 30 characters and 14 traits. Using the phylogenetic species
concept, characters were used to recognize 68 species. These are widely distributed from southern Mexico to Bolivia and
Paraguay, and reach the Lesser Antilles and Hispaniola. Analysis of each species for traits, geographic distribution, and
quantitative variables led to recognition of 90 subspecies in 18 species, giving a total of 140 taxa. Twelve new species (G.
bernalii, G. concinnoidea, G. deneversii, G. dindoensis, G. fosteri, G. galeanoae, G. gentryi, G. operculata, G. peruviana,
G. sanmartinensis, G. schizocarpa, G. venosa) and 33 new subspecies (G. brongniartii subsp. pascoensis, G. concinna
subsp. simplex, G. concinnoidea subsp. coclensis, G. concinnoidea subsp. jefensis, G. congesta subsp. osensis, G. cuneata
subsp. guanacastensis, G. cuneata subsp. indivisa, G. cuneata subsp. minor, G. cuneata subsp. rubra, G. deversa subsp.
belizenesis, G. deversa subsp. peninsularis, G. deversa subsp. quadriflora, G. ferruginea subsp. nicaraguensis, G.
lehmannii subsp. corrugata, G. longivaginata subsp. copensis, G. longivaginata subsp. sanblasensis, G. longivaginata
subsp. vallensis, G. maxima subsp. dispersa, G. maxima subsp. multiramosa, G. maxima subsp. sigmoidea, G. pohliana
subsp. linharenseis, G. pohliana subsp. rodriguesii, G. pohliana subsp. unaensis, G. stricta subsp. antioquiensis, G. stricta
subsp. bracteata, G. stricta subsp. divaricata, G. stricta subsp. pendula, G. stricta subsp. pliniana, G. stricta subsp.
quibdoensis, G. stricta subsp. submontana, G. undata subsp. tacarcunensis, G. undata subsp. tumucensis, G. undata
subsp. venezuelana) are described. Forty-one new combinations are made. Several of the most variable species are
considered to be species complexes and are divided into morphotypesgroups of similar specimens with no formal
taxonomic status. Nomenclature, descriptions and distribution maps are provided for each species and subspecies.
Images of the type specimens of all new taxa are also provided. A phylogenetic analysis, using the same 30 characters
used for the taxonomic revision was carried out using parsimony analysis. A sample tree and consensus tree are shown
and a discussion is given of the various clades.
Key words: Palmae, palms, phylogeny, morphometrics, Neotropics
Introduction
Geonoma Willdenow (1805: 174) is one of the largest three palm genera in the Neotropics, approximately
equal in size to Bactris Jacquin ex Scopoli (1777: 70) and Chamaedorea Willdenow (1806: 638). Species of
Geonoma are common in lowland and montane tropical moist forests in Central and South America, and reach
Hispaniola and the Lesser Antilles. They occur from sea level to just over 3000 m elevation. Geonoma are
usually rather small, understory palms, although some reach the canopy of montane forests.
REVISION OF GEONOMA (ARECACEAE)
The genus was described over 200 years ago by Willdenow (1805), based on just two species from
Venezuela. Five more species from French Guiana were added by Poiteau (1822), although these were placed
in a different genus, Gynestum Poiteau (1822: 387), based on a misinterpretation of the flowers. These species
were transferred to Geonoma by Kunth (1841). Martius (18231837) greatly increased the number of species
with the description of his Amazon collections, and of dOrbignys Bolivian collections (Martius, 1843).
The first revision of the whole genus was that of Burret (1930a), a German botanist who worked in the
Berlin herbarium. In this revision, published as Geonomeae americanae, 172 species were recognized.
Burrets work was based in part on an unpublished manuscript by the German horticulturalist Hermann
Wendland (18251903). Burret went on after 1930 to describe a further 49 new species of Geonoma so that by
the time the last was published, in 1940, he recognized a total of 221 species. Disaster struck in 1943 when the
Berlin herbarium was bombed and most of Burrets type specimens, as well as Wendlands manuscript were
destroyed.
Burret has been criticized for employing an extremely narrow species concept. Wessels Boer (1968) wrote
As a result geonomoid palms are grossly overnamed in the present authors opinion; species have been
distinguished on, as will be shown, irrelevant characters and identification is next to impossible.Wessels
Boers (1968) treatment of Geonoma was based on extensive field experience in Suriname and Venezuela. In
his treatment, few of Burrets (1930a) names survived synonymy and only 75 species were recognized.
However, Wessels Boers revision was criticized by Moore (1969), particularly for inaccuracies in the key and
descriptions.
Since Wessels Boers revision in 1968, 16 new species have been described. There have been several
regional treatments, e.g., Henderson (1995) for the Amazon region, Borchsenius et al. (1998) for Ecuador,
Hammel et al. (2003) for Costa Rica, and Galeano & Bernal (2010) for Colombia. Henderson et al. (1995)
provided a treatment of the whole genus in field guide format, where Geonoma was considered to be difficult
taxonomically because of the occurrence of several widespread and variable species complexes. Because of
this, a pragmatic approach was taken and species complexes were recognized as single species.
Geonoma thus presents a classic problem in taxonomyhow to reconcile the splitting of Burret who
recognized 221 species, with the lumping of Wessels Boer who recognized 75 species. In the present study,
explicit, repeatable, quantitative methods, leading to testable hypotheses are employed, in an attempt to
understand variation within the genus.
Materials and Methods

Species concept
In this study the Phylogenetic Species Concept (PSC) is used. Under this concept, species are defined as: "the
smallest aggregation of populations.... diagnosable by a unique combination of character states in comparable
individuals" (Nixon & Wheeler 1990). Individual specimens are considered comparable because all are
fertile. The terms character and trait are used in the sense of the PSC. Characters are qualitative variables the
same states of which are found in all comparable individuals within a terminal lineage (i.e., species); traits are
qualitative variables with more than one state found within species (although some species may have only one
state of a given trait). The PSC is chosen here because it has an explicit definition, theoretical background and
discovery operation, as described below. This is discussed in more detail in Henderson (2005a; see also
Henderson 2004 and 2005b).
Two operational modifications are necessary in order to apply the PSC. According to Davis & Nixon
(1992), phylogenetic species are delimited by successive rounds of aggregation of local populations, based on
analysis of characters and traits. Because palm specimens are seldom collected on a population basis and
because there is no a priori method of placing specimens in populations and consequently distinguishing a
priori between characters and traits, all specimens (i.e., treating specimens as populations) and all qualitative
variables (i.e., traits and characters) were used in the analysis (see below).
HENDERSON
A second modification of the PSC involves subspecific variation. Some groups of specimens with unique
combinations of qualitative character states (i.e., species) may vary internally in trait state distributions and
quantitative variables, and may occur in disjunct geographic areas. Based on these criteria subgroups may be
recognizable. Luckow (1995), in her discussion of the PSC, stated that groups of populations that differ not
by fixed characters, but by differences in mean values, would be recognized as subspecies or varieties [under
the PSC]. A slightly modified version of this is followed here. If subgroups can be delimited by traits (i.e.,
with unique combinations of trait states) and/or by geographic disjunctions, and these subgroups are
supported by analysis of quantitative variables (see below), then a phylogenetic subspecies concept is applied.
In summary, the PSC is applied to groups of specimens with unique combinations of qualitative character
states, and a PSC subspecies concept is applied to subgroups that can be delimited by analysis of traits,
geography and quantitative variables.
Species delimited under the PSC are testable hypotheses (Wheeler & Platnick 2000). This testing depends
on distinguishing characters from traits, i.e., the test is that characters are not traits and traits are not
characters. In the first case, a supposed character may turn out to be distributed as a trait. Such a
misinterpretation would give an overestimation of the number of species. In the latter case, a supposed trait
may be distributed as a character, giving an underestimation of the number of species.
Data matrix construction
Four thousand, nine hundred and ninety (4990) specimens from the following herbaria were examined and
scored: AAU, BH, BM, BR, C, COAH, COL, CR, F, FTG, G, GH, HUA, INB, INPA, K, LE, M, MBML,
MEXU, MO, NY, P, PMA, RB, SP, SPF, U, US, and USM (herbarium abbreviations from Holmgren et
al.1990). Fragmentary type specimens from some herbaria (e.g., from C, M) were examined but not
necessarily scored. Sometimes, more than one duplicate of a collection was used in scoring.
Morphological attributes that could be scored or measured from specimens were divided into qualitative
(binary or multistate) or quantitative (continuous, meristic) variables. A search was made for qualitative
variables in which two or more states of the variable were present among the specimens and could be scored
unequivocally. This search was based on those variables used in previous monographs (e.g., Wessels Boer,
1968) and on a survey of specimens. A dissecting microscope was used to survey floral variables. Forty-four
qualitative variables were found and scored (Appendix I). One attribute (rachillae hairs) has been used
previously in Geonoma (e.g., Wessels Boer 1968) but was not used here because potential states could not be
scored unequivocally, and hairs are early deciduous and often cannot be seen on rachillae from inflorescences
past anthesis.
A search was made for quantitative variables that could be measured from specimens or taken from
specimen labels (where, in case of ranges, median values were used). Variables were counted or measured
with a ruler, digital calipers, or protractor. Twenty-seven quantitative variables were found and scored
(Appendix II). Five are from stems, 11 from leaves and 11 from reproductive structures. Twenty-four are
continuous and three are meristic.
A data matrix was constructed with specimens as rows and variables as columns (http://sciweb.nybg.org/
Science2/res/Henderson/Geonoma.xls.zip). Additional columns recorded a specimen identification number,
collector, collectors number, herbarium, country, latitude, longitude, and elevation. Specimen identification
numbers are unique within species, but not between species. Latitude and longitude were taken from the
specimen label, when available. On specimens lacking coordinates, these were estimated from the collection
locality using either maps or electronic gazetteers.
For each of the 4990 specimens in the matrix, three spatial variables and 70 morphological variables were
recorded, giving a potential total of 364,270 data points. Nevertheless, approximately 40% of these potential
data are missing in the matrix, because specimens are often fragmentary or incomplete and various organs are
often not preserved (e.g., bracts, flowers, fruits). Data on plant and stem height and branching are often
missing from labels.
Data analyses
Some inferential statistics were used in this study. Although random samples are required for statistical
inference, the samples of herbarium specimens are not random. However, there is no reason to believe that
collectors favored any particular kind of specimen over others. Therefore inferential statistics were used, but
the results should be considered accordingly. Statistical analyses were carried out using the programs NTSYS
(Rohlf 2000) and Systat (Wilkinson 1997). Specimens with missing values were excluded. Analyses are thus
based on subsets of the data. Because some quantitative variables were not normally distributed, data were
log10-transformed before analysis.
Species delimitation
All specimens were assigned a preliminary species identification, either based on a previous determination or
on the key in Wessels Boer (1968). Cluster analysis (CA) was used to divide qualitative variables into either
characters or traits. The SIMQUAL module of NTSYS with the simple matching coefficient (for binary and
multistate variables) was used to produce a similarity matrix. The SAHN module of NTSYS was used to
subject the similarity matrix to the unweighted pair group method, arithmetic average (UPGMA) clustering
algorithm. Successive analyses were used, with all variables used in the first analysis. Suspected traits (i.e.,
those variables both states of which occur in adjacent and otherwise homogeneous groupings) were removed,
and the analysis run again until groups were found with unique combinations of states. These groups were
recognized as species.
One variable with many missing values (stem branching) was excluded from this analysis and was treated
as a trait. Because of missing data in other variables (e.g., few specimens contained both flowers and fruits
and consequently had missing data for one of other of these variables), analyses were made separately on leaf,
inflorescence, and flower variables, and then leaf, inflorescence, and fruit variables.
Specimens that had not been included in these analyses because of missing data were then assigned to
their respective species based on their morphology and geography.
Subspecies delimitation
Variation within each species was examined, based on analysis of trait state distributions, geographic
distributions, and quantitative variables. The purpose of these analyses was to look for evidence of presence
of discrete subgroups (i.e., subspecies).
For each species, all traits were examined and those that varied within a species were used to recognize
subgroups with unique combinations of trait states (trait subgroups). In general, little value was placed on two
traitsstem branching and leaf divisionbecause these seldom appeared to have any taxonomic value.
Next, geographic distribution of species was analyzed by examining distribution maps produced by
Arcview GIS 3.2 (Environmental Systems Research Institute, Inc.) using latitude and longitude data for each
specimen. Each dot on the maps represents at least one specimen. Geographic subgroups were recognized if
specimens clustered in discrete groups separated from other such groups.
Finally, quantitative variation was analyzed. A t-test(two-sample, separate variance test on log10transformed variables) or, with more than two subgroups, an one-way ANOVA (on log 10 -transformed
variables) was used to test for trait and geographic subgroup differences for each quantitative variable. The
Bonferroni pair wise procedure was used to see which pairs of means differed significantly (P <0.05). If there
were too few specimens, usually less than 10, then tests were not carried out. If subgroups delimited by traits
and/or by geographic disjunctions were supported by analysis of quantitative variables, then they were
recognized as subspecies.
Morphotypes
Subspecies delimitation in several species is problematic. About 13 species, or 20% of the total, are
considered to be species complexes or polymorphic species (see section on Infraspecific Variation). These are
widespread, variable species in which numerous local variants occur.
HENDERSON
Some groups of specimens within species complexes can be recognized as subspecies. This is based, as
discussed above, on unique trait combinations and/or geographic separation, and this recognition is usually
supported by quantitative data. However, this leaves a residual group of specimens which are often
widespread and differ slightly from site to site. These patterns of variation make taxonomic treatment of
species complexes problematic. Recognition of some subspecies within a species complex means that by
default residual specimens must also be treated as subspecies, although the patterns of variation suggest that
these would be artificial taxa. Alternatively, no subspecies may be recognized within species complexes, but
rather they may be divided into morphotypes: informal groups of similar specimens with no formal taxonomic
status. Of the two options for treating species complexesdivision into subspecies some of which may be
artificial, or division into morphotypes some of which may be subspeciesneither is satisfactory. Depending
on the evidence, some species complexes are divided into subspecies, and some of these may be divided into
morphotypes, and other species complexes are not divided subspecifically but are divided into morphotypes.
If type specimen(s) are present in morphotype groups, then the name of the earliest type is given to the
morphotype.
Environmental variation
Linear regression was used to analyze relationships within species and subspecies between log10-transformed
quantitative variables and latitude, longitude, and elevation. The first two of these were taken as proxies for
correlated variation in environmental variables. If there was a significant (P <0.05) correlation between
variables, squared multiple R is reported. This shows the amount of variance in the dependent variable
explained by the independent variable.
Taxonomic treatment
A genus description of Geonoma is given (based on characters and traits) (see also section on Morphology for
an illustrated discussion of morphology). This is followed by a key to all species, based on all attributes
(characters, traits, quantitative variables) and geography.
For each species, arranged in alphabetical order, complete synonymy is given. Most types (or images of
types) of names of Geonoma have been examined for this study and these are followed by a !. Those which
have not been examined are followed by n.v.. Excluded names are listed in Appendix III. Plates of type
images of all new taxa are given in Appendix IV, and images of types of new taxa deposited at NY are
available at the website http://www.nybg.org/bsci/herbarium_imaging/. A numerical list of taxa and a list of
specimens examined, ordered by collector, are given in Appendix V.
Phylogeny
For phylogenetic analysis of the species of Geonoma, three outgroups were usedAsterogyne spicata
(Moore) Wessels Boer (1968: 82), A. yaracuyense Henderson & Steyermark (1986: 309), and A. martiana
(Wendland) Wendland ex Drude (1889: 59)based on recent, molecular-based, family-level studies (e.g.,
Asmussen et al. 2006, Baker et al. 2009) and tribal-level analyses (Roncal et al. 2005). The ingroup contains
the 68 species of Geonoma recognized here, and the characters used are those identified by the methods
discussed above (Appendix I). Some authors have suggested using quantitative variables in phylogenetic
analysis (e.g., Goloboff et al. 2006). This was attempted, using the coding method of Thiele (1993), but the
results gave no resolution. Therefore only characters were used.
The phylogeny data matrix (Table 1) was constructed and edited with Winclada (Nixon 19992002).
Parsimony analyses were conducted with Nona (Goboloff 1999). The Multiple TBR + TBR(mult*max*)
search strategy was used with the following settings: hold = 100,000, mult*N = 1000, and hold/ = 10. A strict
consensus tree was calculated in WinClada. Branch support was calculated using Bootstrap analysis with
1000 replications, 10 random entry sequences, and one tree saved per replication.
TABLE 1. Phylogeny Matrix ( = missing data)

A.
A.
A.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
spicata
yaracuyense
martiana
aspidiifolia
baculifera
bernalii
braunii
brenesii
brongniartii
calyptrogynoidea
camana
chlamydostachys
chococola
concinna
concinnoidea
congesta
cuneata
deneversii
deversa
dindoensis
divisa
elegans
epetiolata
euspatha
ferruginea
fosteri
frontinensis
galeanoae
gentryi
hollinensis
hugonis
interrupta
lanata
laxiflora
lehmannii
leptospadix
longipedunculata
longivaginata
macrostachys
maxima
monospatha
mooreana
multisecta
occidentalis
oldemanii
oligoclona
operculata
orbignyana
paradoxa
pauciflora
peruviana
pinnatifrons
poeppigiana
pohliana
poiteauana
sanmartinensis
santanderensis
schizocarpa
schottiana
scoparia
simplicifrons
spinescens
stricta
talamancana
tenuissima
triandra
triglochin
trigona
umbraculiformis
undata
venosa
0
5
10
15
20
25
|
|
|
|
|
|
01011101011111012100111101100
01111101011111012100111101100
01011101011111011100111101100
101111100011101011101211101100
001110101111101011001000101111
001111101131101111001011101100
001101101131101111001011101000
001011111011001011001211101000
001011101011000021001011101100
001110101011101011001000100111
001011101011001011311111101110
01011101011001011311111101111
001011101011001011311111110111
001110101111001011001011101110
001110101111001011001011100110
001110101011001011001000100111
001011101011011011001011101200
010111010110010123111
001111101101100011001011101100
00111110113110111100101
001101101101101011101211101200
001011101011101011001010001100
011011101011001011001211101200
00111110111210001001011101000
001101101011101011101011101000
00110111131101111001011101000
0111110101210001001011101000
0011101011111000110010111010
0011101111101111001011101000
001111111101100000001011101000
101011101011001011001211101000
00111100111010001001011101000
001111101131101111001011101000
001101101111100011001011101100
001011101011201011001010101000
001111101111100011001011101100
001111101111100021001011101100
001101101111101011101011101000
001011101011001011221111101010
001111101011001011311111101100
001111111011001011001211101000
001101101131101011001011101100
001011101011001012221111101110
001101101121100011001011101000
011011101011001011311111101111
101101100011101011101211101000
0011011113110111100101101010
001111101011201011001010101000
001011101011001021221111101110
001111101011101011001010001100
20111110111110001100101110100
00111100111210001001011101000
001111101011100021101011101100
001111001011101011001010001100
001011101011001011221111101110
01011101011100021001011101000
101111100011101011101211101000
001011101011001011221111111110
000111101011101011001010001100
201101101131101011001011101100
00111110111010001001011101000
001101111101111001011101000
001111110011101011101011101200
001011121011201011001010101000
001101101131101111001011101100
001111111121100010001011101000
011111101011001011311111101111
01111101011201001001010101000
011111101011001011311111111111
001111001011201011001010101000
011131101111001011101100
HENDERSON
Phylogeny
Results
Heuristic analysis generated 6960 most parsimonious trees (MPTs) of 102 steps with a consistency index (CI)
of 0.42 and a retention index (RI) of 0.80. Figure 1 represents one MPT. Thirty-four nodes collapsed in the
strict consensus tree of 6960 MPTs (Fig. 2).
Geonoma is resolved as monophyletic. There is a dichotomy at the outgroup node leading to two,
unequal-sized, well-supported clades. The smaller of these, the G. macrostachys clade (BS=93%) (all clades
in the following discussion are named after their most widespread species), consists of 13 species. It shares
with the outgroup the character state of staminodial tubes with spreading, acuminate lobes. This clade
contains two well-supported subclades and five other species that are not resolved. The three species of one
subclade (G. oldemanii, G. triglochin, and G. umbraculiformis), without non-homoplasious synapomorphy,
occur in both the extreme western and eastern Amazon. The five species of the second subclade (G.
macrostachys, G. multisecta, G. paradoxa, G. poiteauana, G. schizocarpa) share the non-homoplasious
character states of the thecae inserted onto jointed connectives, and the anthers short at anthesis and remaining
straight and parallel. These species, with the cis-Andean exception of G. paradoxa, are widely distributed in
the Amazon region.
The larger clade of the basal dichotomy contains all other species (BS=89%). It is divided into two clades.
The first, the G. cuneata clade (BS=88%), contains five species (G. cuneata, G. epetiolata, G. hugonis, G.
brenesii, G. monospatha) without non-homoplasious synapomorphy. These species are mostly Central
American, although G. cuneata also ranges all along the Pacific coast and adjacent areas of Colombia and
Ecuador, and just reaches Venezuela.
All other species of Geonoma occur in one large clade (BS=87%). In this there are seven well-supported
subclades, but sister group relationships amongst these are not resolved in the consensus tree (Fig. 2) and nine
species are unresolved.
The first subclade, the G. triandra clade (BS=89%), contains three species (G. occidentalis, G. hollinensis,
G. triandra) without non-homoplasious synapomorphy. The last two species share the non-homoplasious
character state of staminate flowers with three stamens, and have a cis-trans-Andean distribution pattern. They
correspond to Wessels Boers (1968) subgenus Kalbreyera (see below).
The second subclade, the Geonoma schottiana clade (BS=93%), consists of four species (G. elegans, G.
pauciflora, G. pohliana, G. schottiana), sharing the non-homoplasious character state of ovoid fruits. These
species are endemic to the Cerrado and Atlantic Coastal Forest of Brazil and just reach Bolivia and Paraguay.
The third subclade, the Geonoma undata clade (BS=92%), consists of five species (G. orbignyana, G. trigona,
G. undata, G. lehmannii, G. talamancana) mostly confined to higher elevations in Central and northern and
western South America, and just reaching the Lesser Antilles. It corresponds to Roncal et al. s (2005) high
elevation clade (see below). The species of this clade share the non-homoplasious character state of apiculate
and lobed proximal lips of the flower pits.
The fourth subclade, the Geonoma congesta clade (BS=91%), consists of five species. These share the
non-homoplasious character state of the prophyll surfaces with close, equal, parallel, non-dividing ridges,
although this character state also occurs in G. galeanoae (unresolved for subclade in the strict consensus tree;
included in this subclade in the MPT shown in Fig. 1). This subclade consists of two well-supported clades.
The first, with two species (G. concinna, G. concinnoidea) occurs in Panama and Colombia. The second, with
three species (G. baculifera, G. calyptrogynoidea, G. congesta) shares the non-homoplasious character state of
staminodial tubes of non-fertilized pistillate flowers projecting and persistent after anthesis. It has a disjunct
distribution and occurs in Central America (G. congesta), western South America (G. calyptrogynoidea), and
the eastern Amazon region (G. baculifera).
The fifth subclade, the Geonoma interrupta clade (BS=90%), consists of five species (G. euspatha, G.
frontinensis, G. interrupta, G. pinnatifrons, G. simplicifrons). Two species (G. interrupta, G. simplicifrons)
share the non-homoplasious character state of the flower pits densely hairy internally distally only; and the
. . . . . . Figure 1 continued on the next page
10
HENDERSON
Figure 1 continued from the last page
FIGURE 1. One of 6960 MPTs. Character changes are labeled above each horizontal line with numbers corresponding
to Appendix I. Closed circles represent non-homoplasious changes, open circles represent homoplasious changes.
other three species have the flower pits densely hairy internally proximally and distally. Species in this clade
are widely distributed in Central America, the Amazon, Andean, and Choc regions of South America, and
just reach Hispaniola and the Lesser Antilles.
Species of the sixth subclade, the Geonoma stricta clade (BS=87%), do not share any non-homoplasious
synapomorphy. This subclade contains two well-supported clades. The first, containing three species (G.
ferruginea, G. divisa, G. longivaginata) occurs in Central America and just reaches the Pacific coast of
Colombia. The second, containing four species (G. stricta, G. aspidiifolia, G. oligoclona, and G.
santanderensis) is widespread but mostly occurring in the Amazon region. Species shares the nonhomoplasious character state of rachillae surfaces with spiky, fibrous projections or ridges.
The seventh subclade, the Geonoma lanata clade (BS=88%), contains 12 species (G. spinescens, G.
gentryi, G. operculata, G. braunii, G. fosteri, G. lanata, G. bernalii, G. dindoensis, G. venosa, G. tenuissima, G.
mooreana, G. scoparia). These are narrow endemics in mostly montane areas in Costa Rica, Panama,
Venezuela, Colombia, and Ecuador. Most of these species occupy only one degree square and the most
11
widespread species, G. lanata, only occupies four degree squares. A well-supported clade within this seventh
subclade has 10 species which share the non-homoplasious character state of the flower pits alternately
arranged, and they also share the trait state of the rachillae being filiform with extended narrowed sections
between the flower pits.
Discussion
There have been two previous attempts to understand species relationships in Geonoma. Wessels Boer (1968),
using pre-cladistic methodology, placed species into two subgenera. Subgenus Kalbreyera contained one
species, G. triandra. Subgenus Geonoma contained two sections, 16 species groups, and 74 species. In the
present study, neither subgenus nor sections are found to be monophyletic, and only three of the 17 species
groups are monophyleticthe groups numbered 4, 8, and 11 by Wessels Boer.
The only previous phylogeny of the geonomoid palms using cladistic methodology is that of Roncal et al.
(2005), based on molecular data. These authors included 30 species of geonomoid palm, of which 20 were
Geonoma. They found that Geonoma species formed a strongly supported monophyletic group. Roncal et al.
found two strongly supported clades within Geonoma. One of these consisted of three high elevation
speciesG. undata, G. orbignyana, and G. jussieuana (here treated as part of G. orbignyana).
. . . . . . Figure 2 continued on the next page
12
HENDERSON
Figure 2 continued from the last page
FIGURE 2. Strict consensus tree of 6960 MPTs. Bootstrap values are shown above horizontal lines.
13
The second well-supported clade consisted of six speciesG. macrostachys, G. polyandra (here named G.
multisecta), G. congesta, G. triglochin, G. maxima, and G. camana. These species are also members of a wellsupported clade in the present study, with the exception of G. congesta. This species was represented in
Roncal et al.s study by voucher specimens Borchsenius 343 and Borchsenius 348 (AAU). Both these
specimens are included in the present study and are identified as G. calyptrogynoidea.
Roncal et al. (2010, in press) have looked at divergence times in the Geonomateae and diversity patterns
in Geonoma.
Although the present study resolves Geonoma as monophyletic, it illustrates the difficulty of finding and
using morphological characters to resolve species relationships in this genus. The relatively low CI of 0.42
indicates a high amount of homoplasy. Although several clades are well-supported, their sister group
relationships are not resolved. This problem is based on the more fundamental difficulty of defining
characters in Geonoma. As discussed in the taxonomic treatment, several characters are problematic. An
example of this is found in character 18 (Appendix I): connectives. One state of this character requires a
somewhat convoluted definition (thecae diverging or not diverging at anthesis, inserted onto poorly to welldeveloped, non-split, jointed connectives, connectives when well-developed alternately long and short)
because if this state was split into two statesconnectives jointed and well-developed versus poorlydevelopedthen virtually identical and sympatric specimens would become separate species. This problem
occurs in the G. macrostachys clade in both G. macrostachys itself and G. paradoxa. Similar problems are
encountered in character 10, pit arrangement, and other characters. Most of these problems occur in species
complexes, discussed further in the section on Infraspecific Variation.
Distribution
Geonoma species are distributed from 1945N (Haiti) to 2946S (Brazil) and 9640W (Mexico) to
3504W (Brazil) (Fig. 3). The country with the highest number of species, 32, is Colombia, followed by
Ecuador with 29 and Brazil with 24. The areas with the highest number of species (10 species per degree
square) are situated along the mountain regions and adjacent areas of Central and western South America, in
Costa Rica (Alajuela), Panama (Chiriqu, Cocl), Colombia (Antioquia, Choc, Valle, Caquet, Putumayo),
Ecuador (Sucumbios), and Peru (Amazonas, Pasco)(Fig. 4). The degree square with the highest number of
species, 14, is situated along the border between Colombia (Caquet) and Ecuador (Sucumbios). These areas
of high species diversity all have annual precipitation of >4000 mm per year. According to Wessels Boer
(1968) no species of Geonoma occurs in areas with <1000 mm annual precipitation.
There are several, large areas, especially in the Amazon region where there are very few collections of
Geonoma (Fig. 3). Most parts of the Brazilian Amazon, except for Acre, are poorly collected. There are three
areas in Amazonian Peru with few collections: 1) the northwestern part of Loreto, parallel to the border with
Ecuador, and including adjacent Colombia, 2) the southern part of Loreto and adjacent Ucayali, including the
subAndean parts of San Martn, and 3) the southern part of Ucayali and adjacent Madre de Dios.
The areas with the highest morphological diversity, as measured by the coefficient of variation (CV) for
all specimens per degree square (CVs of 18.0 per degree square) are similar to the areas with the highest
number of species and are mostly in montane regions. They are situated in the eastern part of the Cordillera
Central in Panama, northern French Guiana, the Pacific coast of Colombia in the Choc and Valle, the eastern
Cordillera in Colombia, the Pacific coast of northwestern Ecuador, eastern Ecuador, and central Peru (Pasco)
(Fig. 4). These areas all have annual precipitation of >4000 mm per year.
Species of Geonoma occur from sea level to 3370 m elevation (G. undata in Ecuador). Of the 3639
specimens with elevation data, 1974 (54%) occur at 500 m and 2606 (71%) occur at 1000 m. Only a few
species consistently occur at >2000 m elevation, all in the G. undata clade (G. lehmannii, G. orbignyana, G.
talamancana, G. trigona, G. undata).
14
HENDERSON
Species of Geonoma are found in a variety of habitats. Most commonly they occur in lowland rainforest,
in both flooded and non-flooded areas. Some taxa have specialized habitats. A few are reported to be
rheophytes (Geonoma cuneata subsp. linearis, Geonoma interrupta subsp. rivalis, Geonoma undata subsp.
pulcherrima, and some populations of G. deversa). Some occur in gallery forest in otherwise non-forested
areas (e.g., G. pohliana subsp. weddelliana, G. schottiana), others in restinga forest (some morphotypes of G.
pohliana subsp. pohliana), others in brejo vegetation (some morphotypes of G. pohliana subsp. pohliana), and
others at high elevation in pajonal or dwarf forest (e.g., G. trigona). Different morphotypes of some species
complexes occupy a variety of habitats, as discussed in the next section on Infraspecific Variation.
FIGURE 3. Distribution of all Geonoma specimens examined.
Infraspecific Variation
There is a wide range in variability amongst species of Geonoma. Some species have relatively little variation
and others are extremely variable and have been termed species complexes. Cronk (1988) considered that this
kind of variation in larger genera was quite common. He divided species into different categories. Monotypic
species, usually the majority of species in a genus, were those that could be clearly distinguished from other
species, even though some of them might be members of groups of closely related species. Polytypic species
were more variable and had two or more local forms although these were usually geographically separate
from one another. Variation in these could be described taxonomically by the use of subspecies. Polymorphic
species (or ochlospecies) were those that had chaotic infraspecific variation which was intractable to formal
taxonomic treatment. Cronk estimated that in three large genera that had been studied in detail, approximately
15
87% of species were monotypic, 10% polytypic, and 3% polymorphic. Cronk discussed various
characteristics of these different species categories, and his discussion serves as a basis for examining
infraspecific variation in Geonoma.
Cronk listed four, defining characteristics of polymorphic species (note that trait has been substituted for
character here, in line with the usage in this revision)variation was strongly polymorphic but only weakly
polytypic; trait state distributions showed only partial correlation with geography and ecology; traits varied
independently and were not correlated; and complexity of variation was not due to hybridization or some
complex breeding system. Cronk also listed six other, ancillary characteristicspolymorphic species were
usually geographically and ecologically widespread; had distinct variants that were recognizable locally but
not globally; often had closely related satellite species; had similar variants that occurred in widely separated
localities and appeared to be independent in origin; occurred in medium to large genera with a low frequency;
and usually had long synonymies.
FIGURE 4. Degree squares with 10 species (crosses); degree squares with CVs of 18.0 per degree square (circles).
Variability, abundance and range

Table 2 lists the 68 accepted species of Geonoma arranged according to the sum of their coefficients of
variation (CVs) for 26 quantitative variables (hereafter referred to as variability). The number of specimens
examined for each species (hereafter abundance), and the number of one degree squares occupied by each
species (hereafter range) are also given. Although number of specimens examined, and number of one degree
squares occupied are not exact measures of abundance and range, and although both are likely to be
16
HENDERSON
influenced by collecting (smaller-sized palms are more likely to be collected than larger ones, and many areas
are poorly collected), they are at least proxies, and are the only measures available and are therefore used here.
Four species are known from only one specimen each, and thus have no CV values. Ten species have CV
values of less than 4, but most of these are represented by only a few specimens. About one third (23) of the
species have CV values between 5 and 7, and 13 species have CV values of over 10.
TABLE 2. Sums of Coefficients of Variation (CVs) of 26 quantitative variables per species. (orders of inflorescence
branching excluded)(variability), number of specimens examined (abundance), and number of one degree squares
occupied by each species (range).
Species
Variability (sum of CV of
variables)
Abundance (number
of specimens
examined)
Range (number of
degree squares
occupied)
G. deneversii
G. dindoensis
G. gentryi
G. operculata
G. fosteri
0.900
G. venosa
1.151
G. peruviana
1.234
G. spinescens
2.708
G. schizocarpa
2.996
10
G. hollinensis
3.505
G. sanmartinensis
3.514
G. trigona
3.525
G. bernalii
3.610
G. galeanoae
3.650
G. tenuissima
4.030
12
G. lanata
4.849
28
G. chlamydostachys
4.924
G. concinna
5.101
G. oldemanii
5.150
12
G. umbraculiformis
5.217
13
G. scoparia
5.234
G. poiteauana
5.356
28
22
G. brenesii
5.430
13
G. laxiflora
5.518
36
19
G. leptospadix
5.599
57
40
G. epetiolata
5.694
38
G. divisa
5.713
10
G. hugonis
5.846
32
G. occidentalis
5.995
29
15
......continued on the next page
17
TABLE 2 (continued)
G. talamancana
6.041
12
G. santanderensis
6.191
G. calyptrogynoidea
6.294
36
14
G. braunii
6.526
G. aspidiifolia
6.562
32
G. concinnoidea
6.573
30
G. frontinensis
6.663
23
11
G. longipedunculata
6.722
35
10
G. monospatha
6.831
14
G. simplicifrons
6.894
29
G. baculifera
6.974
95
50
G. oligoclona
7.020
14
10
G. mooreana
7.242
G. paradoxa
7.406
16
G. elegans
7.511
75
21
G. triandra
7.802
19
G. ferruginea
7.984
77
G. congesta
8.065
79
18
G. chococola
8.251
32
G. euspatha
8.486
63
37
G. pauciflora
8.797
37
11
G. schottiana
8.963
76
28
G. multisecta
8.999
24
G. deversa
9.383
356
162
G. poeppigiana
9.475
83
23
G. triglochin
9.539
35
10
G. pinnatifrons
10.020
245
82
G. brongniartii
10.076
165
55
G. lehmannii
10.133
118
22
G. interrupta
10.196
154
60
G. camana
10.223
65
29
G. longivaginata
10.720
65
G. pohliana
11.344
241
70
G. orbignyana
11.731
248
58
G. maxima
11.939
373
135
G. cuneata
12.423
349
45
G. stricta
12.839
504
98
G. undata
13.964
386
102
G. macrostachys
14.782
343
73
18
HENDERSON
For abundance, 12 species are known from five or fewer specimens. These species have ranges of 13
degree squares and are mostly local endemics from montane areas in Central and South America. Linear
regression shows there is a significant, positive association between abundance and variability. Squared
multiple R for the regression of variability on abundance is 0.56. The more abundant a species, the more
variable it is. Geonoma stricta has large leverage because it is by far the most abundant species.
For range, 31 species occur in five or fewer degree squares, 48 occur in 20 or less, and only three species
occur in more than 100. Linear regression shows there is a significant, positive association between range and
variability. Squared multiple R for the regression of variability on range is 0.43. The greater the range of a
species, the more variable it is. Geonoma deversa has large leverage and is an outlierit has the widest range
of any species, but a relatively low CV. Geonoma maxima also has large leverageit has the second widest
range. Three speciesG. laxiflora, G. leptospadix, G. baculiferaare relatively widespread (occur in > 40
degree squares) but have relatively low variability (5.5186.974).
Combining variability and range, there are 10 species which have CVs of over 10 and ranges of over 40
(G. brongniartii, G. cuneata, G. interrupta, G. macrostachys, G. maxima, G. orbignyana, G. pinnatifrons, G.
pohliana, G. stricta, and G. undata). Three other species have CVs over 10 but have relatively narrow ranges
(G. lehmannii, G. camana, and G. longivaginata). The two last two of these have high CVs because of their
unusually high value for basal pinna width (this variable has one of the highest CVs for all species combined).
Of the 10 most variable and widespread species, all but one (G. macrostachys) are divided into subspecies.
In some of these (e.g., G. interrupta, G. maxima, G. pinnatifrons, G. pohliana) variation appears polytypic in
that the different morphotypes are mostly allopatric and can be recognized as subspecies based on the criteria
discussed in the Materials and Methods section. Geonoma interrupta, G. maxima, G. pinnatifrons, and G.
pohliana can be considered polytypic species. However, designation of species as polytypic appears
somewhat arbitrary, because there are many other species which are here divided into subspecies, several of
them with relatively low variability, low abundance, and narrow ranges (e.g., G. concinna and G. aspidiifolia).
The remaining variable species (G. brongniartii, G. cuneata, G. macrostachys, G. orbignyana, G. stricta,
and G. undata) are either undivided taxonomically but with various morphotypes (G. macrostachys), or
divided into a mixture of subspecies and morphotypes. These species could be considered polymorphic.
However, they do not appear to satisfy all of Cronks four defining characteristics.
The first of thesethat variation is strongly polymorphic but only weakly polytypicappears to apply to
these variable Geonoma species. In fact, one of the most obvious aspects of these species is that two or more
morphotypes may occur at the same locality. On the other hand, the second and third characteristicsthat trait
state distributions show only partial correlation with geography and ecology, and traits vary independently
and are not correlateddo not always apply. In all of these species it is possible to recognize morphotypes,
and almost by definition these have correlated traits. For example, in G. stricta there is a widespread and
consistent morphotype, trailii, which has pinnate leaves and raised abaxial veins. In G. macrostachys there is a
widespread morphotype, macrostachys, with mostly undivided leaves and non-raised veins. The problem in
these species is that in some areas there are specimens with intermediate trait distributions, such that division
into taxa is not possible. Over their entire range these species exhibit a mosaic of variation that may be both
polytypic and polymorphic, correlated or not correlated, and correlated or not correlated with geography.
Explanations of variability in Geonoma
The fourth of Cronks defining characteristics, in part, was that complexity of variation in polymorphic
species was not due to some complex breeding system. This seems to be the case in Geonoma. There appears
to be no evidence for any cytological or breeding system abnormality in Geonoma species (e.g., polyploidy,
apomixis). They have a chromosome number of 2n = 28 (although only 3 species have been counted), and
there is no evidence of polyploidy (Dransfield et al. 2008). Plants of Geonoma are monoecious and
protandrous, with unisexual flowers borne in triads of one pistillate and two staminate. Where known, they are
self-compatible, outcrossing, and have a variety of pollinators, including bees (especially euglossine bees),
flies, beetles, and wind (reviewed in Henderson 2002).
19
However, the second part of Cronks fourth characteristic was that complexity of variation in polymorphic
species was not due to hybridization. This may not be the case in Geonoma. Hybridization, including
introgression, seems to be widespread and may contribute to variation. In this study, many specimens have
been considered to be hybrids, based on morphological intermediacyand almost all hypothesized cases of
hybridization are found in the most variable species. Nevertheless, it is difficult to demonstrate hybridization
and introgression using morphological data taken from herbarium specimens.
One of the most compelling instances is the presence of a potential, widespread hybrid zone in the eastern
Amazon region between G. deversa and G. leptospadix. There may be other examples of hybrid zones, such as
between G. undata subsp. edulis and G. orbignyana subsp. hoffmanniana in Costa Rica and Panama, and
between G. poeppigiana and G. brongniartii in the western Amazon basin. There are also many examples of
sporadic hybrids between different species, subspecies, and morphotypes within species.
There are other processes that might account for some of the variation seen in polymorphic species of
Geonoma. Geographical variation along environmental or spatial gradients is apparently quite common. One
of the commonest examples of geographical variation concerns leaf shape and elevation. For example, in the
related, variable species G. undata and G. orbignyana, occurring in montane areas, it is common for basal
pinna to become narrower with narrower angles with increasing elevation. A similar change is seen in G.
pohliana subsp. weddelliana, where the number of pinnae increases and they become shorter and narrower
with narrower angles with increasing elevation, giving a distinctive, pinnate leaf. In three unrelated species
occurring in the western Amazon region there is geographical variation with longitude in leaf shape. In G.
brongniartii, G. camana, and G. poeppigiana, there is variation from plants in the in the eastern part of the
range having fewer, wider and longer pinnae with narrower angles to plants in the western part having more,
narrower and shorter pinnae with wider angles. However, these kinds of geographical variation are based on
limited sample sizes and with many missing data, and seem unlikely to contribute greatly to variation within
polymorphic species.
Variation in Geonoma macrostachys
None of the processes discussed above seem to explain satisfactorily the situation in variable species of
Geonoma. For the most variable species, G. macrostacys, there is quite a lot of comparative data on
pollination and habitat preference and this is reviewed here. In the last taxonomic treatment (Henderson
1995), G. macrostachys was divided into three varieties. In the present treatment, one of these varieties is
recognized at the species level (G. poiteauana). The other two varieties are recognized as morphotypes, along
with seven other morphotypes. In the following discussion, these morphotype names are used (based on
voucher specimens included in this revision) instead of the names used by the original authors.
Listabarth (1993) studied pollination of two sympatric morphotypes of Geonoma macrostachys in
Hunuco, Perularge-size and tapajotensis. He found that the two had different phenology and pollinators
the large-size morphotype flowered from December to July and was pollinated mostly by meliponine bees and
syrphid flies whereas the tapajotensis morphotype flowered from September to February and was pollinated
by weevils. Variation in floral scents among different morphotypes of G. macrostachys has been documented
by Knudsen (1999). Roncal (2006) looked at the habitat preferences of four morphotypes of G. macrostachys
at three sites in Peru (Table 3).
In Loreto and Madre de Dios, there is a clear habitat preference of tapajotensis for floodplains, and a
slightly less marked preference of macrostachys for terra firme. In Pasco, acaulis has a preference for
floodplains, and to a lesser extent, large-size for terra firme.
There are several other studies on habitat of G. macrostachys (Kahn & de Granville 1992, Svenning 1999,
Vormisto et al. 2004), confirming habitat preferences for morphotypes of G. macrostachys. There is thus
abundant evidence that at individual sites, different morphotypes have different habitat preferences. In
particular, the wide-angled, pinnate leaved morphotypes, tapajotensis and acaulis, have a preference for
flooded habitats, and the narrow-angled, undivided or few-pinnate leaved forms, macrostachys and large-size,
have a preference for non-flooded habitats. However, there are some conflicting reports. Vormisto et al.
20
HENDERSON
(2004) reported that in Peru acaulis was more abundant than expected on hills and less abundant at lower
positions, and macrostachys was more abundant in valleys and less so on hills (no vouchers have been seen
from this study). Svenning (1999) found that in Ecuador macrostachys, the most abundant palm at the site did
not show any topographic preference. In summary, the data seem to indicate that at the same site different
morphotypes have different habitat preferences, but at different sites the same morphotypes may have
different habitat preferences.
TABLE 3. Number of individual plants per hectare in different habitats at different sites (from Roncal, 2006).
*Identification tentative because staminate flowers lacking.
tapajotensis
macrostachys
acaulis*
largesize
terra firme, Loreto
321
floodplain, Loreto
2805
57
terra firme, Madre de

Dios
221
floodplain, Madre de
Dios
29
72
terra firme, Pasco
12
36
floodplain, Pasco
200
11
Roncal et al. (2007) studied the genetic distinctness of three morphotypes at four sites in Peru using
intersimple sequence repeat variation. Results showed that specimens formed two main, well-supported
clusters. In one main cluster, specimens of the large-size morphotype from the two closest sites, Pasco and
Hunuco, clustered together along with Hunuco tapajotensis. In the second main cluster, specimens of the
macrostachys and tapajotensis morphotypes from the two most distant sites, Loreto and Madre de Dios,
clustered together. There were two exceptions to this pattern from Loreto. One specimen clustered with the
Pasco/Hunuco cluster, and another appeared outside both clusters. One of these specimens is the
macrostachys morphotype and the other is unclear.
Despite the problems of the identity of some morphotypes, these results indicate that the three
morphotypes macrostachys, large-size, and tapajotensis are not homogeneous and cannot be considered as
taxa. Specimens of large-size and tapajotensis from Pasco/Hunuco cluster together, indicating at least in this
case, that parallel, local phenotypic diversification has taken place, resulting in similar but unrelated
morphotypes in different areas. It is also interesting to note that Listabarths (1993) study was carried out in
the same Hunuco locality, where the two morphotypes (large-size and tapajotensis) were shown to have such
different phenology and pollination.
Finally, for Geonoma macrostachys, there is unexplained variation in staminate flowers. As noted under
that species, there appear to be two different kinds of connective, poorly-developed and well-developed, and
these two can occur together at the same locality.
In summary, Geonoma macrostachys is an extremely variable, abundant, and wide ranging species, and
has very complex patterns of variation. Based on traits and quantitative variables of leaves it can be divided
into at least nine different morphotypes. These exhibit a wide range of variation in leaf size and shape, from
small or large undivided leaves with narrow basal angles to small or large pinnate leaves with wide basal
angles (the variables for this species with the highest CVs are petiole length, rachis length, number of pinnae,
basal pinna width, and basal pinna angle). However, these morphotypes are probably not taxa, and the same
morphotype may have arisen independently in different places. Where two or more different morphotypes
occur together at a site, they appear to have different phenology, pollination, habitat preference, and some may
even have different floral morphology. They appear and behave as distinct species. On a larger scale, the
21
differences between the morphotypes break down because of intermediate morphotypes. Within the
macrostachys morphotype, for example, the majority of specimens have small, narrow, undivided leaves with
narrow basal angles. But there are some specimens, from scattered localities, with similar but slightly
different leaves, sometimes larger, sometimes pinnate, sometimes broad with wide angles, sometimes with
raised adaxial veins. These are intermediate between the macrostachys morphotypes and other morphotypes.
There is no evidence that these intermediates are hybrids (in that they can appear in areas without the putative
parents), although there may be hybrids between morphotypes.
Hypotheses of variation
Cronk (1988) put forward two hypotheses to explain polymorphic species; the refugia hypothesis and the
rapid expansion hypothesis (neither of which is mutually exclusive). The refugia hypothesis is still
controversial (e.g., Bush & de Oliveira 2006). The rapid expansion hypothesis was based on the idea that a
species capable of rapid colonization and with wide ecological tolerance could greatly expand its range. It
would then come under diverse selection pressures, such as genetic drift in founder populations and rapid
selection in new habitats, leading to geographically unstructured variation.
A third possible hypothesis that could apply to all polymorphic species is the resource polymorphism
hypothesis. Smith & Sklason (1996) defined resource polymorphisms, for vertebrates, as the occurrence of
discrete intraspecific morphs showing differential niche use, usually through discrete differences in feeding
biology and habitat use. Morphs may differ in morphology, color, behavior, or life history traits, and in many
instances they may differ in more than one characteristic. If one substitutes pollination biology for feeding
biology in this definition, then one has a plausible definition of the situation in G. macrostachys and other
polymorphic Geonoma species.
Smith & Sklason considered that three conditions promoted resource polymorphismsthe existence of
open niches, habitat variability, and the relaxation of interspecific competition. They considered that a process
involving invasion of an unexploited niche and relaxation of interspecific competition followed by divergent
selection would lead to the evolution of polymorphism, with different morphotypes adapted to different
resources. Further divergent selection and reduced gene flow could lead to reproductive isolation,
representing a step toward sympatric speciation
This hypothesis, not excluding elements of the refugia and rapid expansion hypotheses, could explain
polymorphism in Geonoma macrostachys, and probably other species complexes. It is well documented that
forests in the western Amazon basin are heterogeneous for habitat. Salo et al. (1986) considered that up to one
quarter of forest areas in the Peruvian Amazon were a mosaic of different stages of succession caused by river
bank lateral erosion. Contemporary forests exhibit habitat diversity at the same site, based on edaphic factors
and light levels, and also in forest structure and diversity (e.g., Haugaasen & Peres 2006). Roncal (2006)
found that soil factors were more important than light factors in the distribution of Geonoma morphotypes.
This forest heterogeneity presumably exists in other regions where polymorphic Geonoma species occur.
In conclusion, there is a continuum in variability amongst Geonoma species, from those with very low to
those with very high variability, and this is correlated with data on abundance and range. It may not be
possible to assign species to different categories. Polytypic species can have both low (e.g., G. aspidiifolia)
and high (e.g., G. pinnatifrons) variability, and both narrow (e.g., G. longivaginata) and wide (e.g., G. maxima)
ranges. Designation of species as polymorphic also appears somewhat arbitrary, and the kinds of patterns of
variation seen, for example, in G. macrostachys, may occur, to a lesser extent, in many species. Nevertheless,
the species with the highest variability, perhaps 20% of all species of Geonoma, are difficult taxonomically
and their infraspecific variation cannot be understood with morphological data taken from herbarium
specimens. The high levels of variability in these species may be based on resource (habitat) polymorphisms,
and these may represent an intermediate step in sympatric speciation.
22
HENDERSON
Morphology
The genus description given below in the Taxonomic Treatment section is based on the list of characters and
traits used in this study (Appendix I). In the following discussion, morphology is treated in more detail, and
the morphology of several attributes of Geonoma not used in delimiting species is discussed, particularly
flowers and fruits.
Stems of Geonoma are solitary or clustered, and this trait often varies within species. Sixteen species are
reported to have consistently solitary stems and 10 species to have consistently clustered stems. All others
have both solitary and clustered stems. Geonoma are usually rather small palms, and the mean size of stems of
all specimens is 1.9 m tall and 1.0 cm diameter. Internodes, which are usually yellowish and smooth, rarely
brown scaly (Fig. 5A), may be longer than wide, giving a cane-like stem (Fig. 5A), or wider than long, giving
a non-cane-like stem which is often short and subterranean. Cane-like stems are a mean of 1.9 m long, 0.9 cm
wide, and have 2.3 cm long internodes, whereas non-cane-like stems are 1.1 m long, 1.7 cm wide, and have
0.6 cm long internodes.
Leaves of Geonoma are undivided (Fig. 5B) or pinnate (Fig. 5C). There is a high level of variation within
species, sometimes within plants, in leaf division. Only 11 species have consistently undivided leaves whereas
29 species have both undivided and pinnate leaves. All other species have pinnate leaves. These are usually
irregularly pinnate (i.e., the pinnae are unevenly wide) or less often regularly pinnate and then the pinnae
usually with one main vein only. Rarely, in regularly pinnate leaves, the pinnae have one main vein and two
lateral veins on either side of main vein (Fig. 5D). The mean number of pinnae per side of the rachis in pinnate
leaved plants is seven. Leaves may be plicate (Fig. 5E) and this condition is found mostly in species from
higher elevations. The mean elevation of plicate-leaved specimens is 2063 m. The bases of leaf blades usually
run diagonally into the rachis but in a few species are recurved against the rachis (Fig. 5F). Petioles and rachis
dry green or yellowish, but sometimes they dry orange-brown or reddish-brown (Fig. 5G). There is a distinct
hastula on the adaxial surface of the rachis, but this does not appear to have any taxonomic value (but see
Henderson, 2005b). Veins are either raised and rectangular in cross-section adaxially (Fig. 5H) or not raised or
slightly raised and triangular in cross-section adaxially (5I).
Inflorescences are either interfoliar or infrafoliar in Geonoma, but this is neither consistent within species
nor easy to score from specimens. Furthermore, inflorescences can be interfoliar at anthesis and infrafoliar in
fruit. There are two bracts on the inflorescencea prophyll and a peduncular bract (and usually a few, smaller
bracteoles on the peduncle). The form of these bracts is useful taxonomically. In most species with
unbranched inflorescences the prophyll and peduncular bract are ribbed with elongate, unbranched fibers and
both bracts are tubular, narrow, elongate, closely sheath the peduncle, and are more or less persistent. (Fig.
6A). The prophyll itself also exhibits useful characters. In some species the prophyll is short, asymmetrically
apiculate, and the margins curve around the stem. The surfaces are flat with dense, felty, brown tomentum
(Fig. 6B). In other species, prophyll surfaces may be ridged with close, equal, parallel, non-dividing ridges
(Fig. 6C), or they may be ridged with unequally wide ridges, these dividing from and rejoining other ridges. In
this case, the prophylls usually split irregularly between the ridges (Fig. 6D) and the margins have irregular,
spine-like projections (Fig. 6E). Peduncular bracts may be vestigial (Fig. 6F) or rarely absent.
Inflorescences are either unbranched or branched. The number of rachillae per inflorescences varies
widely in Geonoma. Eighteen species have consistently unbranched inflorescences. There are several other
species with mostly unbranched inflorescences but with a few individuals with branched inflorescences.
There are also a few species which can have high numbers of rachillaeup to 138 rachillae have been
counted on one inflorescence of G. scoparia. Rachillae surfaces can be spiky with fibrous projections or ridges
(Fig. 6G) or have faint to pronounced, short, transverse ridges (Fig. 6H). Rachilla width varies considerably,
with some species having filiform rachillae with extended narrowed sections between the flower pits (Fig.
6H). Rachillae are often covered with various hairs but these have not been found useful taxonomically.
23
FIGURE 5. A. Internodes covered with brownish scales, especially in their distal part; stem cane-like, the internodes longer than
wide. B. Leaf undivided. C. Leaf irregularly pinnate, some pinnae with 1 main vein only. D. Pinna with 1 main vein and 2 lateral veins
on either side of main vein. E. Leaves plicate. F. Base of leaf blade recurved against the rachis (arrow). G. Rachis drying reddishbrown. H. Veins raised and rectangular in cross-section adaxially; leaf pinnate. I. Veins not raised or slightly raised and triangular in
cross-section adaxially; leaf undivided. Scale bar = 1 cm.
24
HENDERSON
FIGURE 6. A. Prophylls and peduncular bracts ribbed with elongate, unbranched fibers, both bracts tubular, narrow, elongate, closely
sheathing the peduncle, more or less persistent. B. Prophyll short, asymmetrically apiculate, the margins curved around the stem, the
surfaces flat with dense, felty, brown tomentum. C. Prophyll surface ridged, with close, equal, parallel, non-dividing ridges, scarcely
tomentose between the ridges. D. Prophyll surface ridged, the ridges unequally wide, dividing from and rejoining other ridges, the
prophylls usually splitting irregularly between the ridges. E. Prophyll margins with irregular, spine-like projections. F. Peduncular
bract vestigial (arrowed), the prophyll three times or more long; stem cane-like, the internodes longer than wide; internodes yellowish
and smooth. G. Rachilla surface with spiky, fibrous projections or ridges. H. Rachillae drying brown, the surfaces with faint to
pronounced, short, transverse ridges; rachillae filiform with extended narrowed sections between the flower pits; flower pits
alternately arranged; proximal and distal lips joined laterally with no clear gap between them, forming a raised cupule, the margins not
overlapping. Scale bar = 1 cm.
25
Geonoma is monoecious with unisexual flowers arranged in triads borne in pits in the rachillae. The
arrangement of the flower pits is very variable. Most commonly they are spirally arranged (Fig. 6G), but they
may be alternately arranged (Fig. 6H), tricussately arranged (Fig. 7A), or decussately arranged (Fig. 7B). Pits
have proximal and distal lips, although the latter are sometimes lacking (Fig. 7C). Proximal lips appear to
represent modified triad bracteoles (i.e., the outer bracteole of the triad, as illustrated in Fig. 1.7ag of
Dransfield et al. 2008). The lateral margins of the proximal lip overlap the distal, just as the outer bracteole of
the triad does. The proximal lip often has a central notch before anthesis (Fig. 7D). The distal lip appears to
represent part of the rachilla and not any bracteole. In a few species the proximal and distal lips are not clearly
separate and appear to form a raised cupule (Fig. 6H). Within the flower pit there are three further, ciliate
bracteoles, each subtending a flower. In some species the flower pits are densely hairy internally distally, or
proximally and distally. In a small number of other species (e.g., G. brongniartii) there are a few hairs along
the margins of the proximal and distal lips within the pit, but the pits are never densely hairy.
Staminate flowers of Geonoma have three, free, imbricate, narrow, keeled sepals. The three petals are
connate proximally for about half their length into a tube, free and valvate distally. In a few cases, staminate
petals appear valvate throughout (Fig. 7E). Staminate flowers are usually deciduous after anthesis, rarely
persistent (Fig. 7F). There are six stamens, rarely three or more than six. The filaments are united into a tube
proximally and are free distally. The connection of the anther to filament, and form of the anther are rather
complex in Geonoma. The connective (i.e., the part of the stamen that connects filament to anther) is
sometimes bifid and then scarcely (Fig. 7G) to well-developed. On bifid connectives the thecae (i.e., the two
locules of a single anther) are clearly separate from one another and diverge at anthesis. Typically these
separate thecae curl over at anthesis (Fig. 7G). In other species the connective is not bifid and is attached to
the apex of the filament by a joint (Fig. 7H), so that before anthesis the connective and anthers are inflexed in
relation to the filament. Jointed connectives are again scarcely to well-developed. Well-developed, jointed
connectives appear to have alternately long and short connectives. On non-bifid connectives the thecae are
parallel to one another and not obviously split (i.e., non-diverging). In a third variation, thecae are diverging
but inserted directly onto the apex of the filament, without any obvious connective (Fig. 7I). In this case the
thecae do not curl over at anthesis. There is a small pistillode at the base of the stamen tube, although in some
species (e.g., G. leptospadix) this is well-developed. Pollen of Geonoma is ellipsoid, sulcate, with perforate or
rugulate exine (Dransfield et al. 2008). Where known, species of Geonoma are protandrous (see Henderson
2002 for a review of reproductive biology in Geonoma).
Pistillate flowers of Geonoma have sepals and petals similar in form to those of the staminate flower. The
staminodes form a tube and the apex of the tube is a useful character. In most species the staminodial tubes are
crenulate or shallowly lobed at the apex (Fig. 8A). In a smaller group of species, the staminodial tubes are
lobed at the apex and the lobes are acuminate and spread star-like at anthesis (Fig. 8B). In a third, even smaller
group of species, the staminodial tubes are lobed at the apex but, the lobes do not spread at anthesis and they
are not acuminate (Fig. 8B). In three species, the staminodial tubes of non-fertilized pistillate flowers project
and are persistent after anthesis (Fig. 8C). Stauffer and Endress (2003), who also discussed morphology of
pistillate flowers of Geonoma, considered that one of these species, G. baculifera, had an apically swollen and
slightly calyptrate staminodial tube. The gynoecium is tricarpellate but only one carpel develops. The style is
basally inserted and elongate, exerted above the mouth of the staminodial tube.
Fruits are variously shaped in Geonoma, mostly globose to ellipsoid. They are usually purple black, but
some species have blue (e.g., some morphotypes of G. stricta) or reddish (e.g., G. aspidiifolia) fruits. Wessels
Boer (1968) used fruit color as a character, but it has not been found useful here, and is not observable on
specimens and seldom accurately recorded on labels. Fruit bases may have a prominent, asymmetric stipe
(Fig. 8D) and the apices may be conical (Fig. 8E). Fruits commonly have bumpy surfaces from the numerous,
subepidermal, tangential, short fibers present, and these come to a point at the apices (Fig. 8D). In a few
species, the surfaces split deeply and longitudinally at maturity to reveal the mesocarp with a dense layer of
radial fibers (Fig. 8F), and in a few other species the fruit surfaces have the radial fibers emerging so that the
26
HENDERSON
FIGURE 7. A. Flower pits tricussately arranged, the groups of pits closely spaced; proximal lips hood-shaped. B.
Proximal and distal lips drying darker brown than the rachilla; flower pits decussately arranged, the groups not closely
spaced nor consistently arranged throughout the rachillae. C. Flower pits densely hairy internally; proximal lips hoodshaped; distal lips of flower pits absent; flower pits spirally arranged. D. Proximal lips of flower pits with a central notch
before anthesis, the two sides of the notch overlapping, the lips more or less heart-shaped. E. Staminate and pistillate
petals emergent, valvate throughout. F. Staminate flowers persistent post-anthesis. G. Stamens 6; thecae diverging at
anthesis, inserted almost directly onto the filament apices, the connectives bifid but scarcely developed; anthers short and
curled over at anthesis. H. Thecae not diverging at anthesis, inserted onto well-developed, non-split, jointed connectives;
anthers short at anthesis, remaining straight and parallel. I. Anthers not short and curled at anthesis, elongate, remaining
straight. Scale bar = 1 cm.
27
FIGURE 8. A. Staminodial tubes crenulate or shallowly lobed at the apex; proximal lips of flower pits recurved at the
apices after anthesis (arrowed). B. Staminodial tubes lobed at the apex, the lobes spreading at anthesis, acuminate (left);
staminodial tubes lobed at the apex, the lobes not spreading at anthesis, not acuminate (right). C. Staminodial tubes of
non-fertilized pistillate flowers projecting and persistent after anthesis. D. Fruit bases with a prominent, asymmetric
stipe; fruit surfaces bumpy from the numerous, subepidermal, tangential, short fibers present, these coming to a point at
fruit apices. E. Fruits ovoid with conical apices. F. Fruit surfaces splitting deeply and longitudinally at maturity to reveal
mesocarp with dense layer of radial fibers G. Fruit surfaces with fibers emerging. H. Locular epidermis with operculum;
locular epidermis with numerous pores. I. Locular epidermis sculpted, with a raised, meridional ridge. Scale bar = 1 cm.
28
HENDERSON
fruits appear spiny (Fig. 8G). There is a well-developed locular epidermis present, and in some species this
has pores (Fig. 8H). There are two interesting features of the locular epidermis. In a few species there is a
basally situated operculum (a lid or cover above the embryo, Fig. 8H). In some species the locular epidermis
is elaborately sculpted, sometimes with a raised, meridional ridge (Fig. 8I). However, the degree of sculpting
of the locular epidermis ranges from scarcely sculpted without a ridge to elaborately sculpted with a ridge.
Sculpting is found in only 11 species, most of them species complexes. It is remarkable that such an elaborate
structure occurs in some specimens but not others within the same species. The seed is globose and the
endosperm is homogeneous. Eophylls are bifid and germination is adjacent.
Taxonomic Treatment
Analysis of the 44 qualitative variables divided them into 30 characters and 14 traits (Appendix I). Analysis of
the 30 characters divided the 4990 specimens into 68 species, 12 of them undescribed. Analysis of traits,
quantitative variables, and geography of these species divided 18 of them into 90 subspecies, giving a total of
140 taxa.
Geonoma Willdenow (1805: 174).
Lectotype (designated by Moore 1963): Geonoma simplicifrons Willd.
Gynestum Poiteau (1822: 387). Lectotype (designated by Moore 1963): Geonoma maxima (Poiteau) Kunth
Roebelia Engel (1865: 680). Type: Roebelia solitaria Engel
Kalbreyera Burret (1930a: 142). Type: Kalbreyera triandra Burret
Taenianthera Burret (1930a: 267). Type: Taenianthera macrostachys Burret
Stems solitary, or clustered, not cane-like, the internodes usually wider than long (ratio of stem diameter to
internode length greater than 2.2), or cane-like, the internodes usually longer than wide (ratio of stem diameter
to internode length less than 1.8); internodes yellowish and smooth, or, if short and congested, not scaly, or
covered with reddish or brownish scales, especially in their distal part, or covered with dense, brown scales.
Leaves undivided, or regularly pinnate, the pinnae with 1 main vein and 2 lateral veins on either side of main
vein, or irregularly pinnate, if regularly pinnate the pinnae with 1 main vein only (rarely with several lateral
veins), not plicate, or plicate, the bases of leaf blades running diagonally into the rachis, or recurved against
the rachis; petioles (and rachis) drying orange-brown or reddish-brown, or drying green or yellowish; veins
raised and rectangular in cross-section adaxially, or not raised or slightly raised and triangular in cross-section
adaxially. Inflorescences unbranched, or branched; prophylls and peduncular bracts ribbed with elongate,
unbranched fibers, both bracts tubular, narrow, elongate, closely sheathing the peduncle, more or less
persistent, or not ribbed with elongate, unbranched fibers, flattened (if tubular, narrow, and elongate then not
ribbed), deciduous or persistent; prophylls short, asymmetrically apiculate, the margins curved around the
stem, the surfaces flat with dense, felty, brown tomentum, prophyll equal to and early deciduous with the
peduncular bract, or not short and asymmetrically apiculate, the surfaces ridged with close, equal, parallel,
non-dividing ridges, scarcely tomentose between the ridges, or not ridged, or if ridged then densely tomentose
with widely to closely spaced ridges, these sometimes dividing, ridged, the ridges unequally wide, often
dividing from and rejoining other ridges, the prophyll margins with irregular, spine-like projections (rarely
these absent), the prophylls usually splitting irregularly between the ridges, or without unequally wide ridges;
peduncular bracts well-developed, or vestigial, the prophyll three times or more long, sometimes the
peduncular bract apparently well-developed but then soon disintegrating, or absent; rachillae surfaces with
spiky, fibrous projections or ridges, or without spiky, fibrous projections or ridges, drying brown or yellowbrown, the surfaces without short, transverse ridges, or drying brown, the surfaces with faint to pronounced,
short, transverse ridges, filiform with extended narrowed sections between the flower pits, or not filiform and
not or scarcely narrowed between the flower pits; flower pits tricussately or quadricussately arranged
29
throughout the rachillae, the groups of pits closely spaced, or usually spirally arranged, sometimes decussately
or tricussately, then the groups not closely spaced nor consistently arranged throughout the rachillae, or
decussately arranged throughout the rachillae, the groups of pits closely spaced, or alternately arranged
(sometimes distorted by twisting and contracting of rachillae), densely hairy internally distally only (rarely
some hairs on lateral margins of the pit), or glabrous internally, or densely hairy internally proximally and
distally; proximal and distal lips not joined laterally, with a clear gap between them, not forming a raised
cupule, the proximal lip margins usually overlapping the distal lip margins, or joined laterally with no clear
gap between them, often forming a raised cupule, the margins not overlapping, drying the same color as the
rachillae, or proximal and distal lips drying darker brown than the rachillae; proximal lips of flower pits with
a central notch before anthesis, often the two sides of the notch overlapping, the lips more or less heartshaped, or without a central notch before anthesis (but often tearing in the center after anthesis), not heartshaped, or apiculate and lobed before anthesis, not recurved at the apices after anthesis, or recurved at the
apices after anthesis, hood-shaped at anthesis (the margin of the proximal lip straight when viewed from
above), sometimes splitting post-anthesis, or not hood-shaped at anthesis; distal lips of flower pits absent, or
well-developed, or a scarcely raised rim; staminate and pistillate petals emergent, valvate throughout, or not
emergent, not valvate throughout; staminate flowers persistent post-anthesis, or deciduous post-anthesis;
stamens 3, or 6, or more than 6; thecae diverging at anthesis, inserted almost directly onto the filament apices,
the connectives bifid but scarcely developed, or diverging at anthesis, inserted onto bifid and well-developed,
non-jointed connectives, or diverging or not diverging at anthesis, inserted onto poorly to well-developed,
non-split, jointed connectives, connectives when well-developed alternately long and short, or diverging at
anthesis, inserted directly onto the apiculate filament apices; anthers short and curled over at anthesis, or not
short and curled at anthesis, usually elongate, spiraled and twisted or sometimes remaining straight, or short at
anthesis, remaining straight and parallel; gynoecium unilocular; non-fertilized pistillate flowers persistent
after anthesis, or deciduous after anthesis; staminodial tubes crenulate or shallowly lobed at the apex, or lobed
at the apex, the lobes spreading at anthesis, acuminate, or lobed at the apex, the lobes not spreading at
anthesis, not acuminate, those of non-fertilized pistillate flowers projecting and persistent after anthesis, or not
projecting, deciduous after anthesis; fruits bases with a prominent, asymmetric stipe, or without a prominent
stipe, ovoid, usually with conical apices, or not ovoid and without conical apices, the surfaces not splitting at
maturity, or splitting deeply and longitudinally at maturity to reveal mesocarp with dense layer of radial
fibers, with fibers emerging, or without fibers emerging, bumpy from the numerous, subepidermal, tangential,
short fibers present, these coming to a point at fruit apices, or not bumpy and not apiculate, or ridged from the
numerous, subepidermal, meridional, elongate fibers present, these coming to a point at fruit apices; locular
epidermis without operculum, or with operculum, smooth, or sculpted and then usually also with a raised,
meridional ridge, without pores or with very few pores, or with numerous pores.
Key to the species of Geonoma

1
2
3
4
5
6
Hispaniola, Lesser Antilles, Trinidad and Tobago ....................................................................................................... 2

Central and South America .......................................................................................................................................... 3
Distal lips of flower pits absent; flower pits densely hairy internally .................................................... G. pinnatifrons
Distal lips of flower pits well-developed; flower pits not densely hairy internally.........................................G. undata
Central America (Mexico, Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Panama) ...................................... 4
South America ............................................................................................................................................................ 24
Distal lips of flower pits absent; flower pits densely hairy internally distally ............................................................. 5
Distal lips of flower pits well-developed; flower pits not densely hairy internally..................................................... 6
Flower pits densely hairy internally distally only (rarely some hairs on lateral margins of the pit); pinnae 18(447)
per side of rachis; rachillae 71(22120).................................................................................................... G. interrupta
Flower pits densely hairy internally proximally and distally; pinnae 8(239) per side of rachis; rachillae 18(445) ...
............................................................................................................................................................... G. pinnatifrons
Flower pits tricussately arranged throughout the rachillae, the groups of pits closely spaced ..................... G. deversa
30
HENDERSON
7
8
9
10
11
12
13
-
14
15
16
17
18
19
20
21
22
23
24
-
Flower pits spirally, alternately, or rarely decussately arranged................................................................................... 7

Flower pits decussately arranged; peduncular bracts vestigial; stamens 3; eastern Panama ....................... G. triandra
Flower pits spirally or alternately arranged; peduncular bracts well-developed, rarely absent; stamens 6 or more;
Mexico, Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Panama ................................................................... 8
Prophyll surfaces ridged with close, equal, parallel, non-dividing ridges, scarcely tomentose between the ridges; locular epidermis with operculum ..................................................................................................................................... 9
Prophyll surfaces not ridged, or if ridged then densely tomentose with widely to closely spaced ridges, these sometimes dividing; locular epidermis without operculum ................................................................................................ 11
Rachillae 1.4(0.81.9) mm in diameter; staminodial tubes of non-fertilized pistillate flowers not projecting, deciduous after anthesis; Panama ................................................................................................................... G. concinnoidea
Rachillae 5.1(4.18.3) mm in diameter; staminodial tubes of non-fertilized pistillate flowers projecting and persistent after anthesis ........................................................................................................................................................ 10
Proximal lips of flower pits with a central notch before anthesis, often the two sides of the notch overlapping, the
lips more or less heart-shaped; Honduras, Nicaragua, Costa Rica, western and central Panama................ G. congesta
Proximal lips of flower pits without a central notch before anthesis (but often tearing in the center after anthesis), not
heart-shaped; eastern Panama .......................................................................................................G. calyptrogynoidea
Rachillae filiform with extended narrowed sections between the flower pits; rachillae 110(85138) ...................... 12
Rachillae not filiform and not or scarcely narrowed between the flower pits; rachillae 13(138) ............................ 13
Internodes covered with dense, brown scales; pinnae 3 per side of rachis; Costa Rica on the Osa Peninsula and adjacent areas...................................................................................................................................................... G. scoparia
Internodes yellowish and smooth; pinnae 33(2649) per side of rachis; western Panama ........................G. mooreana
Staminodial tubes lobed at the apex, the lobes spreading at anthesis, acuminate; stamens more than 6; Panama ........
................................................................................................................................................................... G. deneversii
Staminodial tubes lobed at the apex, the lobes not spreading at anthesis, not acuminate, or staminodial tubes crenulate or shallowly lobed at the apex; stamens 6; Mexico, Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Panama ............................................................................................................................................................................. 14
Prophylls short, asymmetrically apiculate, the margins curved around the stem, the surfaces flat with dense, felty,
brown tomentum, prophyll equal to and early deciduous with the peduncular bract ................................................ 15
Prophylls not short and asymmetrically apiculate ...................................................................................................... 16
Rachillae surfaces with faint to pronounced, short, transverse ridges; rachillae 22.9(10.042.0) cm long ...................
............................................................................................................................................................. G. longivaginata
Rachillae surfaces without short, transverse ridges; rachillae 8.1(3.815.0) cm long .............................G. ferruginea
Proximal lips of flower pits apiculate and lobed before anthesis, tearing in the center after anthesis; fruit bases with
a prominent, asymmetric stipe .................................................................................................................................... 17
lips more or less heart-shaped; fruit bases without a prominent stipe ....................................................................... 20
Peduncular bracts absent; inflorescences unbranched; Costa Rica and western Panama......................G. talamancana
Peduncular bracts well-developed; inflorescences branched or unbranched; Mexico, Guatemala, Belize, Honduras,
Nicaragua, Costa Rica, Panama .................................................................................................................................. 18
Inflorescences branched; prophyll surfaces ridged, the ridges unequally wide, often dividing from and rejoining
other ridges, the prophyll margins with irregular, spine-like projections (rarely these absent), the prophylls usually
splitting irregularly between the ridges ...........................................................................................................G. undata
Inflorescences branched or unbranched; prophyll surfaces without unequally wide ridges ...................................... 19
Prophylls and peduncular bracts ribbed with elongate, unbranched fibers, both bracts tubular, narrow, elongate,
closely sheathing the peduncle, more or less persistent; inflorescences unbranched; western Panama G. lehmannii
Prophylls and peduncular bracts not ribbed with elongate, unbranched fibers, flattened, deciduous or persistent;
inflorescences branched, rarely unbranched; Nicaragua, Costa Rica, and Panama................................ G. orbignyana
Staminodial tubes crenulate or shallowly lobed at the apex .......................................................................... G. cuneata
Staminodial tubes lobed at the apex, the lobes not spreading at anthesis, not acuminate .......................................... 21
Peduncular bracts vestigial ........................................................................................................................................ 22
Peduncular bracts well-developed .............................................................................................................................. 23
Peduncles 5.4(3.87.0) cm long; rachillae 2(13); Costa Rica and Panama .......................................... G. monospatha
Peduncles 24.3(15.037.5) cm long; rachillae 1; Costa Rica ........................................................................ G. brenesii
Petioles absent; Costa Rica and Panama ....................................................................................................G. epetiolata
Petioles 7.0(1.012.0) cm long; Panama ...................................................................................................... G. hugonis
Western Andean slopes and Pacific lowlands of Colombia and Ecuador, including Cauca and Magdalena valleys in
Colombia .................................................................................................................................................................... 25
Andes of Colombia (including the Sierra Nevada de Santa Marta), Venezuela (including the Coastal Cordillera and
the Pennsula de Paria), Ecuador, Peru, and Bolivia, Amazon region of Venezuela, Colombia, Ecuador, Peru,
31
25
26
27
28
29
30
31
32
33
34
35
36
37
-
38
39
40
-
Bolivia, Brazil, and the Guianas, including the Guayana Highland, Cerrado of central Brazil and adjacent Paraguay
and Bolivia, and the Atlantic Coastal Forest of Brazil .............................................................................................. 44
Distal lips of flower pits absent; flower pits densely hairy internally distally ........................................................... 26
Distal lips of flower pits well-developed; flower pits not densely hairy internally.................................................... 27
Flower pits densely hairy internally distally only (rarely some hairs on lateral margins of the pit); pinnae 18(447)
per side of rachis; rachillae 71(22120)..................................................................................................... G. interrupta
Flower pits densely hairy internally proximally and distally; pinnae 8(239) per side of rachis; rachillae 18(445) ...
............................................................................................................................................................... G. pinnatifrons
Flower pits tricussately arranged throughout the rachillae, the groups of pits closely spaced...................... G. deversa
Flower pits spirally, alternately, or rarely decussately arranged................................................................................. 28
Flower pits decussately arranged; peduncular bracts vestigial; stamens 3................................................... G. triandra
Flower pits spirally or alternately arranged; peduncular bracts well-developed, rarely vestigial; stamens 6 ............ 29
Prophyll surfaces ridged with close, equal, parallel, non-dividing ridges, scarcely tomentose between the ridges; locular epidermis with operculum ................................................................................................................................... 30
Prophyll surfaces not ridged, or if ridged then densely tomentose with widely to closely spaced ridges, these sometimes dividing; locular epidermis without operculum ................................................................................................ 31
Fruit surfaces without fibers emerging; staminodial tubes of non-fertilized pistillate flowers not projecting, deciduous after anthesis; rachillae 1.7(1.22.7) mm in diameter; Colombia (Valle) on the Western Cordillera ......................
.................................................................................................................................................................... G. concinna
Fruit surfaces with fibers emerging; staminodial tubes of non-fertilized pistillate flowers projecting and persistent
after anthesis; rachillae 6.1(4.58.3) mm in diameter; Pacific coast of Colombia and Ecuador, and the Magdalena
and Cauca valleys in Colombia.......................................................................................................G. calyptrogynoidea
Rachillae filiform with extended narrowed sections between the flower pits; western Ecuador .............................. 32
Rachillae not filiform and not or scarcely narrowed between the flower pits; western Andean slopes and Pacific lowlands of Colombia and Ecuador, including Cauca and Magdalena valleys in Colombia ........................................... 33
Fruits 5.3(4.66.0) cm long, 4.8(4.15.5) mm in diameter; veins not raised or slightly raised and triangular in crosssection adaxially; Montaas de Ila at 597(520700) m ............................................................................G. tenuissima
Fruits 8.1 mm long, 7.2 mm in diameter veins raised and rectangular in cross-section adaxially; Reserva Biolgica
Los Cedros at 1465(14601470) m elevation .................................................................................................G. venosa
Staminodial tubes lobed at the apex, the lobes spreading at anthesis, acuminate ...................................................... 34
Staminodial tubes staminodial tubes crenulate or shallowly lobed at the apex, rarely lobed at the apex, the lobes not
spreading at anthesis, not acuminate .......................................................................................................................... 36
Rachillae 21(1236); operculum absent ........................................................................................................ G. maxima
Rachillae 1(14); operculum present ......................................................................................................................... 35
Fruit surfaces splitting deeply and longitudinally at maturity to reveal mesocarp with dense layer of radial fibers;
distal lip of flower pit well developed; inflorescence branched or unbranched; rachis 111.7(80.0137.0) cm long .....
................................................................................................................................................................... G. chococola
Fruit surfaces not splitting at maturity; distal lip of flower pit a scarcely raised rim; inflorescences unbranched;
rachis 36.6(21.563.0) cm long ................................................................................................................. G. paradoxa
Prophylls short, asymmetrically apiculate, the margins curved around the stem, the surfaces flat with dense, felty,
brown tomentum, prophyll equal to and early deciduous with the peduncular bract; Pacific coast of northwestern
Colombia .......................................................................................................................................................... G. divisa
Prophylls not short and asymmetrically apiculate; Pacific coast of Colombia and Ecuador, and the Magdalena and
Cauca valleys in Colombia ......................................................................................................................................... 37
a prominent, asymmetric stipe; higher elevations at >1150 m on western Andean slopes ....................................... 38
lips more or less heart-shaped; fruit bases without a prominent stipe; lower elevations at <1375 m on the Pacific
coast of Colombia and Ecuador, and the Magdalena and Cauca valleys in Colombia ............................................... 40
Inflorescences branched; prophyll surfaces ridged, the ridges unequally wide, often dividing from and rejoining
other ridges, the prophyll margins with irregular, spine-like projections (rarely these absent), the prophylls usually
splitting irregularly between the ridges ...........................................................................................................G. undata
Inflorescences unbranched; prophyll surfaces without unequally wide ridges ......................................................... 39
closely sheathing the peduncle, more or less persistent; inflorescences unbranched ................................G. lehmannii
inflorescences branched, rarely unbranched ............................................................................................ G. orbignyana
Inflorescences unbranched.......................................................................................................................................... 41
Inflorescences branched ............................................................................................................................................. 42
32
HENDERSON
41. Peduncles 51.3(13.7117) cm long; Western Andean slopes and Pacific lowlands of Colombia and Ecuador, including Cauca and Magdalena valleys in Colombia ............................................................................................ G. cuneata
Peduncles 5.0(0.517) cm long; Pacific coast of Colombia (Choc).............................................................. G. stricta
42 Veins raised and rectangular in cross-section adaxially; rachillae 4(25); Ecuador on western Andean slopes at
746(2001800) m elevation ............................................................................................................................ G. lanata
Veins not raised or slightly raised and triangular in cross-section adaxially; rachillae 15(1218); Pacific coast of
Colombia at 100150 m elevation .............................................................................................................................. 43
43 Peduncle 5.9 cm long; rachillae 18; Valle..................................................................................................G. dindoensis
- Peduncle 18.7 cm long; rachillae 12; Choc ................................................................................................. G. gentryi
44 Andes of Colombia (including the Sierra Nevada de Santa Marta) and Venezuela (including the Coastal Cordillera
and the Pennsula de Paria) ......................................................................................................................................... 45
Andes of Ecuador, Peru, and Bolivia, Amazon region of Venezuela, Colombia, Ecuador, Peru, Bolivia, Brazil, and
the Guianas, including the Guayana Highland, Cerrado of central Brazil and adjacent Paraguay and Bolivia, and the
Atlantic Coastal Forest of Brazil................................................................................................................................. 63
45 Distal lips of flower pits absent; flower pits densely hairy internally distally .......................................................... 46
Distal lips of flower pits well-developed; flower pits not densely hairy internally.................................................... 50
46 Flower pits densely hairy internally distally only (rarely some hairs on lateral margins of the pit) ......................... 47
Flower pits densely hairy internally proximally and distally ..................................................................................... 48
47 Rachis 129.6(55.5200) cm long; pinnae 18(447) per side of rachis; rachillae 71(22120); Andes of Colombia and
Venezuela .................................................................................................................................................. G. interrupta
Rachis 35.4(24.045.0) cm long; pinnae 2(14) per side of rachis; rachillae 3(25); Coastal Cordillera in Venezuela
............................................................................................................................................................... G. simplicifrons
48 Rachillae surfaces without short, transverse ridges; Western, Central, and Eastern Cordilleras in Colombia...............
................................................................................................................................................................. G. frontinensis
Rachillae surfaces with faint to pronounced, short, transverse ridges; Andes of Colombia (including the Sierra
Nevada de Santa Marta) and Venezuela (including the Coastal Cordillera and the Pennsula de Paria).................... 49
49 Rachis 89.1(36.0163.0) cm long; pinnae 8(239) per side of rachis; prophyll surfaces ridged, the ridges unequally
wide, often dividing from and rejoining other ridges, the prophyll margins with irregular, spine-like projections
(rarely these absent), the prophylls usually splitting irregularly between the ridges; Andes of Colombia (including
the Sierra Nevada de Santa Marta) and Venezuela (including the Coastal Cordillera and the Pennsula de Paria) ......
................................................................................................................................................................ G. pinnatifrons
Rachis 45.4(22.085.0) cm long; pinnae 3(16) per side of rachis; prophyll surfaces without unequally wide ridges;
eastern Andean slopes in Colombia ............................................................................................................. G. euspatha
50 Flower pits tricussately arranged throughout the rachillae, the groups of pits closely spaced...................... G. deversa
Flower pits spirally, alternately, or rarely decussately arranged ................................................................................ 51
51 Prophyll surfaces ridged with close, equal, parallel, non-dividing ridges, scarcely tomentose between the ridges; locular epidermis with operculum; Colombia (Antioquia) in the Central Cordillera ...................................... G. concinna
- Prophyll surfaces not ridged, or if ridged then densely tomentose with widely to closely spaced ridges, these sometimes dividing; locular epidermis without operculum; Andes of Colombia (including the Sierra Nevada de Santa
Marta) and Venezuela (including the Coastal Cordillera and the Pennsula de Paria) ............................................... 52
52 Rachillae filiform with extended narrowed sections between the flower pits........................................................... 53
Rachillae not filiform and not or scarcely narrowed between the flower pits............................................................ 55
53 Fruit surfaces not bumpy and not apiculate; Central Cordillera in Colombia (Antioquia) ........................... G. bernalii
Fruit surfaces bumpy from the numerous, subepidermal, tangential, short fibers present, these coming to a point at
fruit apices; Coastal Cordillera of Venezuela ............................................................................................................. 54
54 Operculum absent; Yaracuy ...........................................................................................................................G. braunii
Operculum present; Miranda ................................................................................................................... G. operculata
55 Staminodial tubes lobed at the apex, the lobes spreading at anthesis, acuminate; Central and Eastern Cordillera in
Colombia (Antioquia) ..................................................................................................................... G. chlamydostachys
Staminodial tubes staminodial tubes crenulate or shallowly lobed at the apex, rarely lobed at the apex, the lobes not
spreading at anthesis, not acuminate; Andes of Colombia (including the Sierra Nevada de Santa Marta) and Venezuela (including the Coastal Cordillera and the Pennsula de Paria).............................................................................. 56
56 Proximal lips of flower pits apiculate and lobed before anthesis, tearing in the center after anthesis; fruit bases with
a prominent, asymmetric stipe .................................................................................................................................... 57
lips more or less heart-shaped; fruit bases without a prominent stipe; elevation and area ........................................ 59
57 Prophyll surfaces ridged, the ridges unequally wide, often dividing from and rejoining other ridges, the prophyll
margins with irregular, spine-like projections (rarely these absent), the prophylls usually splitting irregularly
between the ridges; inflorescences branched ..................................................................................................G. undata
33
Prophyll surfaces without unequally wide ridges; inflorescences unbranched or branched ..................................... 58
58 Prophylls and peduncular bracts ribbed with elongate, unbranched fibers, both bracts tubular, narrow, elongate,
closely sheathing the peduncle, more or less persistent; inflorescences unbranched .................................G. lehmannii
inflorescences branched, rarely unbranched ........................................................................................... G. orbignyana
59 Inflorescences unbranched......................................................................................................................................... 60
Inflorescences branched ............................................................................................................................................. 62
60 Staminodial tubes lobed at the apex, the lobes not spreading at anthesis, not acuminate; fruit surfaces bumpy from
the numerous, subepidermal, tangential, short fibers present, these coming to a point at fruit apices Eastern Cordillera in Colombia .................................................................................................................................G. santanderensis
Staminodial tubes crenulate or shallowly lobed at the apex; fruit surfaces ridged from the numerous, subepidermal,
meridional, elongate fibers present, these coming to a point at fruit apices; Central Cordillera in Colombia and foothills of Andes and Sierra Nevada de Santa Marta ..................................................................................................... 61
61. Peduncles 51.3(13.7117) cm long; foothills of Andes and Sierra Nevada de Santa Marta......................... G. cuneata
Peduncles 5.0(0.517) cm long; Central Cordillera in Colombia (Antioquia) ............................................... G. stricta
62 Proximal and distal lips not joined to form a raised cupule; Coastal Cordillera in Venezuela (Aragua, Carabobo)......
.................................................................................................................................................................. G. spinescens
Proximal and distal lips not joined to form a raised cupule; Central Cordillera in Colombia................... G. galeanoae
63 Andes of Ecuador, Peru, and Bolivia and Amazon region of Venezuela, Colombia, Ecuador, Peru, Bolivia, Brazil,
and the Guianas, including the Guayana Highland .................................................................................................... 64
Cerrado of central Brazil and adjacent Paraguay and Bolivia, and the Atlantic Coastal Forest of Brazil ................. 94
64 Distal lips of flower pits absent; flower pits densely hairy internally distally ........................................................... 65
Distal lips of flower pits present; flower pits not densely hairy internally................................................................. 67
65 Flower pits densely hairy internally distally only (rarely some hairs on lateral margins of the pit); rachis 129.6(55.5
200) cm long; pinnae 18(447) per side of rachis; rachillae 71(22120)................................................. G. interrupta
Flower pits densely hairy internally proximally and distally; rachis 67.9(22.0163.0) cm long; pinnae 7(139) per
side of rachis; rachillae 14(245)................................................................................................................................ 66
66 Rachis 89.1(36.0163.0) cm long; pinnae 9(239) per side of rachis; prophyll surfaces ridged, the ridges unequally
wide, often dividing from and rejoining other ridges, the prophyll margins with irregular, spine-like projections
(rarely these absent), the prophylls usually splitting irregularly between the ridges; eastern Andean slopes in Ecuador .......................................................................................................................................................... G. pinnatifrons
Rachis 45.4(22.085.0) cm long; pinnae 3(16) per side of rachis; prophyll surfaces without unequally wide ridges;
eastern Andean slopes in Ecuador, Peru, and Bolivia, the Guayana Highland region and outlying montane areas in
Venezuela, Brazil, Guyana, Suriname, and French Guiana, and just reaching the Amazon region of Brazil (Par,
Rndonia) .................................................................................................................................................... G. euspatha
67 Flower pits tricussately arranged throughout the rachillae, the groups of pits closely spaced.................................. 68
Flower pits spirally or alternately arranged, rarely decussately arranged, or rarely tricussately arranged and then
loosely spaced ............................................................................................................................................................ 69
68 Peduncular bracts well-developed; stamens 6; Andes of Ecuador, Peru, and Bolivia and Amazon region of Venezuela, Colombia, Ecuador, Peru, Bolivia, Brazil, and the Guianas, including the Guayana Highland ............. G. deversa
Peduncular bracts vestigial; stamens 3; eastern Andean slopes in Ecuador ............................................. G. hollinensis
69 Prophyll surfaces ridged with close, equal, parallel, non-dividing ridges, scarcely tomentose between the ridges; locular epidermis with operculum; central and northeastern Amazon region of Brazil, the Guianas, and Venezuela .......
................................................................................................................................................................... G. baculifera
- Prophyll surfaces not ridged, or if ridged then densely tomentose with widely to closely spaced ridges, these sometimes dividing; locular epidermis without operculum; Andes of Ecuador, Peru, and Bolivia and Amazon region of
Venezuela, Colombia, Ecuador, Peru, Bolivia, Brazil, and the Guianas, including the Guayana Highland ............. 70
70 Rachillae filiform with extended narrowed sections between the flower pits; eastern Andean slopes in Ecuador........
........................................................................................................................................................................ G. fosteri
Rachillae not filiform and not or scarcely narrowed between the flower pits; Andes of Ecuador, Peru, and Bolivia
and Amazon region of Venezuela, Colombia, Ecuador, Peru, Bolivia, Brazil, and the Guianas, including the Guayana
Highland ..................................................................................................................................................................... 71
71 Staminodial tubes lobed at the apex, the lobes spreading at anthesis, acuminate 72 Staminodial tubes staminodial
tubes crenulate or shallowly lobed at the apex, rarely lobed at the apex, the lobes not spreading at anthesis, not acuminate ............................................................................................................................................................................ 80
72 Peduncles 8.2(3.219.5) cm long; rachillae 19(450); operculum absent.................................................... G. maxima
Peduncles 55.3(19.2143.0) cm long; rachillae 1(19); operculum present .............................................................. 73
73 Stamens more than 6; western Amazon region of Colombia and Ecuador ............................................... G. multisecta
Stamens 6; Andes of Ecuador, Peru, and Bolivia and Amazon region of Venezuela, Colombia, Ecuador, Peru,
34
HENDERSON
74
75
76
77
78
-
79
80
81
82
83
84
85
86
87
88
89
90
Bolivia, Brazil, and the Guianas, including the Guayana Highland. .......................................................................... 74
Bases of blades recurved against the rachis .............................................................................................................. 75
Bases of leaf blades running diagonally into the rachis.............................................................................................. 77
Rachilla 1, 32.7(26.045.0) cm long; eastern Amazon region in Guyana, French Guiana, and Brazil ....G. oldemanii
Rachillae 5(39), 17.6(7.533.5) cm long; eastern and western Amazon region....................................................... 76
Fruit surfaces not splitting; western Amazon region in Colombia, Ecuador, and Peru .............................G. triglochin
eastern Amazon region in Guyana, French Guiana, and Brazil ...................................................... G. umbraculiformis
Peru (Amazonas) ................................................................................................................................... G. schizocarpa
Fruits surfaces not splitting......................................................................................................................................... 78
Thecae diverging at anthesis, inserted directly onto the apiculate filament apices; anthers not short and curled at
anthesis, usually elongate, spiraled and twisted or sometimes remaining straight ........................................ G. camana
Thecae diverging or not diverging at anthesis, inserted onto poorly to well-developed, non-split, jointed connectives,
connectives when well-developed alternately long and short; anthers short at anthesis, remaining straight and parallel ............................................................................................................................................................................... 79
fruit apices; western Amazon region in Venezuela, Colombia, Ecuador, Peru, and Brazil .................G. macrostachys
Fruit surfaces not bumpy and not apiculate; eastern and central Amazon region of the Guianas, Venezuela, Colombia, and Brazil ..........................................................................................................................................G. poiteauana
a prominent, asymmetric stipe ................................................................................................................................... 81
lips more or less heart-shaped; fruit bases without a prominent stipe; elevation and area ........................................ 84
Prophyll surfaces ridged, the ridges unequally wide, often dividing from and rejoining other ridges, the prophyll
between the ridges; inflorescences branched ..................................................................................................G. undata
Prophyll surfaces without unequally wide ridges; inflorescences unbranched or branched .................................... 82
Distal lips of flower pits absent; eastern Andean slopes in Ecuador and Peru...............................................G. trigona
Distal lips present (rarely absent); Andes of Ecuador, Peru, and Bolivia................................................................... 83
closely sheathing the peduncle, more or less persistent; inflorescences unbranched ................................G. lehmannii
inflorescences branched, rarely unbranched ........................................................................................... G. orbignyana
Rachillae surfaces with spiky, fibrous projections or ridges ...................................................................................... 85
Rachillae surfaces without spiky, fibrous projections or ridges ................................................................................. 86
Peduncular bracts vestigial, the prophyll three times or more long, sometimes the peduncular bract apparently welldeveloped but then soon disintegrating; internodes yellowish and smooth, or, if short and congested, not scaly .........
........................................................................................................................................................................ G. stricta
Peduncular bracts well-developed; internodes covered with reddish or brownish scales, especially in their distal part
.................................................................................................................................................................................... 86
Fruit surfaces not bumpy and not apiculate; Guyana, and the central Amazon region in Brazil ............ G. aspidiifolia
fruit apices; central-western Amazon region of Colombia, Venezuela, and Brazil ..................................G. oligoclona
Distal lips of flower pits a scarcely raised rim; prophyll 20.1(5.740) cm long; rachillae 3(110)........................... 88
Distal lips of flower pits well-developed; prophyll 7.2(2.311.2) cm long; rachillae 10(233) ................................ 91
Rachillae surfaces with faint to pronounced, short, transverse ridges; western Amazon and sub-Andean regions of
Colombia, Ecuador, and Peru .........................................................................................................G. longipedunculata
Rachillae surfaces without short, transverse ridges; widespread ............................................................................... 89
Prophylls and peduncular bracts not ribbed with elongate, unbranched fibers, flattened (if tubular, narrow, and elongate then not ribbed), deciduous or persistent ........................................................................................G. poeppigiana
closely sheathing the peduncle, more or less persistent ............................................................................................. 90
lips more or less heart-shaped; fruit surfaces not bumpy and not apiculate western Amazon and sub-Andean regions
of Ecuador, Peru, Bolivia, and Brazil, with an outlier in Venezuela...................................................... G. brongniartii
Proximal lips of flower pits without a central notch before anthesis (but often tearing in the center after anthesis), not
heart-shaped; fruit surfaces bumpy from the numerous, subepidermal, tangential, short fibers present, these coming
to a point at fruit apices Peru (San Martn)........................................................................................ G. sanmartinensis
35
91 Internodes covered with dense, brown scales; eastern Andean slopes in Peru.......................................... G. peruviana
Internodes yellowish and smooth; widespread ........................................................................................................... 92
92 Rachis 43.8(35.057.5) cm long; pinnae 4(27); rachillae 24(1333); fruit surfaces bumpy from the numerous, subepidermal, tangential, short fibers present, these coming to a point at fruit apices ............................... G. occidentalis
Rachis 26.5(11.543.0) cm long; pinnae 1(13) per side of rachis; rachillae 5(212); fruit surfaces not bumpy and
not apiculate .............................................................................................................................................................. 93
93 Prophylls short, asymmetrically apiculate, the margins curved around the stem, the surfaces flat with dense, felty,
brown tomentum, prophyll equal to and early deciduous with the peduncular bract; western Amazon region of
Colombia, Brazil, Ecuador, Peru, and Bolivia ..............................................................................................G. laxiflora
Prophylls not short and asymmetrically apiculate; Amazon region of Colombia, Venezuela, Guyana, Suriname,
French Guiana, Brazil, Peru, Ecuador, and Bolivia ................................................................................ G. leptospadix
94 Leaf blades regularly pinnate, the pinnae with 1 main vein and two lateral veins ................................... G. schottiana
Leaf blades undivided or irregularly pinnate, if regularly pinnate the pinnae with 1 main vein only (rarely with several lateral veins)......................................................................................................................................................... 95
95 Prophylls and peduncular bracts ribbed with elongate, unbranched fibers, both bracts tubular, narrow, elongate,
closely sheathing the peduncle, more or less persistent; Atlantic Coastal Forest of Brazil from southern Bahia and
Minas Gerais to Santa Catarina ..................................................................................................................... G. elegans
Prophylls and peduncular bracts not ribbed with elongate, unbranched fibers, flattened (if tubular, narrow, and elongate then not ribbed), deciduous or persistent; Cerrado of central Brazil and adjacent Paraguay and Bolivia, and the
Atlantic Coastal Forest of Brazil................................................................................................................................. 96
96 Veins raised and rectangular in cross-section adaxially; prophyll surfaces ridged, the ridges unequally wide, often
dividing from and rejoining other ridges, the prophyll margins with irregular, spine-like projections (rarely these
absent), the prophylls usually splitting irregularly between the ridges; Cerrado of central Brazil and adjacent Paraguay and Bolivia, and the Atlantic Coastal Forest of Brazil ...................................................................... G. pohliana
Veins not raised or slightly raised and triangular in cross-section adaxially; prophyll surfaces without unequally
wide ridges; Atlantic Coastal Forest of Brazil from Pernambuco to Bahia .............................................. G. pauciflora
1. Geonoma aspidiifolia Spruce (1871: 112). Type: BRAZIL. Amazonas: Tarum, February 1855, R. Spruce
75 (holotype K!, isotype NY!).
Plants 1.8(1.03.0) m tall; stems 1.8(1.03.0) m tall, 0.8(0.51.6) cm in diameter, solitary or clustered, canelike; internodes 2.7(1.17.8) cm long, covered with reddish or brownish scales, especially in their distal part.
Leaves 9(612) per stem, irregularly pinnate, not plicate, bases of blades running diagonally into the rachis;
sheaths 9.5(8.511.5) cm long; petioles 20.6(8.029.0) cm long, drying green or yellowish; rachis 19.5(12.0
38.0) cm long, 2.3(1.33.5) mm in diameter; veins raised and rectangular in cross-section adaxially; pinnae
3(24) per side of rachis; basal pinna 13.7(8.225.3) cm long, 1.5(0.53.8) cm wide, forming an angle of
74(4590) with the rachis; apical pinna 13.9(9.019.8) cm long, 8.7(4.513.8) cm wide, forming an angle of
35(2543) with the rachis. Inflorescences branched 1 order; prophylls and peduncular bracts not ribbed with
elongate, unbranched fibers, flattened, persistent; prophylls 5.7(3.58.8) cm long, not short and
asymmetrically apiculate, the surfaces not ridged, without unequally wide ridges; peduncular bracts 4.9(3.2
8.2) cm long, well-developed, inserted 0.5(0.20.8) cm above the prophyll; peduncles 5.1(3.48.2) cm long,
3.5(2.24.8) mm in diameter; rachillae 3(24), 7.3(4.016.0) cm long, 3.4(2.34.9) mm in diameter, the
surfaces with spiky, fibrous projections or ridges, drying brown or yellow-brown, without short, transverse
ridges, not filiform and not narrowed between the flower pits; flower pits spirally arranged, glabrous
internally; proximal lips without a central notch before anthesis, not recurved after anthesis, not hood-shaped;
proximal and distal lips drying the same color as the rachillae, not joined to form a raised cupule, the proximal
lip margins overlapping the distal lip margins; distal lips well-developed; staminate and pistillate petals not
emergent, not valvate throughout; staminate flowers deciduous after anthesis; stamens 6; thecae diverging at
anthesis, inserted onto bifid and well-developed, non-jointed connectives; anthers short and curled over at
anthesis; non-fertilized pistillate flowers deciduous after anthesis; staminodial tubes lobed at the apex, the
lobes not spreading at anthesis, not acuminate, those of non-fertilized flowers not projecting and persistent
after anthesis; fruits 9.5(7.911.2) mm long, 6.7(6.07.8) mm in diameter, the bases without a prominent
36
HENDERSON
stipe, the apices not conical, the surfaces not splitting at maturity, without fibers emerging, not bumpy, not
apiculate; locular epidermis without operculum, smooth, without pores.
Taxonomic notes:Geonoma aspidiifolia is a member of the G. stricta clade and is closely related to two
other speciesG. oligoclona and G. santanderensis. It differs from these two in its fruits surfaces which are
not bumpy. Wessels Boer (1968) misunderstood G. aspidiifolia, as shown by Henderson (1995). However,
both Henderson (1995) and Henderson et al. (1995) included G. fusca as a synonym of G. aspidiifolia. As
explained below, the two are here recognized at the subspecific level.
Subspecific variation:One trait (stem branching) varies within this species. There is geographic
discontinuity and two subgroups can be recognizedGuyana, and the central Amazon region in Brazil.
Specimens from Guyana differ significantly from Brazilian specimens in eight variables (stem diameter,
rachis length, basal pinna length, prophyll length, peduncular bract length, peduncle length, peduncle width,
and fruit diameter)(t-test, P <0.05). Based on these results, and geographic discontinuity, the two subgroups
are recognized as subspecies (subspp. aspidiifolia, fusca).
Key to the subspecies of G. aspidiifolia
1
-
Peduncles 4.3(3.46.0) cm long; peduncular bracts 4.0(3.24.7) cm long; central Amazon region of Brazil ..............
...........................................................................................................................................................subsp. aspidiifolia
Peduncles 6.1(4.58.2) cm long; peduncular bracts 6.6(4.58.2) cm long; Guyana.................................. subsp. fusca
1a. Geonoma aspidiifolia subsp. aspidiifolia

Inflorescences peduncles 4.3(3.46.0) cm long; peduncular bracts 4.0(3.24.7) cm long.
Distribution and habitat:From 200308S and 59436040W in the central Amazon region of
Brazil (Amazonas) at low elevations in non-flooded lowland rainforest (Fig. 9).
1b. Geonoma aspidiifolia subsp. fusca (Wessels Boer) Henderson, comb. & stat. nov.
Basionym: Geonoma fusca Wessels Boer (1972: 93). Type: GUYANA. Upper Mazaruni River basin, Mt. Ayanganna,
700800 m, 5 August 1960, S. & C. Tillett 45047 (holotype NY!).
Inflorescences peduncles 6.1(4.58.2) cm long; peduncular bracts 6.6(4.58.2) cm long.

Distribution and habitat:From 458526N and 59066002W in the Pakaraima mountains of
Guyana at 870(1021350) m elevation in lowland to montane rainforest (Fig. 9).
There are three populations of this subspeciesan eastern, low elevation one at 282(102442) m
elevation, and two higher elevation ones, a northern one on Mount Ayanganna at 1160(10501350) m
elevation and a southern one on Mount Wokumung at 872(6861120) m elevation. There appear to be some
differences between these three populations, particularly the larger leaves and inflorescences of the low
elevation population, although there are too few specimens to test for differences. However, these separate
populations may be an artifact of insufficient collecting.
2. Geonoma baculifera (Poiteau) Kunth (1841: 233). Gynestum baculiferum Poiteau (1822: 389). Type:
FRENCH GUIANA. Without locality, no date, A. Poiteau s. n. (holotype, P!).
Geonoma acutiflora Martius (1823: 10). Lectotype (designated by Wessels Boer 1968): BRAZIL. Par: without locality,
no date, C. Martius s.n. (lectotype M!).
Geonoma macrospatha Spruce (1871: 105). Geonoma baculifera var. macrospatha (Spruce) Drude (1882: 490). Type:
VENEZUELA. Amazonas: Ro Casiquiare, December 1853, R. Spruce 42 (holotype K!, isotype P!).
Plants 2.3(0.56.0) m tall; stems 1.6(1.02.5) m tall, 1.6(1.32.3) cm in diameter, solitary or clustered, canelike; internodes 4.1(1.49.3) cm long, yellowish and smooth. Leaves 9(611) per stem, undivided or
37
irregularly pinnate, not plicate, bases of blades running diagonally into the rachis; sheaths 17.7(11.030.0) cm
long; petioles 18.0(6.530.0) cm long, drying green or yellowish; rachis 56.2(39.580.0) cm long, 4.2(2.6
7.2) mm in diameter; veins raised and rectangular in cross-section adaxially; pinnae 2(110) per side of
rachis; basal pinna 55.4(42.076.0) cm long, 17.2(6.733.0) cm wide, forming an angle of 20(1228) with
the rachis; apical pinna 28.8(14.039.5) cm long, 19.512.540.0) cm wide, forming an angle of 25(2035)
with the rachis. Inflorescences branched 12 orders; prophylls and peduncular bracts not ribbed with elongate,
unbranched fibers, flattened, persistent; prophylls 24.2(12.733.0) cm long, not short and asymmetrically
apiculate, the surfaces ridged with close, equal, parallel, non-dividing ridges, scarcely tomentose between the
ridges, without unequally wide ridges; peduncular bracts 23.6(13.531.0) cm long, well-developed, inserted
2.8(1.27.0) cm above the prophyll; peduncles 29.1(13.344.2) cm long, 4.0(2.18.4) mm in diameter;
rachillae 6(311), 16.7(4.031.0) cm long, 3.2(2.24.3) mm in diameter, the surfaces without spiky, fibrous
projections or ridges, drying brown, with faint to pronounced, short, transverse ridges, not filiform and not
narrowed between the flower pits; flower pits spirally arranged, glabrous internally; proximal lips without a
central notch before anthesis, not recurved after anthesis, not hood-shaped; proximal and distal lips drying the
same color as the rachillae, not joined to form a raised cupule, the proximal lip margins overlapping the distal
lip margins; distal lips well-developed; staminate and pistillate petals not emergent, not valvate throughout;
staminate flowers deciduous after anthesis; stamens 6; thecae diverging at anthesis, inserted almost directly
onto the filament apices, the connectives bifid but scarcely developed; anthers short and curled over at
anthesis; non-fertilized pistillate flowers persistent after anthesis; staminodial tubes crenulate at the apex,
those of non-fertilized flowers projecting and persistent after anthesis; fruits 10.6(8.212.8) mm long,
7.8(6.19.6) mm in diameter, the bases with a prominent, asymmetric stipe, the apices not conical, the
surfaces not splitting at maturity, with fibers emerging to give spiny fruits, not bumpy, not apiculate; locular
epidermis with operculum, smooth, with pores.
Distribution and habitat:From 650N747S and 45306645W in the central and northeastern
Amazon region of Brazil, the Guianas, and Venezuela at 192(7725) m elevation in lowland rainforest (Fig.
9). Galeano and Bernal (2010) report this species from extreme eastern Colombia in Amazonas and Guaina.
Taxonomic notes:Geonoma baculifera is closely related to G. congesta and G. calyptrogynoidea. These
three species all have the staminodial tubes of non-fertilized pistillate flowers projecting and persistent after
anthesis. Geonoma baculifera differs from its two relatives in its rachillae surfaces with faint to pronounced,
short, transverse ridges, and fruit surfaces without fibers emerging.
Subspecific variation:Two traits (stem branching, leaf division) vary within this species. There is no
evidence of geographic discontinuity, although there are a few outlying specimens from the central Amazon
region.
3. Geonoma bernalii Henderson, sp. nov. (Appendix IV, Plate 1)

A speciebus affinibus prophyllis haud brevibus necnon inaequaliter apiculatis, atque fructibus haud apiculatis crusta
haud tuberculata differt.
Type: COLOMBIA. Antioquia: Mun. San Luis, 105 km al este de Medelln en la carretera a Bogot, vereda Riosol, 1200
m, 9 October 1987, R. Bernal 1399 (holotype COL!, isotype NY!).
Plants 1.3 m tall; stems 1.5 m tall, 0.5 cm in diameter, solitary or clustered, cane-like; internodes 1.7(1.02.3)
cm long, yellowish and smooth. Leaves 8 per stem, undivided or irregularly pinnate, not plicate, bases of
blades running diagonally into the rachis; sheaths 6.3(6.06.5) cm long; petioles 5.8(2.09.5) cm long, drying
green or yellowish; rachis 13.3(12.514.1) cm long, 1.6(1.31.9) mm in diameter; veins not raised or slightly
raised and triangular in cross-section adaxially; pinnae 3 per side of rachis; basal pinna 9.8(9.010.5) cm long,
1.4(1.21.6) cm wide, forming an angle of 41(3744) with the rachis; apical pinna 7.0(6.08.0) cm long,
5.2(4.26.2) cm wide, forming an angle of 36(3338) with the rachis. Inflorescences branched 1 order;
38
HENDERSON
prophylls and peduncular bracts not ribbed with elongate, unbranched fibers, flattened, persistent; prophylls
5.3(4.85.7) cm long, not short and asymmetrically apiculate, the surfaces not ridged, without unequally wide
ridges; peduncular bracts 4.5 cm long, well-developed, inserted 0.6 cm above the prophyll; peduncles
6.6(6.46.7) cm long, 1.5(1.41.6) mm in diameter; rachillae 4(34), 4.3(4.14.4) cm long, 1.3(1.11.5) mm
in diameter, the surfaces without spiky, fibrous projections or ridges, drying brown with faint to pronounced,
short, transverse ridges, filiform with extended narrowed sections between the flower pits; flower pits
alternately arranged (sometimes distorted by twisting and contracting of rachillae), glabrous internally;
proximal lips without a central notch before anthesis, not recurved after anthesis, not hood-shaped; proximal
and distal lips drying the same color as the rachillae, joined to form a raised cupule, the margins not
overlapping; distal lips well-developed; staminate and pistillate petals not emergent, not valvate throughout;
anthesis; non-fertilized pistillate flowers deciduous after anthesis; staminodial tubes crenulate or shallowly
lobed at the apex, those of non-fertilized flowers not projecting and persistent after anthesis; fruits 6.0 mm
long, 6.2 mm in diameter, the bases without a prominent, asymmetric stipe, the apices not conical, the surfaces
not splitting at maturity, without fibers emerging, not bumpy and not apiculate; locular epidermis without
operculum, smooth, without pores.
Distribution and habitat:From 605607N and 75007502W on the eastern slopes of the
Central Cordillera in Colombia (Antioquia) at 1105(10101200) m elevation in montane rainforest (Fig. 9).
FIGURE 9. Distribution maps of Geonoma aspidiifolia subsp. aspidiifolia, G. aspidiifolia subsp. fusca, G. baculifera,
and G. bernalii.
39
Taxonomic notes:The two specimens examined have been previously determined as Geonoma
leptospadix. They bear a superficial resemblance to that species but differ in their alternately arranged flower
pits, not hood-shaped proximal lips, and cupular proximal and distal lips. Geonoma bernalii is similar to
several species in the G. lanata clade. It differs from these in its prophylls not short and asymmetrically
apiculate, and fruit surfaces not bumpy and not apiculate.
Subspecific variation:The only trait to vary between the two specimens examined is stem branching.
On both specimens pinnate leaves were measured, but one specimen also has undivided leaves present.
4. Geonoma braunii (Stauffer) Henderson, comb. & stat. nov.

Basionym: Geonoma spinescens var. braunii Stauffer (1997: 5). Type: VENEZUELA. Yaracuy: Mun. Nirgua,
ca. 5 km N of Nirgua, Cerro La Chapa, 1012N, 6833W, 12001300 m, 28 November1 December 1996,
A. Fernndez, F. Stauffer, R. Riina, K. Walter-Weissbeck & O. Kunert 10087 (holotype VEN n.v.).
Plants 2.0(1.52.5) m tall; stems 2.0(1.52.5) m tall, 1.0(0.81.2) cm in diameter, solitary or clustered, not
cane-like or cane-like; internodes 1.0(0.32.1) cm long, yellowish and smooth. Leaves 12(1013) per stem,
undivided or irregularly pinnate, not plicate, bases of blades running diagonally into the rachis; sheaths 6.5 cm
2.8) mm in diameter; veins not raised or slightly raised and triangular in cross-section adaxially; pinnae 2(1
4) per side of rachis; basal pinna 24.9(17.533.0) cm long, 4.7(3.010.5) cm wide, forming an angle of 35(21
53) with the rachis; apical pinna 15.9(13.521.8) cm long, 10.3(6.517.8) cm wide, forming an angle of
24(1828) with the rachis. Inflorescences branched 23 orders; prophylls and peduncular bracts not ribbed
with elongate, unbranched fibers, flattened, deciduous; prophylls 5.3(4.56.1) cm long, short and
asymmetrically apiculate, the margins curved around the stem, the surfaces flat with dense, felty, brown
tomentum, prophyll equal to and early deciduous with the peduncular bract, the surfaces not ridged, without
unequally wide ridges; peduncular bracts 5.8 cm long, well-developed, inserted 0.2(0.10.4) cm above the
prophyll; peduncles 10.6(6.014.5) cm long, 3.4(2.25.3) mm in diameter; rachillae 24(2226), 7.0(6.08.5)
cm long, 0.8(0.71.0) mm in diameter, the surfaces without spiky, fibrous projections or ridges, drying brown,
with faint to pronounced, short, transverse ridges, filiform with extended narrowed sections between the
flower pits; flower pits alternately arranged (sometimes distorted by twisting and contracting of rachillae),
glabrous internally; proximal lips without a central notch before anthesis, not recurved after anthesis, not
hood-shaped; proximal and distal lips drying the same color as the rachillae, joined to form a raised cupule,
the margins not overlapping; distal lips well-developed; staminate and pistillate petals not emergent, not
valvate throughout; staminate flowers deciduous after anthesis; stamens 6; thecae diverging at anthesis,
inserted almost directly onto the filament apices, the connectives bifid but scarcely developed; anthers short
and curled over at anthesis; non-fertilized pistillate flowers deciduous after anthesis; staminodial tubes
crenulate or shallowly lobed at the apex, those of non-fertilized flowers not projecting and persistent after
anthesis; fruits 7.4(6.58.1) mm long, 6.0(5.16.9) mm in diameter, the bases without a prominent stipe, the
apices not conical, the surfaces not splitting at maturity, without fibers emerging, bumpy from the numerous,
subepidermal, tangential, short fibers present, these coming to a point at fruit apices; locular epidermis
without operculum, smooth, without pores.
Distribution and habitat:From 10091015N and 68286837W in the Coastal Cordillera in
Venezuela (Yaracuy), at 1192(10451300) m elevation in montane rainforest (Fig. 10).
Taxonomic notes:Stauffer (1997) recognized this species as a variety of Geonoma spinescens. It is
here recognized at the species level, differing from G. spinescens in its alternately arranged flower pits.
Subspecific variation:Three traits (stem branching, stem type, leaf division) vary within this species,
but all specimens come from the same area.
40
HENDERSON
FIGURE 10. Distribution maps of Geonoma braunii, G. brenesii, G. brongniartii subsp. brongniartii, and G.
brongniartii subsp. pascoensis.
5. Geonoma brenesii Grayum (1998: 322). Type: COSTA RICA. Alajuela: Reserva Biolgica de San Ramn,
road from Las Lagunas to Colonia Palmarea, 1004N, 8432W, 8501100 m, 30 May 1986, G. de Nevers,
B. Hammel & C. Gmez 7789 (holotype MO!, isotypes CR!, NY!).
Plants 0.8(0.51.0) m tall; stems 0.3(0.20.4) m tall, 1.2 cm in diameter, solitary, not cane-like; internodes 0.5
cm long, not scaly. Leaves 9(811) per stem, irregularly pinnate, not plicate, bases of blades running
diagonally into the rachis; sheaths 8.2(6.510.0) cm long; petioles 31.8(21.038.5) cm long, drying green or
yellowish; rachis 24.6(18.829.8) cm long, 2.8(1.94.3) mm in diameter; veins raised and rectangular in
cross-section adaxially; pinnae 3(34) per side of rachis; basal pinna 28.3(21.536.5) cm long, 4.1(2.010.2)
cm wide, forming an angle of 43(3053) with the rachis; apical pinna 16.7(13.521.0) cm long, 9.5(6.5
12.0) cm wide, forming an angle of 29(2532) with the rachis. Inflorescences unbranched; prophylls and
peduncular bracts ribbed with elongate, unbranched fibers, both bracts tubular, narrow, elongate, closely
sheathing the peduncle, more or less persistent; prophylls 13.5(10.017.5) cm long, not short and
7.1) cm long, vestigial, inserted 4.9(3.47.0) cm above the prophyll; peduncles 24.3(15.037.5) cm long,
2.1(1.42.8) mm in diameter; rachillae 1, 12.5(8.516.0) cm long, 3.6(2.85.0) mm in diameter, the surfaces
without spiky, fibrous projections or ridges, drying brown or yellow-brown, without short, transverse ridges,
not filiform and not narrowed between the flower pits; flower pits spirally arranged, glabrous internally;
41
proximal lips with a central notch before anthesis, often the two sides of the notch overlapping, the lip, more
or less heartshaped, not recurved after anthesis, not hood-shaped; proximal and distal lips drying the same
color as the rachillae, not joined to form a raised cupule, the proximal lip margins overlapping the distal lip
margins; distal lips well-developed; staminate and pistillate petals not emergent, not valvate throughout;
anthesis; non-fertilized pistillate flowers persistent after anthesis; staminodial tubes lobed, the lobes not
spreading at anthesis, not acuminate, those of non-fertilized flowers not projecting and persistent after
Distribution and habitat:From 10021023N and 83598451W in Costa Rica at 1293(967
1550) m elevation in montane rainforest (Fig. 10).
Taxonomic notes:Geonoma brenesii is a member of a group of four Central American species, part of
the G. cuneata clade, including G. monospatha, G. hugonis, and G. epetiolata. They all have unbranched or
few-branched inflorescences and share the character state of the staminodial tubes being lobed at the apex, but
the lobes are not spreading at anthesis and are not acuminate. Geonoma brenesii is most similar to G.
monospatha, differing in its tubular, narrow, elongate, sheathing, persistent prophylls and peduncular bracts
which are ribbed with unbranched fibers.
Subspecific variation:No traits vary within this species.
6. Geonoma brongniartii Martius (1843: 24). Type: BOLIVIA. Cochabamba: Prov. Carrasco, no date, A.
dOrbigny 39 (holotype P n.v., isotype F!).
cane-like; internodes 0.5(0.30.8) cm long, not scaly. Leaves 8(412) per stem, undivided or irregularly
pinnate, not plicate, bases of blades running diagonally into the rachis; sheaths 13.6(5.022.0) cm long;
petioles 48.3(14.090.0) cm long, drying green or yellowish; rachis 53.4(20.080.0) cm long, 3.7(1.96.2)
mm in diameter; veins raised and rectangular in cross-section adaxially or not raised or slightly raised and
triangular in cross-section adaxially; pinnae 5(119) per side of rachis; basal pinna 30.0(16.054.0) cm long,
11.5(2.037.5) cm wide, forming an angle of 32(2052) with the rachis. Inflorescences unbranched or
branched 1 order; prophylls and peduncular bracts ribbed with elongate, unbranched fibers, both bracts
tubular, narrow, elongate, closely sheathing the peduncle, more or less persistent; prophylls 19.6(5.735.5) cm
long, not short and asymmetrically apiculate, the surfaces not ridged, without unequally wide ridges;
peduncular bracts 19.4(12.631.5) cm long, well-developed, inserted 5.7(0.518.5) cm above the prophyll;
peduncles 34.6(18.064.0) cm long, 2.9(1.56.5) mm in diameter; rachillae 1(16), 24.4(8.646.0) cm long,
3.7(1.77.3) mm in diameter, the surfaces without spiky, fibrous projections or ridges, drying brown or
yellow-brown, the surfaces without short, transverse ridges, not filiform and not narrowed between the flower
pits; flower pits usually spirally arranged, sometimes tricussately, then the groups not closely spaced nor
consistently arranged throughout the rachillae, glabrous internally (sometimes some hairs along the proximal
and distal lip margins); proximal lips without a central notch before anthesis, not recurved after anthesis, not
hood-shaped; proximal and distal lips drying the same color as the rachillae, not joined to form a raised
cupule, the proximal lip margins overlapping the distal lip margins; distal lips a scarcely raised rim; staminate
and pistillate petals not emergent, not valvate throughout; staminate flowers deciduous after anthesis; stamens
6; thecae diverging at anthesis, inserted almost directly onto the filament apices, the connectives bifid but
scarcely developed; anthers short and curled over at anthesis; non-fertilized pistillate flowers deciduous after
42
HENDERSON
anthesis; staminodial tubes crenulate or shallowly lobed at the apex, those of non-fertilized flowers not
projecting and persistent after anthesis; fruits 6.8(5.38.3) mm long, 5.7(4.56.5) mm in diameter, the bases
without a prominent stipe, the apices not conical, the surfaces not splitting at maturity, without fibers
emerging, bumpy from the numerous, subepidermal, tangential, short fibers present, these coming to a point at
fruit apices; locular epidermis without operculum, smooth or sculpted and then usually also with a raised,
meridional ridge, without pores.
Taxonomic notes:The holotype of Geonoma brongniartii at P has not been seen. A specimen from F,
labeled Geonoma elegans Mart., Bolivia, Yapacani, 400 m, Otto Kuntze VI 92, has written on it, apparently
in Wessels Boers handwriting, leg. dOrbigny 39, Type of G. brongniartii. This specimen, clearly not
representing G. elegans, is referred to by Wessels Boer (1968) as being an isotype of G. brongniartii. Although
it is not clear why Wessels Boer thought the specimen was an isotype, it is treated as such here.
Subspecific variation:Five traits (stem branching, leaf division, adaxial veins, inflorescence
branching, locular epidermis sculpting) vary within this species. There is evidence of geographic
discontinuity, but some of the gaps in the distribution may be a result of insufficient collecting. Leaving aside
stem branching, leaf division, and locular epidermis sculpting (for which there are few data), there is no
correspondence between geography and variation in adaxial veins or inflorescence branching, except for one
subgroup of specimens from sub-Andean Peru. These have raised adaxial veins and mostly branched
inflorescences. These specimens differ significantly from all other specimens in eight variables (plant height,
number of pinnae, basal pinna width, apical pinna width, rachillae length, rachillae width, fruit length, and
fruit diameter)(t-test, P <0.05). Based on this, they are recognized as a separate subspecies (subsp.
pascoensis). All other specimens are included in subsp. brongniartii.
Key to the subspecies of G. brongniartii
1
-
Inflorescences unbranched; fruits 6.9(5.38.3) mm long; widespread ............................................subsp. brongniartii

Inflorescences unbranched or usually branched 1 order; fruits 5.9(5.76.1) mm long; Peru (Junn, Pasco) .................
............................................................................................................................................................subsp. pascoensis
6a. Geonoma brongniartii subsp. brongniartii

Geonoma metensis Karsten (1856: 409). Type: COLOMBIA. Meta: Villavicencio, no date, H. Karsten s.n. (holotype LE
n.v.).
Geonoma cuneifolia Burret (1931a: 199). Type: PERU. Loreto: Ro Ucayali, 215 m, 17 November 1923, G. Tessmann
3317 (holotype B, destroyed, isotype NY!).
Leaves veins raised and rectangular in cross-section adaxially or not raised or slightly raised and triangular in
cross-section adaxially. Inflorescences unbranched; fruits 6.9(5.38.3) mm long, 5.8(4.96.5) mm in diameter.
Distribution and habitat:From 714N1750S and 63397720W in the western Amazon and
sub-Andean regions of Ecuador, Peru, Bolivia, and Brazil, with an outlier in Venezuela, at 373(751720) m
elevation, often in low-lying, flooded areas, in lowland or, less often, montane rainforest (Fig. 10).
In southern Colombia and Ecuador, specimens are relatively uniform and have pinnate leaves, raised
adaxial veins, more pinnae than usual, and slender rachillae. There is an absence of specimens from
southeastern Ecuador and adjacent Peru. In this area, Geonoma macrostachys is abundant.
In northeastern Peru (Loreto) and adjacent Colombia and Brazil, specimens are similar to those from
southern Colombia and Ecuador, except pinnae are fewer and some leaves undivided, and rachillae larger.
Some specimens are considerably larger than others (e.g., Vsquez 9744). Geonoma brongniartii is again
absent from southwest of Iquitos, where G. macrostachys is abundant.
In these northern populations, in Colombia, Ecuador, northeastern Peru and adjacent Brazil, there is
geographical variation. Regression shows there are significant associations between longitude and eight leaf
variables. Squared multiple R for the regression of petiole length on longitude is 0.58, number of pinnae 0.24,
basal pinna length 0.50, basal pinna width 0.29, basal pinna angle 0.34, apical pinna length 0.20, apical pinna
43
width 0.17, and apical pinna angle 0.50. There is a change in leaf shape, from specimens in the east having
shorter petioles and fewer, wider and longer pinnae with narrower angles to those in the west having longer
petioles, more, narrower and shorter pinnae with wider angles.
In Brazil (Acre) and adjacent Peru most specimens are relatively uniform. There are some specimens
(e.g., Daly 10582, 10947) with thicker rachillae and the flower pits arranged in closer spirals, as in central
Peru (see below).
There is extreme variation in central Peru on eastern Andean slopes and adjacent areas. In Ayacucho,
Hunuco, Pasco, San Martn, and Ucayali some specimens have larger leaves and longer, thicker rachillae
along which the flower pits are wider and are arranged in closer spirals (large-sized morphotype). These
larger-sized specimens sometimes occur together with the more usual-sized specimens. Some resemble G.
poeppigiana and two (Schunke 16280, 9912) have bracts more like those of G. poeppigiana), and may be
hybrids with that species.
Two other specimens (Foster 7846, Roncal 182) from Pasco have longer stems. One (Foster 7846) has a
stem that is reported to be 23 m tall, and the other (Roncal 182) has cane-like stems with the internodes
longer than wide. Both have short inflorescences, and in Roncal 182 they appear to be pendulous. One other
specimen (Smith 3849) appears similar. These occur near to an isolated population of Geonoma deversa
subsp. deversa, killipii morphotype, and it is possible they represent hybrids with that morphotype. These and
other possible hybrids are excluded from the above description.
In San Martn, one specimen (Schunke 8080) has the shortest prophyll (5.7) cm and interbract distance
(0.5 cm) of all specimens, and the adaxial veins are not raised. It occurs sympatrically with another isolated
population of G. deversa subsp deversa, and may also be a hybrid.
One specimen from Hunuco (Moore 8355), with branched inflorescences, has unusually short rachillae
and comes from an unusually high elevation (1575 m). It may be a hybrid.
Most specimens from southern Peru (Cusco, Madre de Dios, Puno) and Bolivia have non-raised adaxial
veins, and thin, elongate rachillae along which the pits may be tricussately arranged, especially in the central
part of the rachilla. A few specimens from northern Bolivia (Moreno 124, Fuentes 3911, Macia 3986, Beck
18258, Croat 51638, Beck 16466, Williams 941, Williams 939) and southern Peru (Foster 9721, 9576) have
thicker rachillae and the flower pits arranged in closer spirals, as found in the large central Peruvian
specimens. A few specimens (Moreno 227, Gerlach 214, Foster 13393, Hodge 6079, Plowman 5062) from
southern Peru and Bolivia have exceptionally long interbract distances (10.318.5 cm).
There is variation in connectives in this species. Specimens are scored as having the thecae inserted
almost directly onto the filament apices, the connectives bifid but scarcely developed. However, in some
specimens the connectives appear not to be bifid, and are similar to those of G. macrostachys.
6b. Geonoma brongniartii subsp. pascoensis Henderson, subsp. nov. (Appendix IV, Plates 2 & 3)
Geonomae brongniartii subsp. brongniartii inflorescentiis saepe ramosis atque fructibus parvioribus differt.
Type: PERU. Pasco: near Pozuzo, opposite the town, steep slope above river opposite margin of Ro Pozuzo, along road
that runs south, 11 September 1998, A. Henderson, E. Ferreira & M. Arakaki 3012 (holotype USM!, isotype NY!).
Leaves veins raised and rectangular in cross-section adaxially. Inflorescences unbranched or usually branched
1 order; fruits 5.9(5.76.1) mm long, 4.8(4.55.2) mm in diameter.
Distribution and habitat:From 9371033S and 74557534W in sub-Andean regions of Peru
(Junn, Pasco) at 417(2001000) m elevation in lowland rainforest (Fig. 10).
7. Geonoma calyptrogynoidea Burret (1930a: 223). Type: COLOMBIA. Antioquia: La Mesa, no date, W.
Kalbreyer 1398 (holotype B, destroyed). Neotype (designated by de Nevers & Grayum 1988): COLOMBIA.
Choc: Zona de Urab, Cerro del Cuchillo, sector Cuchillo Blanco, 1020 m, 15 October 1987, D. Crdenas
668 (neotype MO!).
44
HENDERSON
Plants 3.4(2.05.0) m tall; stems 2.9(1.54) m tall, 2.1(2.02.2) cm in diameter, solitary or clustered, canelike; internodes 2.9(2.63.1) cm long, yellowish and smooth. Leaves 12(815) per stem, undivided or
9.9) mm in diameter; veins raised and rectangular in cross-section adaxially; pinnae 3(18) per side of rachis;
basal pinna 64.9(42.082.5) cm long, 22.0(11.540.5) cm wide, forming an angle of 27(1542) with the
rachis; apical pinna 37.1(17.548.0) cm long, 23.2(14.333.0) cm wide, forming a 29(2235) angle with the
rachis. Inflorescences branched 1 order; prophylls and peduncular bracts not ribbed with elongate, unbranched
fibers, flattened, deciduous or persistent; prophylls 22.1(16.028.2) cm long, not short and asymmetrically
apiculate, the surfaces ridged with close, equal, parallel, non-dividing ridges, scarcely tomentose between the
ridges, without unequally wide ridges; peduncular bracts 25.3(21.530.0) cm, well-developed, inserted
projections or ridges, drying brown or yellow-brown, without short, transverse ridges, not filiform and not
anthesis; non-fertilized pistillate flowers persistent after anthesis; staminodial tubes crenulate at the apex,
those of non-fertilized flowers projecting and persistent after anthesis; fruits 13.2(11.815.5) mm long,
surfaces not splitting at maturity, with fibers emerging, not bumpy, not apiculate; locular epidermis with
operculum, smooth, with pores.
Distribution and habitat:From 820N102 and 74117842W in eastern Panama, the Pacific
coast of Colombia and Ecuador, and the Magdalena and Cauca valleys in Colombia, at 212(15550) m
elevation in lowland rainforest (Fig. 11).
Taxonomic notes:This species was included as a synonym of Geonoma congesta by Henderson et al.
(1995), but this placement was disputed by de Nevers & Grayum (1998). While closely related, G.
calyptrogynoidea differs from G. congesta by its proximal lips of flower pits without a central notch before
anthesis.
Subspecific variation:Two traits (stem branching, leaf division) vary within this species. There is
evidence of geographic discontinuity and the species occurs in two areaseastern Panama and the Pacific
coast of Colombia and Ecuador (the gap in southern Colombia is likely to be an artifact of insufficient
collecting), and the Magdalena and Cauca valleys in Colombia. The Magdalena and Cauca population has
inflorescences with more, longer rachillae, but there are too few specimens to test for differences between the
two areas.
8. Geonoma camana Trail (1876: 324). Taenianthera camana (Trail) Burret (1930a: 270)
Type: BRAZIL. Amazonas: San Antonia da Boa Vista, Rio Javari, 4 December 1874, J. Trail 977/CLXXXII (holotype K
n.v., isotype P!).
Geonoma lagesiana Dammer (1907: 121). Taenianthera lagesiana (Dammer) Burret (1930a: 270). Type: BRAZIL.
Amazonas: Rio Juru, Jurumirim, August 1901, E. Ule 5745 (holotype not known, isotype MG!).
Plants 2.0(0.84.0) m tall; stems 0.5(0.11.3) m tall, 2.3(1.72.8) cm in diameter, solitary, not cane-like;
internodes 0.6(0.50.6) cm long, not scaly. Leaves 9(514) per stem, irregularly pinnate or regularly pinnate
and the pinnae with 1 main vein only, not plicate, the bases of blades running diagonally into the rachis;
sheaths 19.2(5.050.0) cm long; petioles 71.0(24.0140.0) cm long, drying green or yellowish; rachis
45
83.1(46.0137.0) cm long, 5.7(2.89.4) mm in diameter; veins not raised or slightly raised and triangular in
cross-section adaxially; pinnae 17(238) per side of rachis; basal pinna 30.5(13.457.0) cm long, 4.3(0.3
24.0) cm wide, forming an angle of 53(2095) with the rachis; apical pinna 19.9(6.836.0) cm long,
12.6(1.731.0) cm wide, forming an angle of 44(3160) with the rachis. Inflorescences unbranched;
prophylls and peduncular bracts ribbed with elongate, unbranched fibers, both bracts tubular, narrow,
elongate, closely sheathing the peduncle, more or less persistent; prophylls 11.6(4.525.5) cm long, not short
and asymmetrically apiculate, the surfaces not ridged, without unequally wide ridges; peduncular bracts
20.4(12.028.5) cm long, well-developed, inserted 2.7(0.64.4) cm above the prophyll; peduncles 44.9(21.5
100.0) cm long, 3.9(2.15.8) mm in diameter; rachillae 1, 23.8(10.536.0) cm long, 5.8(2.88.3) mm in
diameter, the surfaces without spiky, fibrous projections or ridges, drying brown or yellow-brown, without
short, transverse ridges, not filiform and not narrowed between the flower pits; flower pits spirally arranged,
glabrous internally; proximal lips pits with a central notch before anthesis, often the two sides of the notch
overlapping, not recurved after anthesis, not hood-shaped; proximal and distal lips drying the same color as
the rachillae, not joined to form a raised cupule, the proximal lip margins overlapping the distal lip margins;
distal lips well-developed; staminate and pistillate petals not emergent, not valvate throughout; staminate
flowers deciduous after anthesis; stamens 6; thecae diverging at anthesis, inserted directly onto the apiculate
filament apices; anthers not short and curled at anthesis, usually elongate, spiraled and twisted or sometimes
remaining straight; non-fertilized pistillate flowers deciduous after anthesis; staminodial tubes lobed at the
apex, the lobes spreading at anthesis, acuminate, those of non-fertilized flowers not projecting and persistent
after anthesis; fruits 10.7(8.713.5) mm long, 6.9(6.08.9) mm in diameter, the bases without a prominent
stipe, the apices not conical, the surfaces not splitting at maturity, without fibers emerging, not bumpy, not
apiculate; locular epidermis with operculum, smooth, without pores.
Distribution and habitat:From 0081321S and 68467819W in the western Amazon region of
Colombia, Ecuador, Peru, and Brazil at 210(87850) m elevation in lowland rainforest (Fig. 11).
Taxonomic notes:In previous treatments (e.g., Henderson et al., 1995), Geonoma camana was said to
be characterized by its thick, smooth pinnae with conspicuous, submarginal veins. In the present work, these
potential characters have been found to be inconsistent and difficult to score, and are therefore not used.
Geonoma camana differs from similar species in the G. macrostachys clade (G. chlamydostachys, G.
chococola, G. deneversii, and G. maxima) in its fruit surfaces not splitting, and locular epidermis with
operculum and without pores.
Subspecific variation:No traits vary within this species. There is a gap in the distribution of G.
camana, between Ecuador and adjacent Colombia and Peru in the west, and Colombia, Peru, and Brazil in the
east. However, the intervening area is poorly collected for palms, and this gap may be an artifact of
insufficient collecting.
There is geographical variation in this species. Regression shows there are significant associations
between longitude and two plant, seven leaf, and three inflorescence variables. Squared multiple R for the
regression of plant height on longitude is 0.36, stem height 0.24, petiole length 0.40, rachis width 0.54, basal
pinna length 0.31, basal pinna angle 0.12, apical pinna length 0.50, apical pinna width 0.10, apical pinna angle
0.19, peduncle width 0.25, rachilla length 0.19, and rachilla width 0.18. From east to west, specimens become
smaller. In particular, there is a change in leaf size and shape, from specimens in the east having longer basal
and apical pinnae with narrower angles, to those in the west shorter basal and apical pinnae with wider angles.
Four specimens from Acre, Brazil (Ferreira 125, 136, 163, 165) are much smaller than others and may
represent hybrids between G. camana and G. macrostachys. Other specimens from Loreto, Peru (Balick 1131)
and from Amazonas, Brazil (Pardini 45) also appear intermediate between these two species. These potential
hybrids are not included in the above description or analysis.
9. Geonoma chlamydostachys Galeano (1986: 71). Type: COLOMBIA. Antioquia: Mun. San Luis, Ro
Claro, 4001000 m, 13 January 1983, A. Hernndez et al. 685 (holotype HUA n.v.).
46
HENDERSON
FIGURE 11. Distribution maps of Geonoma calyptrogynoidea, G. camana, G. chlamydostachys, G. chococola subsp.
chococola, and G. chococola subsp. awaensis.
Plants 1.8(1.03.0) m tall; stems 1.4(0.62.5) m tall, solitary. Leaves irregularly pinnate, not plicate, the bases
of blades running diagonally into the rachis; sheaths 19.0 cm long; petioles 8.0 cm long, drying green or
yellowish; rachis 44.6(36.557.0) cm long, 4.3(2.86.0) mm in diameter; veins not raised or slightly raised
and triangular in cross-section adaxially; pinnae 5(39) per side of rachis; basal pinna 33.8(22.041.0) cm
long, 11.6(1.817.0) cm wide, forming an angle of 32(2343) with the rachis; apical pinna 17.0(12.523.0)
cm long, 11.7(8.414.5) cm wide, forming an angle of 38(3245) with the rachis. Inflorescences unbranched;
glabrous internally; proximal lips with a central notch before anthesis, often the two sides of the notch
overlapping, not recurved after anthesis, not hooded; proximal and distal lips drying the same color as the
rachillae, not joined to form a raised cupule, the proximal lip margins overlapping the distal lip margins; distal
lips well-developed; staminate and pistillate petals not emergent, not valvate throughout; staminate flowers
deciduous after anthesis; stamens 6; thecae diverging at anthesis, inserted directly onto the apiculate filament
47
apices; anthers not short and curled at anthesis, usually elongate, spiraled and twisted or sometimes remaining
straight; non-fertilized pistillate flowers deciduous after anthesis; staminodial tubes lobed at the apex, the
lobes spreading at anthesis, acuminate, those of non-fertilized flowers not projecting and persistent after
anthesis; fruits 12.3 mm long, 9.2 mm in diameter, the bases without a prominent stipe, the apices not conical,
the surfaces not splitting at maturity, without fibers emerging, not bumpy, not apiculate; locular epidermis
with operculum, smooth, with pores.
Distribution and habitat:From 718533N and 74187536W in the Central Cordillera in
Colombia (Antioquia), with an outlier in the Eastern Cordillera, at 890(5501450) m elevation in lowland to
montane rainforest (Fig. 11).
Taxonomic notes:Geonoma chlamydostachys differs from similar species in the G. macrostachys clade
(G. camana, G. chococola, G. deneversii, and G. maxima) in its 6 stamens; thecae diverging at anthesis and
inserted directly onto the apiculate filament apices; and fruit surfaces not splitting.
Subspecific variation: No traits vary within this species, nor is there any geographic discontinuity.
10. Geonoma chococola Wessels Boer (1968: 103). Type: COLOMBIA. Valle del Cauca: Buenaventura, 6
May 1926, O. Cook 84 (holotype US!).
Plants 2.8(1.54.0) m tall; stems 1.9(1.03.0) m tall, 3.5 cm in diameter, solitary or clustered, cane-like;
internodes 3.7 cm long, yellowish and smooth. Leaves 11(721) per stem, undivided or irregularly pinnate,
not plicate, bases of blades running diagonally into the rachis; sheaths 16.5(8.023.0) cm long; petioles
37.9(15.361.0) cm long, drying green or yellowish; rachis 111.7(80.0137.0) cm long, 7.6(4.612.5) mm in
diameter; veins raised and rectangular in cross-section adaxially; pinnae 3(17) per side of rachis; basal pinna
48.5(30.080.0) cm long, 13.2(4.543.5) cm wide, forming an angle of 27(1638) with the rachis; apical
pinna 30.1(21.740.5) cm long, 26.5(20.535.0) cm wide, forming an angle of 28(2040) with the rachis.
Inflorescences unbranched or branched 1 order; prophylls and peduncular bracts ribbed with elongate,
unbranched fibers, both bracts tubular, narrow, elongate, closely sheathing the peduncle, more or less
persistent; prophylls 35.0(23.749.5) cm long, not short and asymmetrically apiculate, the surfaces not ridged,
without unequally wide ridges; peduncular bracts 60.8(38.583.0) cm long, well-developed, inserted 2.2(1.5
3.0) cm above prophyll; peduncles 86.3(46.0143.0) cm long, 6.4(3.98.6) mm in diameter; rachillae 1(14),
27.8(16.547.0) cm long, 10.1(5.815.2) mm in diameter, the surfaces without spiky, fibrous projections or
ridges, drying brown or yellow-brown, without short, transverse ridges, not filiform and not narrowed
between the flower pits; flower pits spirally arranged, glabrous internally; proximal lips with a central notch
before anthesis, often the two sides of the notch overlapping, not recurved after anthesis, not hood-shaped;
lip margins overlapping the distal lip margins; distal lips well-developed; staminate and pistillate petals not
emergent, not valvate throughout; staminate flowers persistent after anthesis; stamens 6; thecae diverging at
anthesis, inserted directly onto the apiculate filament apices; anthers not short and curled at anthesis, usually
elongate, spiraled and twisted or sometimes remaining straight; non-fertilized pistillate flowers deciduous
after anthesis; staminodial tubes lobed at the apex, the lobes spreading at anthesis, acuminate, those of nonfertilized flowers not projecting and persistent after anthesis; fruits 18.9(14.824.9) mm long, 15.7(11.317.8)
mm in diameter, the bases without a prominent stipe, the apices not conical, the surfaces splitting deeply and
longitudinally at maturity to reveal mesocarp with dense layer of radial fibers, with fibers emerging, not
bumpy, not apiculate; locular epidermis with operculum, smooth, with pores.
Taxonomic notes:Geonoma chococola was said by Henderson et al. (1995) to have 12 stamens, but
this appears to be a mistake. All eight specimens examined with staminate flowers have 6 stamens. In the
following treatment, G. awaensis is recognized as a subspecies of G. chococola; it is not recognized at the
species level because it shares the same character state combinations as G. chococola.
48
HENDERSON
Subspecific variation:Three traits vary within this species (stem branching, leaf division,
inflorescence branching). There is geographic discontinuity and specimens occur in two separate areas. Based
on one trait distribution (inflorescence branching) and geography, two subgroups can be recognized and these
are treated as subspecies (subspp. chococola, awaensis). There are too many missing data for most variables
for analysis, but subsp. chococola has significantly longer and wider rachillae (t-test, P <0.05).
Key to the subspecies of G. chococola
1
-
Inflorescences unbranched................................................................................................................. subsp. chococola

Inflorescences branched with 24 rachillae.......................................................................................... subsp. awaensis
10a. Geonoma chococola subsp. chococola

Inflorescences unbranched; rachillae 31.1(20.047.0) cm long, 11.2(8.115.2) mm in diameter.
Distribution and habitat:From 302545N and 76157732W on the Pacific coast in western
Colombia, and one outlying specimen in the Cauca Valley, at 109(0500) m elevation in lowland rainforest
(Fig. 11).
10b. Geonoma chococola subsp. awaensis (Henderson, Borchsenius & Balslev) Henderson, comb. & stat.
nov.
Basionym: Geonoma awaensis Henderson, Borchsenius & Balslev (2008: 60). Type: ECUADOR. Esmeraldas: Aw
Reserve, footpath to Ro Mira, 115N, 7840W, 216 m, 20 September 1993, H. Beck, A. Ortiz, H. Cantincuz & A.
Cantincuz 2176 (holotype NY!, isotype QCNE n.v.).
Inflorescences branched; rachillae 18.6(16.520.5) cm long, 6.5(5.87.1) mm in diameter.

Distribution and habitat:From 048115N and 78267844W on western Andean slopes in
northwestern Ecuador at 275(200500) m elevation in lowland rainforest (Fig. 11).
11. Geonoma concinna Burret (1930a: 229). Type: COLOMBIA. Antioquia: Tabor, 1950 m, 27 January
1880, W. Kalbreyer 1367 (holotype B, destroyed). Neotype (selected by Bernal et al. 1989): COLOMBIA.
Antioquia: carretera Granada-San Luis, 5.5 km adelante de El Choc, 1750 m, 2021 September 1987, R.
Bernal & L. Tobn 1358 (neotype COL!, isoneotype NY!).
Plants 1.8(1.52.5) m tall; stems 3.5 m tall, 1.0(0.91.2) cm in diameter, clustered, cane-like; internodes
3.5(2.54.5) cm long, yellowish and smooth. Leaves 8 per stem, undivided or irregularly pinnate, not plicate,
bases of blades running diagonally into the rachis; sheaths 14.0 cm long; petioles 9.0 cm long, drying green or
yellowish; rachis 27.4(20.338.5) cm long, 3.4(3.13.8) mm in diameter; veins raised and rectangular in
12.0) cm wide, forming an angle of 38(3242) with the rachis. Inflorescences branched 2 orders; prophylls
and peduncular bracts not ribbed with elongate, unbranched fibers, flattened, deciduous; prophylls 10.8(9.0
12.5) cm long, not short and asymmetrically apiculate, the surfaces ridged with close, equal, parallel, nondividing ridges, scarcely tomentose between the ridges, without unequally wide ridges; peduncular bracts 7.2
cm long, well-developed, inserted 0.4(0.20.6) cm above the prophyll; peduncles 11.1(8.213.2) cm long,
surfaces without spiky, fibrous projections or ridges, drying brown, with faint to pronounced, short, transverse
internally; proximal lips without a central notch before anthesis, not recurved after anthesis, hood-shaped at
anthesis, sometimes splitting post-anthesis; proximal and distal lips drying the same color as the rachillae, not
49
joined to form a raised cupule, the proximal lip margins overlapping the distal lip margins; distal lips welldeveloped; staminate and pistillate petals not emergent, not valvate throughout; staminate flowers deciduous
after anthesis; stamens 6; thecae diverging at anthesis, inserted almost directly onto the filament apices, the
connectives bifid but scarcely developed; anthers short and curled over at anthesis; non-fertilized pistillate
flowers deciduous after anthesis; staminodial tubes crenulate or shallowly lobed at the apex, those of nonfertilized flowers not projecting and persistent after anthesis; fruits 7.7(7.18.3) mm long, 6.7(6.17.3) mm in
diameter, the bases without a prominent stipe, the apices not conical, the surfaces not splitting at maturity,
without fibers emerging, bumpy from the numerous, subepidermal, tangential, short fibers present, these
coming to a point at fruit apices; locular epidermis with operculum, smooth, without pores.
Taxonomic notes:Geonoma concinna is a member of the G. congesta clade, and most closely related to
G. concinnoidea, from which it differs by its fruits without fibers emerging. Geonoma concinna was
considered by Henderson et al. (1995) to be a poorly understood species. This is still true because of the lack
of specimens (only four specimens known). In Henderson et al., specimens from Panama were included in G.
concinna; here they are recognized as a distinct species, G. concinnoidea.
Subspecific variation:One trait (leaf division) varies within this species. There is geographic
discontinuity and specimens occur in two separate areas in Colombia. There are too few specimens for
analysis. Based on the trait distribution and geography, two subgroups can be recognized and these are treated
as subspecies (subspp. concinna, simplex). Subspecies concinna occurs at higher elevations than subsp.
simplex; 1475(12001750) versus 777(755800) m elevation .
Key to the subspecies of G. concinna
1
-
Leaves pinnate; Central Cordillera ....................................................................................................... subsp. concinna

Leaves undivided; Western Cordillera.....................................................................................................subsp. simplex
11a. Geonoma concinna subsp. concinna

Leaves pinnate.
Distribution and habitat:From 605654N and 75047505W in Colombia (Antioquia) in the
Central Cordillera at 1475(12001750) m elevation in montane rainforest (Fig. 12).
11b. Geonoma concinna subsp. simplex Henderson, subsp. nov. (Appendix IV, Plates 4 & 5)
A Geonoma concinna subsp. concinna foliis simplicibus differt.
Type: COLOMBIA. Valle: near Yatacu, Alto Anchicaya, near CVC hydroelectric plant headquarters, valley of Ro
Dagua, 338N 7645W, 710800 m, 16 July 1984, A. Gentry & M. Monsalve 48188 (holotype NY!, isotype MO!).
Leaves undivided.
Distribution and habitat:From 338340N and 76457650W in Colombia (Valle) on the
Western Cordillera at 777(755800) m elevation in lowland rainforest (Fig. 12).
12. Geonoma concinnoidea Henderson, sp. nov. (Appendix IV, Plates 6 & 7)
Geonomae concinnae crusta fructuum fibris emergentibus differt.
Type: PANAMA. Comarca de San Blas: Yar Bired (San Jos), between Cangandi and San Jos, 920N
7908W, 400500 m, 5 February 1986, G. de Nevers & H. Herrera 6942 (holotype NY!, isotype MO!).
50
HENDERSON
FIGURE 12. Distribution maps of Geonoma concinna subsp. concinna, G. concinna subsp. simplex, G. concinnoidea
subsp. concinnoidea, G. concinnoidea subsp. coclensis, and G. concinnoidea subsp. jefensis.
Plants 2.0(1.03.1) m tall; stems 2.7(1.64.0) m tall, 0.6(0.41.0) cm in diameter, solitary or clustered, canelike; internodes 1.8(0.74.5) cm long, yellowish and smooth. Leaves 6(58) per stem, undivided or irregularly
pinnate, not plicate, bases of blades running diagonally into the rachis; sheaths 7.5(4.512.0) cm long; petioles
23.1(20.025.6) cm long, 5.7(3.66.8) cm wide, forming an angle of 30(2244) with the rachis; apical pinna
14.7(9.020.5) cm long, 5.7(3.06.4) cm wide, forming an angle of 30(2037) with the rachis. Inflorescences
branched 2 orders; prophylls and peduncular bracts not ribbed with elongate, unbranched fibers, flattened,
deciduous; prophylls 5.5(3.09.1) cm long, not short and asymmetrically apiculate, the surfaces ridged with
close, equal, parallel, non-dividing ridges, scarcely tomentose between the ridges, without unequally wide
ridges; peduncular bracts 4.5 cm long, well-developed, inserted 0.2(0.10.3) cm above the prophyll;
1.4(0.81.9) mm in diameter, the surfaces without spiky, fibrous projections or ridges, drying brown, with
faint to pronounced, short, transverse ridges, not filiform and not narrowed between the flower pits; flower
pits spirally arranged, glabrous internally; proximal lips without a central notch before anthesis, not recurved
after anthesis, hood-shaped at anthesis, sometimes splitting post-anthesis; proximal and distal lips drying the
51
lobed at the apex, those of non-fertilized flowers not projecting and persistent after anthesis; fruits 7.3(6.4
7.7) mm long, 5.4(4.65.9) mm in diameter, the bases without a prominent stipe, the apices not conical, the
surfaces not splitting at maturity, with fibers emerging, bumpy from the numerous, subepidermal, tangential,
short fibers present, these coming to a point at fruit apices; locular epidermis with operculum, smooth, without
pores.
Taxonomic notes:Closely related to G. concinna (which see), differing from that species in its fruit
surfaces with emerging fibers.
geographic discontinuity, and specimens come from three different areas in Panama; the eastern end of the
Central Cordillera; the mountain systems east of the Canal Zone; and the western end of the Serrana de San
Blas, with an outlier on the Serrana de Maj. This gives three geographic subgroups, excluding the outlier.
ANOVA shows that for pair wise comparison probabilities, 10 variables (plant height, stem diameter, sheath
length, number of pinnae, apical pinna length, peduncle length, peduncle width, rachilla length, rachilla width,
number of rachillae) differ significantly (P <0.05) between one pair of subgroups, and one variable (rachis
length) differs amongst all three groups. Based on these results, the three Panamanian subgroups are
recognized as subspecies (subspp. concinnoidea, coclensis, jefensis).
Key to the subspecies of G. concinnoidea
1
2
-
Rachis 28.5(20.037.0) cm long; western end of the Serrana de San Blas, with an outlier on the Serrana de Maj...
..................................................................................................................................................... subsp. concinnoidea.
Rachis 17.5(9.627.0) cm long; all other areas ........................................................................................................... 2
Rachillae 1.0(0.81.2) mm in diameter; eastern end of the Central Cordillera.................................... subsp. coclensis
Rachillae 1.4(1.11.7) mm in diameter; east of the Canal Zone (Cerro Azul, Cerro Brewster, Cerro Bruja, Cerro
Jefe) ......................................................................................................................................................... subsp. jefensis
12a. Geonoma concinnoidea subsp. concinnoidea

Leaves rachis 28.5(20.037.0) cm long; apical pinna 16.5(12.320.5) cm long. Inflorescences rachillae
1.6(1.31.9) mm in diameter.
Distribution and habitat:From 850923N and 78257908W in Panama on western end of the
Serrana de San Blas, with an outlier on the Serrana de Maj, at 401(350550) m elevation in lowland
rainforest (Fig. 12).
12b. Geonoma concinnoidea subsp. coclensis Henderson, subsp. nov. (Appendix IV, Plate 8)
A subspeciebus aliis rachide breviore et rhachillis tenuioribus differt.
Type: PANAMA. Cocl: El Cop, 840N 8035W, 724 m, 9 May 1999, A. Henderson & E. Ferreira 3028 (holotype
PMA!, isotype NY!).
Leaves rachis 19.9(14.527.0) cm long; apical pinna 12.1(9.016.2) cm long. Inflorescences rachillae 1.0(0.8
1.2) mm in diameter.
Distribution and habitat:From 835845N and 80258041W in Panama on the eastern end of
the Central Cordillera at 737(500900) m elevation in lowland rainforest (Fig. 12).
12c. Geonoma concinnoidea subsp. jefensis Henderson, subsp. nov. (Appendix IV, Plate 9)
A subspeciebus aliis rachide breviore et rhachillis crassis differt.
Type: PANAMA. Panama: Cerro Jefe, ca. 1000 m, 25 August 1975, S. Mori & L. Joly 7933 (holotype MO!).
52
HENDERSON
Leaves rachis 15.0(9.622.5) cm long; apical pinna 15.0(11.418.5) cm long. Inflorescences rachillae 1.4(1.1
1.7) mm in diameter.
Distribution and habitat:From 910928N and 79167933W in Panama on mountains east of
the Canal Zone (Cerro Azul, Cerro Brewster, Cerro Bruja, Cerro Jefe) at 895(8001000) m elevation in
lowland rainforest (Fig. 12).
Subspecific variation:Most specimens from Cerro Jefe have pinnate leaves.
13. Geonoma congesta Wendland ex Spruce (1871: 112). Type: COSTA RICA. Heredia: Ro Sarapiqu,
between Pedegral and San Miguel, 1857, H. Wendland s.n. (holotype K!).
15.2) mm in diameter; veins raised and rectangular in cross-section adaxially; pinnae 4(124) per side of
rachis; basal pinna 59.2(33.082.0) cm long, 18.0(4.535.5) cm wide, forming an angle of 25(1538) with
the rachis; apical pinna 34.9(21.748.0) cm long, 20.7(7.142.0) cm wide, forming an angle of 26(2032)
unbranched fibers, flattened, deciduous or persistent; prophylls 15.9(6.029.7) cm long, not short and
asymmetrically apiculate, the surfaces ridged with close, equal, parallel, non-dividing ridges, scarcely
tomentose between the ridges, without unequally wide ridges; peduncular bracts 16.5(13.026.5) cm long,
well-developed, inserted 0.6(0.31.0) cm above the prophyll; peduncles 7.8(4.213.0) cm long, 6.2(3.49.8)
mm in diameter; rachillae 9(316), 12.8(6.023.0) cm long, 5.8(4.17.2) mm in diameter, the surfaces without
spiky, fibrous projections or ridges, drying brown or yellow-brown, without short, transverse ridges, not
filiform and not narrowed between the flower pits; flower pits spirally arranged, glabrous internally; proximal
lips without a central notch before anthesis, not recurved after anthesis, not hood-shaped; proximal and distal
lips drying the same color as the rachillae, not joined to form a raised cupule, the proximal lip margins
overlapping the distal lip margins; distal lips well-developed; staminate and pistillate petals not emergent, not
and curled over at anthesis; non-fertilized pistillate flowers persistent after anthesis; staminodial tubes
crenulate at the apex, those of non-fertilized flowers projecting and persistent after anthesis; fruits 12.4(9.7
15.2) mm long, 9.8(8.111.7) mm in diameter, the bases with a prominent, asymmetric stipe, the apices not
conical, the surfaces not splitting at maturity, with fibers emerging, not bumpy, not apiculate; locular
epidermis with operculum, smooth, with pores.
Taxonomic notes:Henderson et al. (1995) included Geonoma calyptrogynoidea as a synonym of G.
congestasee notes under that species.
Subspecific variation:No trait apart from stem branching and leaf division varies within this species.
There is geographic discontinuity and there is an isolated population on the Pacific slope of Costa Rica (Osa
Peninsula and adjacent areas). There are thus two potential subgroups.
Specimens from Osa differ significantly from other specimens in 10 variables (plant height, rachis length,
rachis width, number of pinnae, apical pinna length, peduncle width, rachillae length, rachillae width, number
of rachillae, fruit diameter)(t-test, P <0.05). Specimens from Osa have larger mean values for all these
variables. Based on these results, and geographic discontinuity, the two subgroups are recognized as
subspecies (subspp. congesta, osensis).
53
Key to the subspecies of G. congesta

1
-
Rachillae 11.9(6.021.5) cm long; Honduras, Nicaragua, Costa Rica (excluding Osa Peninsula and adjacent areas),
and western and central Panama ............................................................................................................subsp. congesta
Rachillae 19.1(15.023.0) cm long; Osa Peninsula and adjacent areas on the Pacific slope in Costa Rica ..................
................................................................................................................................................................. subsp. osensis
13a. Geonoma congesta subsp. congesta

Inflorescences rachillae 11.9(6.021.5) cm long.
Distribution and habitat:From 8301542N and 79458534W in Central America in
Honduras, Nicaragua, Costa Rica, and western and central Panama as far east as the Canal Zone at 255(25
1000) m elevation in lowland tropical rainforest (Fig. 13).
FIGURE 13. Distribution maps of Geonoma congesta subsp. congesta, G. congesta subsp. osensis, G. cuneata subsp.
cuneata, G. cuneata subsp. guanacastensis.
13b. Geonoma congesta subsp. osensis Henderson, subsp. nov. (Appendix IV, Plates 1015)
A Geonoma congesta subsp. congesta rhachillis longioribus differt.
Type: COSTA RICA. Puntarenas: Esquinas Forest Preserve between Palmar Sur and Golfito on United Fruit
Company railroad, 9 March 1953, H. Moore 6534 (holotype NY!, isotype BH!).
54
HENDERSON
Inflorescences rachillae 19.1(15.023.0) cm long.

Distribution and habitat:From 831917N and 83178346W in the Osa Peninsula and adjacent
areas on the Pacific slope in Costa Rica at 263(97700) m elevation in lowland tropical rainforest (Fig. 13).
14. Geonoma cuneata Wendland ex Spruce (1871: 104). Type: COSTA RICA. Heredia: Sarapiqu, 1857, H.
Wendland s.n. (holotype K!).
cane-like or cane-like; internodes 0.9(0.22.2) cm long, yellowish and smooth, or, if short and congested, not
scaly. Leaves 10(417) per stem, undivided or irregularly pinnate, not plicate, bases of blades running
diagonally into the rachis; sheaths 20.0(9.051.0) cm long; petioles 30.7(0.2113.5) cm long, drying orangebrown, reddish-brown, or green or yellowish; rachis 46.4(12.0250.0) cm long, 4.1(1.39.0) mm in diameter;
veins raised and rectangular in cross-section adaxially or not raised or slightly raised and triangular in crosssection adaxially; pinnae 3(135) per side of rachis; basal pinna 32.7(12.568.0) cm long, 5.8(0.321.5) cm
wide, forming an angle of 31(395) with the rachis; apical pinna 22.9(8.544.0) cm long, 10.7(1.537.0) cm
wide, forming an angle of 31(1050) with the rachis. Inflorescences unbranched; prophylls and peduncular
bracts ribbed with elongate, unbranched fibers, both bracts tubular, narrow, elongate, closely sheathing the
peduncle, more or less persistent; prophylls 16.5(4.038.0) cm long, not short and asymmetrically apiculate,
the surfaces not ridged, without unequally wide ridges; peduncular bracts 32.6(16.567.0) cm long, welldeveloped, inserted 1.4(0.45.7) cm above the prophyll; peduncles 51.3(13.7117.0) cm long, 3.0(1.18.2)
mm in diameter; rachillae 1, 20.4(5.552.0) cm long, 5.4(1.910.5) mm in diameter, the surfaces without
filiform and not narrowed between the flower pits; flower pits spirally arranged, glabrous internally; proximal
lips pits with a central notch before anthesis, often the two sides of the notch overlapping, recurved after
anthesis, not hood-shaped; proximal and distal lips drying the same color as the rachillae, not joined to form a
raised cupule, the proximal lip margins overlapping the distal lip margins; distal lips well-developed;
staminate and pistillate petals not emergent, not valvate throughout; staminate flowers persistent or deciduous
flowers persistent or deciduous after anthesis; staminodial tubes crenulate or shallowly lobed at the apex,
those of non-fertilized flowers not projecting and persistent after anthesis; fruits 7.6(5.011.6) mm long,
5.4(4.46.5) mm in diameter, the bases without a prominent stipe, the apices not conical, the surfaces not
splitting at maturity, without fibers emerging, ridged from the numerous, subepidermal, meridional, elongate
fibers present, these coming to a point at fruit apices; locular epidermis without operculum, smooth or
sculpted and then usually also with a raised, meridional ridge, without pores.
Taxonomic notes:Geonoma cuneata is a member of the G. cuneata clade, along with G. brenesii, G.
epetiolata, G. hugonis, and G. monospatha, from which it differs in its crenulate or shallowly lobed
staminodial tubes. Geonoma cuneata is very variable; in fact it is the fourth most variable species in the
genus. Borchsenius (1999) studied variation within G. cuneata in western Ecuador using morphometric
methods and data taken from living plants. At a local scale he found that four different varieties (based on
Henderson et al., 1995) of G. cuneata could be distinguished. However, when he included plants from other
sites in western Ecuador in the analysis, differences between the varieties broke down. Borchsenius concluded
that the varietal classification of Henderson et al. was not applicable in western Ecuador, much less
throughout the whole range of the species. Borchsenius study is of interest because of its quantitative
approach. However, he used only quantitative variables and not qualitative traits. Of the four varieties
recognized by Borchseniusvar. cuneata (here as ecuador morphotype), var. gracilis (here as esmeraldas
morphotype), var. procumbens (here as multipinnate morphotype), and var. sodiroi (here as subsp. sodiroi)
one, the last, can be recognized as a subspecies based on both quantitative variables and qualitative traits. In
55
the present study, quantitative variables and qualitative traits, as well as geographic distributions are used and
allow for separation into subspecies, as explained below.
Subspecific variation:Eight traits vary within this species (stem branching, stem type, leaf division,
petiole color, adaxial veins, staminate flower persistence, pistillate flower persistence, locular epidermis
sculpting). Excluding those traits with few data (stem branching, stem type, locular epidermis sculpting), the
remaining five traits (leaf division, petiole color, adaxial veins, staminate flower persistence, pistillate flower
persistence,) divide the specimens into seven subgroups. None of these are geographically separate. Geonoma
cuneata has a widespread and almost continuous distribution from Nicaragua through Central America and
western Colombia to western Ecuador. In some cases there are too few specimens for quantitative analysis,
and each subgroup is considered separately.
The first subgroup, with undivided leaves, petioles drying reddish-brown, and raised adaxial veins, is
confined to a small area of the Pacific coast of Colombia, in Valle, and is recognized as a subspecies (subsp.
rubra).
The second subgroup, with undivided leaves, petioles drying orange-brown, and non-raised adaxial veins,
is confined to a small area in northern Costa Rica, on the Cordilleras de Guanacaste and Tilarn, and is
recognized as a subspecies (subsp. guanacastensis).
The third subgroup, with pinnate leaves, petioles drying green, and raised adaxial veins, occurs in western
Colombia and Ecuador and is reported to be a rheophyte, growing along the banks of fast-flowing rivers, and
this is recognized as a subspecies (subsp. linearis).
The fourth subgroup, with pinnate leaves, petioles drying green, and non-raised adaxial veins, is
widespread, from Nicaragua to Ecuador. This subgroup can be separated geographically and morphologically
into specimens from western Ecuador with 57 pinnae per side of the rachis, and specimens from Nicaragua,
Costa Rica, and Panama with 628 pinnae per side. These two are recognized as subspecies (subspp. sodiroi,
procumbens, respectively).
The fifth subgroup, with undivided leaves, petioles drying green, and non-raised adaxial veins is also
widespread. This subgroup can be separated based on morphology, and a lesser extent geography, into
specimens from central Panama (El Cop, Coclecito Road, El Valle) with small leaves with the rachis 19.0
29.0 cm long, and specimens from central Panama (Santa F to the western end of the Serrana de San Bls)
with large leaves with the rachis 40.088.0 cm long. These two are recognized as subspecies (subspp. minor,
indivisa, respectively).
The sixth subgroup, with both undivided and pinnate leaves, petioles drying green, and raised adaxial
veins, is widespread from Nicaragua to Ecuador. It consists of numerous local morphotypes, as discussed
below, and cannot be divided into subspecies. It is recognized as a subspecies (subsp. cuneata).
The seventh subgroup, with pinnate leaves and raised veins, also has persistent staminate and pistillate
flowers. It occurs in western Ecuador and is recognized as a subspecies (subsp. irena).
Key to the subspecies of G. cuneata
1
2
3
4
5
6
7
Staminate and non-fertilized pistillate flowers persistent after anthesis; western Ecuador.........................subsp. irena
Staminate and non-fertilized pistillate flowers deciduous after anthesis; widespread including western Ecuador ..... 2
Petioles drying orange-brown or reddish-brown .......................................................................................................... 3
Petioles drying green or yellowish................................................................................................................................ 4
Petioles drying reddish-brown; Colombia (Valle) ......................................................................................subsp. rubra
Petioles drying orange-brown; Costa Rica (Cordilleras de Guanacaste and Tilarn) ................ subsp. guanacastensis
Leaves with the veins raised and rectangular in cross-section adaxially...................................................................... 5
Leaves with the veins not raised or slightly raised and triangular in cross-section adaxially .................................... 6
Rheophytes; western Colombia and Ecuador ..........................................................................................subsp. linearis
Non-rheophytes; widespread .................................................................................................................. subsp. cuneata
Pinnae 6(57) per side of rachis; western Ecuador...................................................................................subsp. sodiroi
Leaves undivided or pinnate; Nicaragua, Costa Rica, Panama .................................................................................... 7
Pinnae 12(128) per side of rachis; Nicaragua, Costa Rica, Panama...............................................subsp. procumbens
56
HENDERSON
8
-
Leaves undivided; Panama .......................................................................................................................................... 8

Rachis 22.6(19.029.0) cm long; El Cop, Coclecito Road, El Valle ....................................................... subsp. minor
Rachis 56.6(40.088.0) cm long; Santa F to the western end of the Serrana de San Bls .................. subsp. indivisa
14a. Geonoma cuneata subsp. cuneata (Plates V & VI)

Geonoma obovata Wendland ex Spruce (1871: 104). Type: COSTA RICA. Heredia: Sarapiqu, near Pedregal, August
1857, H. Wendland s. n. (holotype K!).
Geonoma gracilis Wendland ex Spruce (1871: 105). Geonoma cuneata var. gracilis (Wendland ex Spruce) Skov ex
Govaerts & Dransfield (2005: 114). Type: COSTA RICA. Heredia: Sarapiqu, August 1857, H. Wendland s.n.
(holotype K!).
Geonoma cuneatoidea Burret (1930a: 167). Type: COLOMBIA. Antioquia: Murr, ca. 1000 m, 21 July 1880, W.
Kalbreyer 1828 (holotype B, destroyed). Neotype (selected by Bernal et al. 1989): COLOMBIA. Antioquia: Mun.
Frontino, Corregimiento de Murr, La Blanquita, 815 m, 22 March 1982, R. Bernal & G. Galeano 286 (neotype
COL!).
Leaves undivided or pinnate; petioles drying green or yellowish; veins raised and rectangular in cross-section
adaxially; rachis 45.1(12.0250.0) cm long; pinnae 3(135) per side of rachis. Inflorescences staminate
flowers deciduous after anthesis; non-fertilized pistillate flowers deciduous after anthesis.
Distribution and habitat:From 1218N148S and 71208523W in Nicaragua, Costa Rica,
Panama, Colombia, Venezuela, and Ecuador at 541(21750) m elevation in lowland or montane rainforest
(Fig. 13).
Geonoma cuneata subsp. cuneata is made up of a series of slightly differing morphotypes. There are a few
specimens that do not fit into any of these morphotypes. These are usually single specimens from isolated
areas.
Specimens from the Atlantic slope in Costa Rica (atlantic morphotype), a few specimens just reaching the
Pacific slope, and a few from adjacent Panama have undivided or pinnate leaves with raised adaxial veins.
When pinnate, the apical pinna is usually wide. The types of G. cuneata, G. gracilis, and G. obovata come
from this area. Regression shows there are significant associations between elevation and three leaf variables.
Squared multiple R for the regression of rachis length on elevation is 0.32, basal pinna angle 0.38, and apical
pinna angle 0.40. Specimens from higher elevations tend to have shorter rachis and wider basal and apical
angles.
There are two unplaced specimens from this region. In Nicaragua there is a single specimen (Nee 28420)
from cloud forest in Chontales. To the south of this, at a lower elevation, is another single specimen (Stevens
8967), with a undivided leaf with a narrow basal angle. Both these are separated from other Atlantic slope
specimens in Costa Rica by a large population of subsp. procumbens.
On the Pacific slope (pacific morphotype), specimens occur only in Costa Rica, and are concentrated in
the Osa Peninsula region. They are similar to Atlantic slope specimens in their undivided or pinnate leaves
with raised adaxial veins. They differ in their shorter plant height, shorter leaf sheaths, shorter and thinner
rachis, more pinnae, wider basal pinna angles, shorter and thinner apical pinnae, thinner peduncles, and
shorter rachillae.
There are three unplaced specimens from this region. Two specimens (Grayum 5960, Jimnez 891) from
the Montaas Jamaica have smaller, undivided leaves with less pronounced raised veins. One specimen
(Davidse 26231) from the Cordillera de Talamanca has orange-brown petioles and rachis.
In western Panama in the Fortuna area (fortuna morphotype), in Chiriqu and Bocas del Toro, there is a
homogeneous morphotype from higher elevations (1180 m mean elevation), compared with 496 m mean
elevation for Atlantic and Pacific slope morphotypes. All specimens except one have undivided leaves with
raised adaxial veins, and small inflorescences. Specimens differ from the nearest others, the Atlantic slope
morphotype in Costa Rica, in their shorter petioles, shorter and narrower rachis, narrower basal pinna angles,
narrower peduncles, and shorter and thinner rachillae.
There is a similar morphotype from the Santa F region in Panama (santafe morphotype). Specimens have
undivided or pinnate leaves with raised adaxial veins. They differ from Fortuna specimens in their longer
57
petioles, more pinnae, wider basal pinna angles, and shorter apical length. About half the specimens have
pinnate leaves, which accounts for their wider basal pinna angles. This morphotype also occurs at lower mean
elevations than the fortuna morphotype (888 versus 1140) m.
At the eastern end of the Central Cordillera at El Cop and the Coclecito Road (elcope morphotype) there
is a morphotype with smaller, pinnate leaves and a slender, short rachilla. Specimens differ from Santa F
ones in nine variables, particularly in their smaller leaves.
At El Valle and El Cop (elvalle morphotype) there is a morphotype with larger, undivided or pinnate
leaves with pronounced adaxial veins and long, stout rachillae.
Specimens from Ro Guanche, Santa Rita Ridge, Cerro Bruja, and Serrana de San Bls (guanche
morphotype) are particularly variable, especially in rachilla size. They have undivided or pinnate leaves with
pronounced adaxial veins. Some specimens have leaves which dry a gray-green color.
Some specimens from the Serrana de San Bls (sanblas morphotype) have small, usually undivided
leaves with raised veins and small inflorescences.
In northwestern Colombia and just reaching eastern Panama, with an outlier on the Sierra Nevada de
Santa Marta there is a morphotype (cuneatoidea morphotype) with large, undivided leaves or with 28 pinnae
per side with pronounced raised veins and large inflorescences. The type of G. cuneatoidea is of this
morphotype.
In the northern half of the Choc region, and extending into eastern Panama, the Magdalena valley, the
Central and Western Cordilleras, and western Venezuela is a morphotype (choco morphotype) with leaves
with 37 pinnae per side, rarely undivided, and raised adaxial veins. Two specimens (Bernal 2174, Juncosa
1228) from this area are unplaced. They are similar to subsp. indivisa except they have orange-brown petioles
and rachis.
Stauffer (1998) reported that G. cuneata occurred in Apure, Venezuela. The specimens cited by Stauffer
have not been seen, but another specimen from the same locality is here determined as G. brongniartii.
Geonoma cuneata does, however occur in Zulia, Venezuela in its cuneatoidea morphotype.
In northwestern Colombia, southwestern Colombia, and northwestern Ecuador (multipinnate
morphotype), there is a pinnate leaved morphotype with 635 pinnae per side of the rachis, raised adaxial
veins, and large inflorescences. There is no significant difference in any variable between the two areas where
this morphotype occurs.
In western Ecuador (Esmeraldas) and extreme southwestern Colombia (esmeraldas morphotype) at low
elevations (50350 m), a few specimens, have small, undivided leaves with raised adaxial veins.
In western Ecuador and just reaching southwestern Colombia (ecuador morphotype) at higher elevations
(2001375 m) there is a morphotype with large, undivided or divided leaves with 26 pinnae per side of the
rachis and prominent raised adaxial veins. This morphotype differs from the similar cuneatoidea morphotype
in its narrower rachis and shorter rachillae. The specimen from the most southerly location (Jativa231) is
smaller than the others.
14b. Geonoma cuneata subsp. guanacastensis Henderson, subsp. nov. (Appendix IV, Plate 16)
A subspeciebus aliis petiolis in sicco brunneoaurantiacis differt.
Type: COSTA RICA. Guanacaste/Alajuela: slopes of Miravalles, above Bijagua, ca. 1500 m, November 1982, L. Gmez
et al. 19053 (holotype NY!, isotype MO!).
Leaves undivided; petioles drying orange-brown; veins not raised or slightly raised and triangular in crosssection adaxially; rachis 27.6(17.731.5) cm long; pinnae 1 per side of rachis. Inflorescences staminate
Distribution and habitat:From 1035N1059N and 84558527W in Costa Rica (Cordilleras de
Guanacaste and Tilarn) at 814(4701500) m elevation in lowland or montane rainforest (Fig. 13).
58
HENDERSON
14c. Geonoma cuneata subsp. indivisa Henderson, subsp. nov. (Appendix IV, Plate 17)
A subspeciebus aliis foliis simplicibus venis haud prominentibus atque rachide longiore differt
Type: PANAMA. San Blas: Cerro Brewster, 918N, 7916W, 800850 m, 20 November 1985, G. de Nevers, A.
Henderson, H. Herrera, G. McPherson & L. Brako 6293 (holotype NY!, isotype MO!).
Leaves undivided; petioles drying green or yellowish; veins not raised or slightly raised and triangular in
cross-section adaxially; rachis 56.6(40.088.0) cm long; pinnae 1 per side of rachis. Inflorescences staminate
Distribution and habitat:From 834900N and 78558107W in central Panama (Santa F to
the western end of the Serrana de San Bls) at 541(501000) m elevation in lowland rainforest (Fig. 14).
FIGURE 14. Distribution maps of Geonoma cuneata subsp. indivisa, G. cuneata subsp. irena, G. cuneata subsp.
linearis, G. cuneata subsp. minor.
14d. Geonoma cuneata subsp. irena (Borchsenius) Henderson, comb. & stat. nov.
Basionym: Geonoma irena Borchsenius (1996: 605). Type. ECUADOR. Pichincha: Hacienda Irena, km 35 on the road
Sto. Domingo de los Colorados-Quevedo, 280) m, 16 September 1995, F. Borchsenius, C. Asmussen & J. Knudsen
286 (holotype AAU!, isotypes COL!, F!, NY!).
Leaves pinnate; petioles drying green or yellowish; veins raised and rectangular in cross-section adaxially;
rachis 68.6(36.0105.0) cm long; pinnae 67 per side of rachis. Inflorescences staminate flowers persistent
after anthesis; non-fertilized pistillate flowers persistent after anthesis.
59
Distribution and habitat:From 103N028S and 78557942W in western Ecuador at 253(185

300) m elevation in lowland rainforest (Fig. 14).
14e. Geonoma cuneata subsp. linearis (Burret) Henderson, comb. & stat. nov.
Basionym: Geonoma linearis Burret (1933a: 861). Type: COLOMBIA. Nario: Barbacoas, Ro Telembi, 10 August
1880, F. Lehmann 51 (holotype B, destroyed). Neotype (selected by Galeano & Skov (1989): COLOMBIA. Nario:
Mun. Barbacoas, Ro Telemb, entre Barbacoas y ca. 15 km ro arriba, 160) m, 20 November 1986, R. Bernal & B.
Hammel 1320 (neotype COL!, isoneotype AAU!).
Leaves pinnate; petioles drying green or yellowish; veins raised and rectangular in cross-section adaxially;
rachis 40.3(24.080.5) cm long; pinnae 7(410) per side of rachis. Inflorescences staminate flowers deciduous
after anthesis; non-fertilized pistillate flowers deciduous after anthesis.
Distribution and habitat:From 035545N and 76257856W in western Colombia and
Ecuador at 112(10300) m elevation in lowland rainforest along the banks of fast-flowing streams (Fig. 14).
14f. Geonoma cuneata subsp. minor Henderson, subsp. nov. (Appendix IV, Plate 18)
A subspeciebus aliis foliis simplicibus venis haud prominentibus atque rachide breviore differt.
Type: PANAMA. Cocl: El Valle de Antn, Cerro Gaital, 837N, 8006W, 26 November 1985, G. de Nevers, A.
Henderson, H. Herrera, G. McPherson & L. Brako 6355 (holotype NY!, isotype MO!).
Leaves undivided; petioles drying green or yellowish; veins not raised or slightly raised and triangular in
cross-section adaxially; rachis 22.6(19.029.0) cm long; pinnae 1 per side of rachis. Inflorescences staminate
Distribution and habitat:From 837842N and 80058039W in central Panama (El Cop,
Coclecito Road, El Valle) at 810(7001030) m elevation in lowland rainforest (Fig. 14).
14g. Geonoma cuneata subsp. procumbens (Wendland ex Spruce) Henderson, comb. & stat. nov.
Basionym: Geonoma procumbens Wendland ex Spruce (1871: 105). Geonoma cuneata var. procumbens (Wendland ex
Spruce) Skov ex Govaerts & Dransfield (2005: 114). Type: COSTA RICA. Heredia: Sarapiqu, 1857, H. Wendland
s.n. (holotype K!).
Leaves pinnate; petioles drying green or yellowish; veins not raised or slightly raised and triangular in crosssection adaxially; rachis 71.3(42.0108.0) cm long; pinnae 12(128) per side of rachis. Inflorescences
staminate flowers deciduous after anthesis; non-fertilized pistillate flowers deciduous after anthesis.
Distribution and habitat:From 7241143N and 77408520W in Nicaragua, Costa Rica, and
Panama at 221(50700) m elevation in lowland rainforest (Fig. 15).
There are two separate populations, in Nicaragua and Costa Rica, and eastern Panama. These do not differ
from each other except for two variables (apical pinna width, rachilla widthboth wider in the Panama
population)(t-test, P <0.05).
Two specimens (Herrera 752, 973) from the same locality in eastern Panama, and from the same locality
as other specimens of subsp. procumbens, differ in their undivided leaves.
14h. Geonoma cuneata subsp. rubra Henderson, subsp. nov. (Appendix IV, Plates 19 & 20)
A subspeciebus aliis petiolis in sicco brunneo-rubris differt.
Type: COLOMBIA. Valle: Ro Yurumangu, Quebrada Querr, arribe de Venerel, 50 m, 9 February 1990, R. Bernal, W.
Devia, E. Linhares & J. Angulo 1770 (holotype COL!).
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HENDERSON
Leaves undivided; petioles drying reddish-brown; veins raised and rectangular in cross-section adaxially;
rachis 29.4(22.042.5) cm long; pinnae 1 per side of rachis. Inflorescences staminate flowers deciduous after
anthesis; non-fertilized pistillate flowers deciduous after anthesis.
Distribution and habitat:From 310359N and 76587720W in western Colombia (Valle) at
74(50120) m elevation in lowland rainforest (Fig. 15).
FIGURE 15. Distribution maps of Geonoma cuneata subsp. procumbens, G. cuneata subsp. rubra, G. cuneata subsp.
sodiroi, and G. deneversii.
14i. Geonoma cuneata subsp. sodiroi (Dammer ex Burret) Henderson, comb. & stat. nov.
Basionym: Geonoma sodiroi Dammer ex Burret (1930a: 165). Geonoma cuneata var. sodiroi (Dammer ex Burret) Skov
ex Govaerts & Dransfield (2005: 114). Type: ECUADOR. Pichincha: Santo Domingo, October 1885, L. Sodiro 187/
1 (holotype P!).
Leaves pinnate; petioles drying green or yellowish; veins not raised or slightly raised and triangular in crosssection adaxially; rachis 30.3(16.144.5) cm long; pinnae 6(57) per side of rachis. Inflorescences staminate
Distribution and habitat:From 015034S and 79097920W in western Ecuador at 475(300
The single specimen from Centinela (Dodson 14822) is smaller than the other two.
61
15. Geonoma deneversii Henderson, sp. nov. (Appendix IV, Plates 2124)
A speciebus affinibus staminibus plus quam sex differt.
Type: PANAMA. Bocas del Toro: E of Gualaca-Chiriqu Grande road, 10 mi. N of continental divide, 1 mi. along side
road E, 855N 8205W, 100500 m, 19 January 1989, G. de Nevers, F. Almeda & G. McPherson 8823 (holotype
NY!, isotypes BH!, K!, MO!).
Plant height no data; stems 1.3 m tall, solitary; internodes no data. Leaves 20 per stem, irregularly pinnate, not
plicate, bases of blades running diagonally into the rachis; sheaths no data; petioles drying green or yellowish;
rachis 104.5 cm long, 5.4 mm in diameter; veins raised and rectangular in cross-section adaxially; pinnae 3
per side of rachis; basal pinna 87.0 cm long, 18.7 cm wide, forming an angle of 31 with the rachis; apical
pinna 25.5 cm long, 27.5 cm wide, forming an angle of 27 with the rachis. Inflorescences unbranched;
elongate, closely sheathing the peduncle, more or less persistent; prophylls 35.5 cm long, not short and
asymmetrically apiculate, the surfaces not ridged, without unequally wide ridges; peduncular bracts 31.0 cm
long, well-developed, inserted 2.7 cm above the prophyll; peduncles 92.5 cm long, 5.2 mm in diameter;
rachillae 1, 31.0 cm long, 9.9 mm wide, the surfaces without spiky, fibrous projections or ridges, drying
brown or yellow-brown, without short, transverse ridges, not filiform and not narrowed between the flower
pits; flower pits spirally arranged, glabrous internally; proximal lips with a central notch before anthesis, often
the two sides of the notch overlapping, not recurved after anthesis, not hood-shaped; proximal and distal lips
drying the same color as the rachillae, not joined to form a raised cupule, the proximal lip margins overlapping
the distal lip margins; distal lips well-developed; staminate and pistillate petals not emergent, not valvate
throughout; staminate flowers deciduous after anthesis; stamens more than 6; thecae diverging at anthesis,
inserted directly onto the apiculate filament apices; anthers not short and curled at anthesis, usually elongate,
spiraled and twisted or sometimes remaining straight; non-fertilized pistillate flowers deciduous after
anthesis; staminodial tubes lobed at the apex, the lobes spreading at anthesis, acuminate; staminodial tubes
projection no data; fruits no data.
Distribution and habitat:At 855N and 8205W on the Caribbean slope in Panama (Bocas del Toro)
at 250 m elevation in lowland rainforest (Fig. 15).
Taxonomic notes:Geonoma deneversii differs from similar species (G. camana, G. chlamydostachys,
G. chococola) in its staminate flowers with more than six stamens. It is also the only species of this group to
occur in Central America.
Subspecific variation: No trait varies within this species, and only one specimen is known.
16. Geonoma deversa (Poiteau) Kunth (1841: 321). Gynestum deversum Poiteau (1822: 390). Type:
FRENCH GUIANA. Without locality, no date, A. Poiteau s.n. (holotype P!).
Plants 2.4(0.55.0) m tall; stems 2.4(0.37.0) m tall, 1.0(0.51.8) cm in diameter, solitary or clustered, canelike or not cane-like; internodes 1.9(0.57.5) cm long, yellowish and smooth. Leaves 11(618) per stem,
undivided or irregularly pinnate, sometimes regularly pinnate and the pinnae with 1 main vein only, not
plicate, bases of blades running diagonally into the rachis; sheaths 12.5(5.027.5) cm long; petioles 20.6(4.2
82.0) cm long, drying green or yellowish; rachis 42.0(17.292.5) cm long, 3.2(1.47.0) mm in diameter; veins
not raised or slightly raised and triangular in cross-section adaxially; pinnae 5(128) per side of rachis; basal
pinna 28.5(10.560.5) cm long, 6.4(0.527.0) cm wide, forming an angle of 42(2093) with the rachis;
apical pinna 19.9(8.835.5) cm long, 12.4(0.626.7) cm wide, forming an angle of 28(1445) with the rachis.
Inflorescences branched 13 orders; prophylls and peduncular bracts not ribbed with elongate, unbranched
fibers, flattened, deciduous; prophylls 6.8(3.013.0) cm long, not short and asymmetrically apiculate, the
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HENDERSON
surfaces not ridged, without unequally wide ridges; peduncular bracts 4.8(3.27.5) cm long, well-developed,
inserted 0.3(0.10.7) cm above the prophyll; peduncles 8.3(2.019.7) cm long, 4.2(1.99.0) mm in diameter;
narrowed between the flower pits; flower pits tricussately or quadricussately arranged throughout the
rachillae, the groups of pits closely spaced, glabrous internally; proximal lips without a central notch before
anthesis, not recurved after anthesis, hood-shaped at anthesis, sometimes splitting post-anthesis; proximal and
distal lips drying the same color as the rachillae, joined to form a raised cupule, the margins not overlapping;
flowers deciduous after anthesis; stamens 6; thecae diverging at anthesis, inserted almost directly onto the
filament apices, the connectives bifid but scarcely developed; anthers short and curled over at anthesis; nonfertilized pistillate flowers deciduous after anthesis; staminodial tubes crenulate or shallowly lobed at the
apex, those of non-fertilized flowers not projecting and persistent after anthesis; fruits 6.6(4.58.1) mm long,
5.6(4.47.0) mm in diameter, the bases without a prominent stipe, the apices not conical, the surfaces not
splitting at maturity, without fibers emerging, not bumpy and not apiculate; locular epidermis without
operculum, smooth, without pores.
Taxonomic notes:Geonoma deversa is the most wide-ranging species in the genus, occurring from
Belize and Guatemala to Brazil and Bolivia. It is variable, but not as much as its wide range would suggest
(see section on Infraspecific Variation). Its treatment here is generally consistent with that of Wessels Boer
(1968), except that several subspecies are recognized. It is recognized by its flower pits which are usually
tricussately arranged (sometimes quadricussately) and closely spaced throughout the rachillae.
Subspecific variation:Three traits (stem branching, stem type, leaf division) vary within this species.
Only two specimens (of 179) are scored as stems not cane-like and this trait, together with stem branching and
leaf division, are not useful for subspecific delimitation in this species.
There is little geographic discontinuity, except that in Central America two subgroups are geographically
isolatedin Belize and Guatemala, and the Osa Peninsula of Costa Rica. Specimens from Belize and
Guatemala differ significantly from other Central American (excluding Osa Peninsula subgroup) specimens in
14 variables (stem diameter, internode length, sheath length, petiole length, rachis width, basal pinna length,
basal pinna width, basal pinna angle, apical pinna length, apical pinna width, prophyll length, peduncular
bract length, peduncle width, number of rachillae)(t-test, P <0.05); and specimens from the Osa Peninsula of
Costa Rica differ from other Central American (excluding Belize and Guatemala) specimens in 12 variables
(stem diameter, sheath length, rachis length, rachis width, number of pinnae, apical pinna length, apical pinna
width, prophyll length, peduncle width, rachillae length, number of rachillae, fruit length). Based on these
results, these two subgroups are recognized as subspecies (subspp. belizensis, peninsularis).
There is considerable variation in the trait pit arrangement in specimens from the western Amazon region.
Some specimens from the northwestern Amazon region of Colombia, Brazil, and Peru have quadricussate
flower pits rather than the more usual tricussate ones. Because of this, the subgroup is recognized as a
subspecies (subspp. quadriflora), and all other specimens are recognized as subsp. deversa.
Key to the subspecies of G. deversa
1
2
3
4.
-
Central America............................................................................................................................................................ 2
South America .............................................................................................................................................................. 4
Belize, Guatemala................................................................................................................................ subsp. belizensis
All other areas .............................................................................................................................................................. 3
Rachis 38.9(17.280.0) cm long; widespread......................................................................................... subsp. deversa
Rachis 64.5(46.585.0) cm long; Osa Peninsula of Costa Rica ...................................................... subsp. peninsularis
Flower pits tricussately arranged; widespread........................................................................................ subsp. deversa
Flower pits quadricussately arranged; northwestern Amazon region of Colombia, Brazil, and Peru...........................
.......................................................................................................................................................... subsp. quadriflora
63
16a. Geonoma deversa subsp. deversa

Geonoma paniculigera Martius (1823: 11). Lectotype (selected by Wessels Boer 1968): BRAZIL. Par et Rio Negro,
no date, C. Martius s.n. (M!).
Geonoma longipetiolata rsted (1858: 36). Type: NICARAGUA. Ro San Juan, no date, A. Orsted s.n. (holotype C,
n.v.).
Geonoma microspatha Spruce (1871: 108). Geonoma paniculigera var. microspatha (Spruce) Trail (1876: 327). Type:
BRAZIL. Amazonas: Rio Negro, Serra do Gama, So Gabriel, February 1853, R. Spruce 28 (holotype K!, isotype
P!).
Geonoma microspatha var. pacimoensis Spruce (1871: 116). Type: BRAZIL. Amazonas: Rio Casiquiare, no date, R.
Spruce 41 (holotype K!, isotype P!).
Geonoma flaccida Wendland ex Spruce (1871: 108). Type: COSTA RICA. Heredia: Ro Sarapiqu, 1857, H. Wendland
s.n. (holotype K!).
Geonoma paniculigera var. papyracea Trail (1876: 326). Type: BRAZIL. Amazonas: Rio Javari, 8 December 1874, J.
Trail 943/CXCIII (holotype K!).
Geonoma paniculigera var. cosmiophylla Trail (1876: 326). Type: BRAZIL. Amazonas: Tabatinga, 30 November 1874,
J. Trail 956/CLXXVII (holotype K!, isotype P!).
Geonoma paniculigera var. cosmiophylla subvar. gramineifolia Trail (1876: 327). Type: BRAZIL. Amazonas: Tabatinga,
30 November 1874, J. Trail 958/CLXXVII (holotype K!, isotype P!).
Geonoma myriantha Dammer (1907: 120). Type: BRAZIL. Acre: Rio Juru, Juru-mirim, September 1901, E. Ule 5882
(holotype B, destroyed, isotype MG!), synon. nov.
Geonoma leptostachys Burret (1930b: 1014). Type: BRAZIL. Amazonas: Rio Negro, Camanaos, 26 September 1928, P.
Luetzelburg 23072 (holotype B, destroyed, isotype M!).
Geonoma macropoda Burret (1930b: 1015). Type: BRAZIL. Amazonas: Manaus, 26 August 1928, P. Luetzelburg 22089
(holotype B, destroyed, isotypes M!, R!).
Geonoma major Burret (1930b: 1016). Type: BRAZIL. Amazonas: Rio Negro, Serra do Cucui, 25 September 1928, P.
Luetzelburg 22273 (holotype B, destroyed, isotypes M!, R!).
Geonoma killipii Burret (1932a: 320). Type: PERU. Junn: Puerto Bermudez, ca. 375 m, 1417 July 1929, E. Killip & A.
Smith 26594 (holotype B, destroyed, isotypes F!, NY!, US!).
Leaf rachis 38.9(17.280.0) cm long. Inflorescences peduncles 3.8(1.97.7) mm in diameter; rachillae

Distribution and habitat:From 1507N1700S and 49318527W in Central and South America
at 246(51200) m elevation in lowland to montane rainforest (Fig. 16).
There is a single specimen (Henderson 3024) from Herrera, Panama, isolated from others, which
resembles specimens of subsp. peninsularis, particularly in its short peduncle and long rachillae, and it may
represent a distinct subspecies.
Specimens from central Panama, from the Santa Rita Ridge to western San Blas have different shaped
leaves, with shorter basal and apical pinnae with wider angles. Such specimens are also found just to the
northeast of the Osa Peninsula, sympatric with subsp. peninsularis.
Several specimens, particularly from white sand regions of the Rio Negro in Venezuela and Brazil, but
also less commonly from other areas, have regularly pinnate leaves with numerous pinnae with 1 main vein
only. Some of these are reported to be rheophytes.
A few specimens, forming an isolated population in Pasco, Peru, are considerably smaller than others.
They have wider basal pinna angles and sigmoid pinnae. The type of G. killipii is one of these specimens, and
they are recognized as the killipii morphotype. Potential hybrids between this morphotype and G. brongniartii
are discussed under that species.
There are several other areas where hybrids are suspected with other taxa. All these potential hybrids have
been excluded from the above descriptions and analyses.
Specimens from the northeastern part of the range of the subspecies, in Suriname, French Guiana, and
Brazil (Amap, Par), are intermediate in morphology between that of subsp. deversa and the sympatric G.
leptospadix. Some of these specimens have leaves like those of subsp. deversa and triad arrangement
approaching that of G. leptospadix; some have leaves of G. leptospadix and triad arrangement of subsp.
deversa. These specimens are hypothesized to be hybrids between the two taxa. There are a few other
64
HENDERSON
specimens from this region which also appear somewhat intermediate, but are determined as subsp. deversa.
A hybrid zone may exist running from Suriname and French Guiana to Brazil (Amap, Par)(see Fig. 23).
There is considerable variation in the western Amazon region, and there may also be hybrid zones here,
especially in Peru (Loreto) and Brazil (Acre). Specimens from near Iquitos (iquitos morphotype) differ from
other specimens of subsp. deversa in 13 variables (plant height, stem diameter, petiole length, rachis length,
rachis width, basal pinna length, interbract distance, peduncle length, peduncle width, rachilla length, rachilla
width, number of rachillae, fruit diameter)(t-test, P <0.05). On the other hand, the same specimens differ from
the sympatric subsp. quadriflora in only four variables (peduncle width, rachilla width, fruit length, fruit
diameter) (t-test, P <0.05). These results indicate that the iquitos morphotype may represent hybrids between
subsp. deversa and subsp. quadriflora (specimens of subsp. quadriflora may have tricussate pits at the apices
of the rachillae, and one specimen of subsp. deversa (Croat 20340) contains two inflorescences, one with
quadricussate pits and the other with tricussate pits).
Some specimens from the western Amazon region of Brazil (Acre, Amazonas) differ from the more
typical subsp. deversa in their longer leaves with more pinnae, and shorter, thicker peduncles. In their leaves
they resemble subsp. quadriflora and in their inflorescences they resemble G. occidentalis, and they occur in
an area between the ranges of these two taxa. They may be hybrids between subsp. quadriflora and G.
occidentalis, or between one of these taxa and subsp. deversa. The type specimen of G. myriantha appears to
represent one of these possible hybrids.
16b. Geonoma deversa subsp. belizensis Henderson, subsp. nov. (Appendix IV, Plate 25)
A Geonoma deversa subsp. deversa pedunculo latiore differt.
Type: BELIZE. Stann Creek District: Cockscomb Basin, Jaguar Preserve, 10 km W of Maya Center, off Southern
Highway, 1645N, 8835W, 400 m, 23 May 1990, M. Balick, R. Arvigo, P. Cocom, R. Cocom, H. Robinson & G.
Shropshire 2698 (holotype NY!).

Distribution and habitat:From 1535N1710N and 88248901W in Belize and Guatemala at
300(100400) m elevation in lowland rainforest (Fig. 16).
16c. Geonoma deversa subsp. peninsularis Henderson, subsp. nov. (Appendix IV, Plate 26)
A Geonoma deversa subsp. deversa rachide longiore atque pedunculo latiore differt.
Type: COSTA RICA. Puntarenas: Reserva Forestal Golfo Dulce, Cantn de Osa, Rancho Quemado, ca. 15 km W of
Rincn, 842N, 8333W, 250 m, 6 June 1992, A. Henderson, G. Galeano & B. Hammel 1817 (holotype CR!,
isotype NY!).

Distribution and habitat:From 827858N and 83148340W in the Osa Peninsula and adjacent
areas of Costa Rica at 287(65744) m elevation in lowland rainforest (Fig. 16).
There is a sympatric population of subsp. deversa occurring just to the northeast of the Osa Peninsula.
16d. Geonoma deversa subsp. quadriflora Henderson, subsp. nov. (Appendix IV, Plates 2730)
A Geonoma deversa subsp. deversa foveis floralibus quarternatis differt.
65
Type: COLOMBIA. Amazonas: road from Leticia to Tarapac, ca. 7 km N of Leticia, 180 m, 17 March 1990, G.
Galeano, R. Bernal, A. Henderson & S. Churchill 2112 (holotype COL!, isotype NY!).

Colombia, Brazil, and Peru at 137(95180) m elevation in lowland rainforest (Fig. 16).
The outlying specimens from Brazil probably appear isolated only because the intervening area is poorly
collected. See discussion under G. deversa subsp. deversa for possible hybrids.
FIGURE 16. Distribution maps of Geonoma deversa subsp. deversa, G. deversa subsp. belizensis, G. deversa subsp.
peninsularis, and G. deversa subsp. quadriflora.
17. Geonoma dindoensis Henderson, sp. nov. (Appendix IV, Plates 31 & 32)
A speciebus affinibus prophyllis haud brevibus necnon inaequaliter apiculatis atque rachide longiore differt.
Type: COLOMBIA. Valle: Dindo area, Bajo Calima, 359N, 7658W, 100 m, 20 July 1984, A. Gentry & M. Monsalve
48419 (holotype NY!, isotype MO!).
Plants 2.0 m tall; stems height no data, 0.6 cm in diameter, cane-like; internodes 2.5 cm long, yellowish and
smooth. Leaves undivided, not plicate, bases of blades running diagonally into the rachis; sheaths 6.5 cm long;
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HENDERSON
petioles 6.5 cm long, drying green or yellowish; rachis 27.0 cm long, 2.6 mm in diameter; veins not raised or
slightly raised and triangular in cross-section adaxially; pinnae 1 per side of rachis; basal pinna forming an
angle of 22 with the rachis; apical pinna 9.5 cm long, forming an angle of 26 with the rachis. Inflorescences
deciduous or persistent; prophylls length no data, not short and asymmetrically apiculate, the surfaces not
ridged, without unequally wide ridges; peduncular bracts length no data, well-developed, inserted 1.3 cm
above the prophyll; peduncle 5.7 cm long, 2.3 mm in diameter; rachillae 18, 4.2 cm long, 0.9 mm in diameter,
the surfaces without spiky, fibrous projections or ridges, drying brown, with faint to pronounced, short,
transverse ridges, filiform with extended narrowed sections between the flower pits; flower pits alternately
arranged (sometimes distorted by twisting and contracting of rachillae), glabrous internally; proximal lips
without a central notch before anthesis, not recurved after anthesis, not hood-shaped; proximal and distal lips
drying the same color as the rachillae, joined to form a raised cupule, the margins not overlapping; distal lips
well-developed; staminate and pistillate petals not emergent, not valvate throughout; staminate flowers
deciduous after anthesis; stamens 6; thecae diverging at anthesis, inserted almost directly onto the filament
apices, the connectives bifid but scarcely developed; anthers short and curled over at anthesis; non-fertilized
pistillate flowers deciduous after anthesis; staminodial tubes crenulate or shallowly lobed at the apex, those of
non-fertilized flowers not projecting and persistent after anthesis; fruits no data.
Distribution and habitat:At 359S and 7658W on the Pacific coast of Colombia (Valle) at 100 m
FIGURE 17. Distribution maps of Geonoma dindoensis, G. divisa, G. elegans, and G. epetiolata.
67
Taxonomic notes:Geonoma dindoensis shares the same character states as G. bernalii, except that
fruits are lacking in the single specimen known. Given the differences in rachis length (27.0 cm long versus
13.3(12.514.1) cm long) and habitat (lowland rainforest in the Choc at 100 m versus montane rainforest on
eastern Andean slopes at 1105(10101200) m, G. dindoensis is recognized as distinct from G. bernalii,
pending more collections.
Subspecific variation: No trait varies within this species and only one specimen is known.
18. Geonoma divisa Moore (1980: 25). Type: COLOMBIA. Choc: N ridge of Alto de Buey, above Dos
Bocas del Ro Mutat, tributary of Ro El Valle, ESE of El Valle, 200500 m, 8 August 1976, A. Gentry & M.
Fallen 17438 (holotype BH!, isotypes COL!, MO!).
Plants 2.8(2.53.0) m tall; stems 2.8(2.03.5) m tall, 0.8(0.61.1) cm in diameter, clustered, cane-like;
internodes 1.9(0.95.4) cm long, yellowish and smooth. Leaves 8(610) per stem, undivided, not plicate,
bases of blades running diagonally into the rachis; sheaths 7.9(4.511.0) cm long; petioles 9.2(6.015.0) cm
long, drying green or yellowish; rachis 21.5(8.831.0) cm long, 3.0(2.34.4) mm in diameter; veins raised and
rectangular in cross-section adaxially; pinnae 1 per side of rachis; basal pinna forming an angle of 28(1238)
with the rachis; apical pinna 27.3(21.234.5) cm long, forming an angle of 20(1530) with the rachis.
Inflorescences branched 1 order; prophylls and peduncular bracts not ribbed with elongate, unbranched fibers,
flattened, deciduous; prophylls 6.7(4.98.1) cm long, short, asymmetrically apiculate, the margins curved
around the stem, the surfaces flat with dense, felty, brown tomentum, prophyll equal to and early deciduous
with the peduncular bract, the surfaces not ridged, without unequally wide ridges; peduncular bracts 5.1(3.5
6.7) cm long, well-developed, inserted 0.2(0.10.3) cm above the prophyll; peduncles 4.3(2.87.3) cm long,
ridges, not filiform and not narrowed between the flower pits; flower pits tricussately arranged throughout the
rachillae, the groups of pits closely spaced, glabrous internally; proximal lips without a central notch before
anthesis, not recurved after anthesis, not hood-shaped; proximal and distal lips drying the same color as the
pistillate flowers deciduous after anthesis; staminodial tubes crenulate or shallowly lobed at the apex, those of
non-fertilized flowers not projecting and persistent after anthesis; fruits 7.3 mm long, 6.5 mm in diameter, the
bases without a prominent stipe, the apices not conical, the surfaces not splitting at maturity, without fibers
emerging, ridged from the numerous, subepidermal, meridional, elongate fibers present, these coming to a
point at fruit apices; locular epidermis without operculum, smooth, without pores.
Distribution and habitat:From 525616N and 76317728W on the Pacific coast of
northwestern Colombia at 346(351500) m elevation in lowland or montane tropical rainforest (Fig. 17).
Taxonomic notes:Geonoma divisa is related to two other species within the G. stricta cladeG.
longivaginata and G. ferruginea, both from Central America. It differs from these in its tricussately arranged,
closely spaced flower pits. It is sympatric with both Geonoma stricta and G. cuneata, and shares some
character states with these species, especially its fruits which are ridged from the numerous, subepidermal,
meridional, elongate fibers present, these coming to a point at fruit apices.
Subspecific variation:No trait varies within this species.
19. Geonoma elegans Martius (1826: 144). Type: BRAZIL. Rio de Janeiro: no locality, no date, H. Schott s. n.
(holotype, not known, presumed lost). Neotype (here designated): BRAZIL. Rio de Janeiro: Macio da Tijuca,
Reserva Florestal da FEEMA, 550 m, 17 October 1977, P. Maas & P. Carauta 3285 (neotype NY!).
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HENDERSON
Geonoma elegans var. robusta Drude (1882: 506). Type: BRAZIL. Rio de Janeiro: Barrire des Orgnes, 12 May 1877, A.
Glaziou 9018 (holotype BR!, isotypes FI!, NY!, P!).
Geonoma bifurca Drude & Wendland in Drude (1882: 504). Type: BRAZIL. Rio de Janeiro: no locality, no date, L.
Riedel 732 (holotype M!, isotypes BR!, C, n.v., F!, P!,).
internodes 1.5(0.63.0) cm long, yellowish and smooth. Leaves 9(810) per stem, undivided or irregularly
pinnate, rarely regularly pinnate and the pinnae with 1 main vein only, not plicate, bases of blades running
diagonally into the rachis; sheaths 8.2(4.513.0) cm long; petioles 12.0(3.029.0) cm long, drying green or
yellowish; rachis 26.0(13.545.0) cm long, 2.5(1.34.3) mm in diameter; veins not raised or slightly raised
and triangular in cross-section adaxially; pinnae 3(114) per side of rachis; basal pinna 22.4(15.534.5) cm
long, 4.9(0.28.2) cm wide, forming an angle of 33(852) with the rachis; apical pinna 14.1(8.026.5) cm
long, 9.4(4.015.3) cm wide, forming an angle of 29(2040) with the rachis. Inflorescences unbranched;
and asymmetrically apiculate, the surfaces without ridges; peduncular bracts 18.5(8.0285.0) cm long, welldeveloped, inserted 6.2(1.711.0) cm above the prophyll; peduncles 26.2(10.045.06) cm long, 2.5(1.34.8)
mm in diameter; rachillae 1, 17.6(3.528.0) cm long, 4.0(1.98.0) mm in diameter, the surfaces without spiky,
fibrous projections or ridges, drying brown or yellow-brown, without short, transverse ridges, not filiform and
not narrowed between the flower pits; flower pits spirally arranged, glabrous internally; proximal lips without
a central notch before anthesis, not recurved after anthesis, not hood-shaped; proximal and distal lips drying
the same color as the rachillae, not joined to form a raised cupule, the proximal lip margins overlapping the
distal lip margins; distal lips well-developed; staminate and pistillate petals not emergent, not valvate
throughout; staminate flowers deciduous after anthesis; stamens 6; thecae diverging at anthesis, inserted
almost directly onto the filament apices, the connectives bifid but scarcely developed; anthers short and curled
over at anthesis; non-fertilized pistillate flowers deciduous after anthesis; staminodial tubes crenulate or
shallowly lobed at the apex, those of non-fertilized flowers not projecting and persistent after anthesis; fruits
9.5(7.612.3) mm long, 7.2(5.89.2) mm in diameter, the bases with a prominent, asymmetric stipe, the apices
conical with rounded apices, the surfaces not splitting at maturity, without fibers emerging, not bumpy, not
apiculate; locular epidermis without operculum, smooth or sculpted and then usually also with a raised,
meridional ridge, without pores.
Distribution and habitat:From 16012655S and 39224949W in the Atlantic Coastal Forest
of Brazil (southern Bahia and Minas Gerais to Santa Catarina) at 265(2800) m elevation in lowland
Taxonomic notes:Wessels Boer (1968) stated that the type specimen of G. elegans was not at either M
or R, and it has not been seen at either herbaria, or any other, in the present study. A neotype is therefore
designated. Geonoma elegans is the first species dealt with here in a group of species from the Atlantic
Coastal Forest and adjacent Cerrado of Brazil (the G. schottiana clade, also including G. pauciflora, G.
pohliana, and G. schottiana). Although the clade is well-supported, all constituent species are extremely
variable internally. Geonoma elegans differs from other species in the group by its prophylls and peduncular
bracts which are ribbed with elongate, unbranched fibers, and both bracts are tubular, narrow, elongate,
closely sheathing the peduncle, and more or less persistent
Subspecific variation:Two traits vary within this species (leaf division, locular epidermis sculpting).
There is no geographic discontinuity, except for an outlying specimen from Minas Gerais. There are many
missing data for locular epidermis sculpting, and leaf division is not consistent, so these traits are not used to
divide specimens. Because of this, no subspecies are recognized. However, G. elegans is a variable species.
A specimen (Fiaschi 3154) from Rio de Janeiro has pinnate leaves with 14 pinnae per side of the rachis
(versus 16 for other specimens) and the pinnae have 1 main vein only (pinnate morphotype).
Three specimens (Riedel 732 (type of G. bifurca), MelloSilva 844, Martinelli 13314) from eastern part of
the Serra do Mar in Rio de Janeiro have narrow, undivided leaves (bifurca morphotype).
69
Six specimens (Bausen 129, Demuner 4032, Fernandes 1833, 1837, Folli 1660, Kollmann 4170) from
Esprito Santo have broad, undivided leaves (broad leaf morphotype). They occur on the Serra do Mar at 400
700 m elevation.
Five specimens (Fernandes 3095, 3103, 3290, Kollmann 6324, 9321) from Esprito Santo have thicker
rachillae than usual (thick rachillae morphotype). They occur on the Serra do Mar at 550800 m elevation.
Specimens from lower elevations (30400) in Esprito Santo and southern Bahia, including the outlier
from Minas Gerais, are smaller than usual (small morphotype). They have smaller leaves and these are mostly
undivided. Two of these (Hatschbach 47799 , Lombardi 5156) have non-sculpted endocarps (as opposed to
the more common sculpted). There is no overlap between the ranges of the small morphotype of G. elegans
and G. pauciflora, but specimens of the small morphotype resemble some of those of the pinnate-branched
morphotype of G. pauciflora and there may be a hybrid zone in the region of southern Bahia and Esprito
Santo between G. elegans and G. pauciflora.
Specimens from Rio de Janeiro to Santa Catarina are less variable and typically have three pinnae per side
of the rachis. A few specimens from So Paulo (Kirizawa 935, Leito Filho 33102, 34584) have undivided
leaves. Two specimens from the southern margin of the range, in Santa Catarina (Lourteig 2374, Smith 5712)
have non-sculpted endocarps (as opposed to the more common sculpted).
20. Geonoma epetiolata Moore (1980: 28). Type: PANAMA. Veraguas: Guabal (Dos Bocas del Ro
Calovbora), about 16 km NW of Santa F, 500 m, 1516 November 1974, R. Dressler 4777 (holotype BH!,
isotypes MO!, PMA! US!).
Plants 1.4(0.63.0) m tall; stems 1.6(0.53.0) m tall, 0.7(0.51.1) cm in diameter, solitary or clustered, canelike; internodes 1.2(0.62.6) cm long, yellowish and smooth. Leaves 9(810) per stem, undivided, not plicate,
bases of blades recurved against the rachis; sheaths 5.8(3.08.2) cm long; petioles absent; rachis 34.1(18.8
50.5) cm long, 4.5(2.27.5) mm in diameter; veins raised and rectangular in cross-section adaxially; pinnae 1
per side of rachis; basal pinna forming an angle of 14(424) with the rachis; apical pinna 9.0(5.214.0) cm
long, forming an angle of 30(1645) with the rachis. Inflorescences unbranched; prophylls and peduncular
the surfaces not ridged, without unequally wide ridges; peduncular bracts 7.6(7.08.7) cm long, welldeveloped, inserted 2.5(1.05.6) cm above the prophyll; peduncles 10.2(5.015.0) cm long, 2.3(1.33.2) mm
in diameter; rachillae 1, 20.3(9.038.0) cm long, 3.4(1.74.9) mm in diameter, the surfaces without spiky,
not narrowed between the flower pits; flower pits usually spirally arranged, sometimes tricussately, then the
groups not closely spaced nor consistently arranged throughout the rachillae, glabrous internally; proximal
lips drying the same color as the rachillae, not joined to form a raised cupule, the proximal lip margins
overlapping the distal lip margins; distal lips well-developed; staminate and pistillate petals not emergent, not
and curled over at anthesis; non-fertilized pistillate flowers persistent after anthesis; staminodial tubes lobed,
the lobes not spreading at anthesis, not acuminate, those of non-fertilized flowers not projecting and persistent
after anthesis; fruits 8.6(7.39.9) mm long, 5.1(4.75.3) mm in diameter, the bases without a prominent stipe,
the apices not conical, the surfaces not splitting at maturity, without fibers emerging, not bumpy, not apiculate,
ridged from the numerous, subepidermal, meridional, elongate fibers present, these coming to a point at fruit
apices; locular epidermis without operculum, smooth, without pores.
Distribution and habitat:From 8311023N and 79168406W in Costa Rica and Panama at
557(3001200) m elevation in lowland to montane tropical rainforest (Fig. 17).
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HENDERSON
Taxonomic notes:Geonoma epetiolata is a member of a group of four Central American species, part
of the G. cuneata clade, including G. brenesii, G. monospatha, and G. hugonis. They all have unbranched or
few-branched inflorescences and share the character state of the staminodial tubes being lobed at the apex, but
the lobes are not spreading at anthesis and are not acuminate. Although petiole length is treated as a
quantitative variable here, the short or virtually absent petiole of G. epetiolata is characteristic. This species is
also characterized by its mottled leaves (Blanco & Martn-Rodrguez 2007), although this mottling is not
obvious from herbarium specimens.
Subspecific variation:There is no variation in traits, except for stem branching. There is geographic
discontinuity, and specimens come from Costa Rica and several areas in Panama.
Specimens from Costa Rica and Santa F, Panama differ significantly from other Panamanian specimens
in nine quantitative variables (stem diameter, rachis length, rachis width, basal pinna angle, apical pinna
length, apical pinna angle, interbract distance, peduncle width, rachilla length)(t-test, P <0.05). However,
there is variation in both depth of staminodial tube lobing (de Nevers & Grayum 1998) and flower pit
arrangement, and this does not correspond to the geographic division.
Most specimens from Costa Rica have spirally arranged flower pits but some tend to be tricussately
arranged. Depth of lobes of staminodial tubes varies from 0.20.3 mm.
Specimens from Santa F, Panama have spirally arranged flower pits and depth of lobes of staminodial
tubes is 0.2 mm.
Specimens from Cerro Tife and El Cop, Panama have spirally to almost tricussately arranged flower pits
and depth of lobes of staminodial tubes varies from 0.20.9 mm.
Specimens from Llano Grande, Panama have spirally to almost tricussately arranged flower pits and depth
of lobes of staminodial tubes varies from 0.20.3 mm.
East of the Canal Zone, specimens from Cerro Brewster and Cerro Bruja have spirally arranged flower
pits and depth of lobes of staminodial tubes varies from 0.40.7 mm.
Independently of staminodial tube lobing, there are at least four distinct populations of G. epetiolata
Costa Rica; Santa F; El Cop/Llano Grande/Cerro Tife; and Cerro Brewster/Cerro Bruja. However, because
of the complexity in staminodial tube lobing and triad arrangement, and because there are few specimens from
most sites, and the gaps between the sites in Panama may be a result of insufficient collecting, no subspecific
taxa are recognized.
21. Geonoma euspatha Burret (1930c: 10). Type: COLOMBIA. Caquet: Sucre, 10 July 1926, G. Woronow &
J. Juzepczuk 5885 (holotype LE!).
Geonoma karuaiana Steyermark (1951: 88). Type: VENEZUELA. Bolvar: Ro Karuai, base of Sororopn-teup, W of
La Laja, 1220 m, 29 November 1944, J. Steyermark 60789 (holotype F!).
undivided or irregularly pinnate, not plicate, bases of blades running diagonally into the rachis; sheaths
16.6(8.522.5) cm long; petioles 46.5(11.076.0) cm long, drying green or yellowish; rachis 45.4(22.085.0)
cm long, 3.2(1.94.7) mm in diameter; veins raised and rectangular in cross-section adaxially; pinnae 3(16)
per side of rachis; basal pinna 38.8(20.063.5) cm long, 5.9(1.519.0) cm wide, forming an angle of 44(27
28(1437) with the rachis. Inflorescences branched 12 orders; prophylls and peduncular bracts not ribbed
with elongate, unbranched fibers, persistent; prophylls 21.2(9.031.0) cm long, not short and asymmetrically
apiculate, the surfaces not ridged, without unequally wide ridges; peduncular bracts 20.2(8.026.0) cm long,
well-developed, inserted 6.8(0.314.0) cm above the prophyll; peduncles 30.3(6.546.0) cm long, 4.3(2.1
7.6) mm in diameter; rachillae 13(221), 12.8(5.519.0) cm long, 2.4(1.53.4) mm in diameter, the surfaces
without spiky, fibrous projections or ridges, drying brown, with faint to pronounced, short, transverse ridges,
71
not filiform and not narrowed between the flower pits; flower pits spirally arranged, densely hairy internally
proximally and distally; proximal lips without a central notch before anthesis, not recurved after anthesis,
hood-shaped at anthesis, sometimes splitting post-anthesis; proximal and distal lips drying the same color as
the rachillae, not joined to form a raised cupule; distal lips absent; staminate and pistillate petals not emergent,
not valvate throughout; staminate flowers deciduous after anthesis; stamens 6; thecae diverging at anthesis,
and curled over at anthesis; non-fertilized pistillate flowers persistent or deciduous after anthesis; staminodial
tubes crenulate or shallowly lobed at the apex, those of non-fertilized pistillate flowers not projecting and
persistent after anthesis; fruits 6.3(4.98.4) mm long, 5.1(3.96.8) mm in diameter, the bases without a
prominent stipe, the apices not conical, the surfaces not splitting at maturity, without fibers emerging, bumpy
from the numerous, subepidermal, tangential, short fibers present, these coming to a point at fruit apices;
locular epidermis without operculum, smooth, without pores.
Distribution and habitat:From 552N1723S and 49157835W on eastern Andean slopes in
Colombia, Ecuador, Peru, and Bolivia, the Guayana Highland region and outlying montane areas in
Venezuela, Brazil, Guyana, Suriname, and French Guiana, and just reaching the Amazon region of Brazil
(Par, Rndonia), at 735(2001630) m elevation in lowland to montane rainforest (Fig. 18).
FIGURE 18. Distribution maps of Geonoma euspatha, G. ferruginea subsp. ferruginea, G. ferruginea subsp.
microspadix, and G. ferruginea subsp. nicaraguensis.
Taxonomic notes:Geonoma euspatha is the first species to be treated here of a group of related species
characterized by its lack of a distal lip of the flower pit and flower pits hairy internally. This group, the G.
72
HENDERSON
interrupta clade, also includes G. frontinensis, G. interrupta, G. pinnatifrons, and G. simplicifrons. These
species have had a checkered taxonomic history. Geonoma euspatha was included in G. interrupta by Wessels
Boer (1965), but later reinstated (Wessels Boer, 1968). It differs from G. interrupta and G. simplicifrons in its
flower pits which are densely hairy internally proximally and distally; from G. frontinensis in rachillae
surfaces with faint to pronounced, short, transverse ridges; and from G. pinnatifrons in its prophyll surfaces
without unequally wide ridges.
Subspecific variation:Three traits vary within this species (stem branching, stem type, leaf division).
There is geographic discontinuity and the species occurs in two areas: eastern Andean slopes of Colombia,
Ecuador, Peru, and Bolivia; and the Guayana Highland region of Venezuela and adjacent Brazil and the
Guianas. There are also outliers in the Amazon region of Brazil. Excluding the Amazon outliers, of which
there are only four specimens, specimens from eastern Andean slopes differ significantly from Guayana
Highland/Guiana specimens in only four variables (rachis width, basal pinna width, peduncular bract length,
number of rachillae)(t-test, P <0.05). Based on these results, no subspecies are recognized.
There is geographical variation in this species. Regression shows there are significant (P <0.05)
associations between elevation and one stem, three leaf, and nine inflorescence variables. Squared multiple R
for the regression of stem diameter on elevation is 0.54, sheath length 0.44, basal pinna angle 0.22, apical
pinna length 0.50, prophyll length 0.38, peduncle length 0.43, peduncle width 0.12, interbract distance 0.42,
rachilla length 0.19, rachilla width 0.08, number of rachillae 0.13, fruit length 0.40, and fruit diameter 0.21.
Stem diameter, sheath length, prophyll length, interbract distance, peduncle length, and rachilla length
decrease with elevation, and basal pinna angle, apical pinna length, peduncle width, rachis width, rachilla
width, number of rachillae, and fruit length and diameter increase with elevation. In particular, inflorescences
change with increasing elevation, with prophylls and interbract distances becoming shorter, peduncles shorter
and wider, rachillae fewer, shorter, and narrower, and fruits larger. Wessels Boer (1968) commented on these
changes with elevation.
22. Geonoma ferruginea Wendland ex Spruce (1871: 110). Type: COSTA RICA. Heredia: Sarapiqu valley,
1857, H. Wendland s.n. (holotype K!).
Plants 2.4(1.04.0) m tall; stems 2.2(1.05.0) m tall, 1.0(0.51.6) cm in diameter, solitary or clustered, canelike; internodes 2.7(0.86.2) cm long, yellowish and smooth. Leaves 9 per stem, irregularly pinnate, not
plicate, bases of the blades running diagonally into the rachis; sheaths 12.3(7.317.3) cm long; petioles
diameter; veins raised and rectangular in cross-section adaxially; pinnae 4(315) per side of rachis; basal
pinna 19.2(11.728.0) cm long, 2.6(0.58.0) cm wide, forming an angle of 65(3595) with the rachis; apical
fibers, flattened, deciduous; prophylls 7.5(4.712.2) cm long, short, asymmetrically apiculate, the margins
curved around the stem, the surfaces flat with dense, felty, brown tomentum, prophyll equal to and early
deciduous with the peduncular bract, the surfaces not ridged, without unequally wide ridges; peduncular
bracts 7.3(5.311.5) cm long, well-developed, inserted 0.3(0.10.5) cm above the prophyll; peduncles
6.4(3.710.7) cm long, 3.7(1.96.8) mm in diameter; rachillae 15(538), 8.2(3.815.0) cm long, 2.5(1.54.0)
mm in diameter, the surfaces without spiky, fibrous projections or ridges, drying brown or yellow-brown,
without short, transverse ridges, not filiform and not narrowed between the flower pits; flower pits spirally
arranged, glabrous internally; proximal lips without a central notch before anthesis, not recurved after
staminate and pistillate petals not emergent, not valvate throughout; staminate flowers deciduous after
anthesis; stamens 6; thecae diverging at anthesis, inserted onto bifid and well-developed, non-jointed
73
connectives; anthers short and curled over at anthesis; non-fertilized pistillate flowers deciduous after
anthesis; staminodial tubes crenulate or shallowly lobed at the apex, those of non-fertilized pistillate flowers
not projecting and persistent after anthesis; fruits 6.9(5.29.3) mm long, 5.4(4.27.3) mm in diameter, the
fruit apices; locular epidermis without operculum, smooth, without pores.
Taxonomic notes:Henderson et al. (1995) suggested that this species was part of G. longivaginata. It is
closely related to that species, differing in its shorter rachillae 8.1(3.815.0) cm long versus 22.9(10.0
42.0) cm longwithout short, transverse ridges.
Subspecific variation:One trait (stem branching) varies within this species. The species has a disjunct
distribution, and occurs in two areasNicaragua and Costa Rica, with outlying specimens in Honduras and
Guatemala. Within Costa Rica, there is an isolated, lower elevation subgroup from the Sarapiqu valley. Apart
from the outliers, there are three subgroups based on geography.
ANOVA shows that for pair wise comparison probabilities, 13 variables (stem diameter, rachis length,
rachis width, pinnae number, basal pinna width, basal pinna angle, apical pinna width, interbract distance,
peduncle length, peduncle width, rachilla width, number of rachillae, fruit diameter) differ significantly (P
<0.05) between one pair of subgroups, and one (rachilla length) differs amongst all three groups. Based on
these results, these three subgroups are recognized as subspecies (subspp. ferruginea, microspadix,
nicaraguensis). There are only three specimens from Honduras and Guatemalatoo few for analysisand
these outliers are unplaced for subspecies.
Key to the subspecies of G. ferruginea
1
2
-
Nicaragua ...................................................................................................................................... subsp. nicaraguensis

Costa Rica ..................................................................................................................................................................... 2
Rachillae 10(525), 9.7(6.415.0) cm long; Cordilleras Tilarn, Central, Talamanca, and Guanacaste .......................
............................................................................................................................................................ subsp. ferruginea
Rachillae 22(1138), 5.5( 3.88.0) cm long; Sarapiqu Valley ....................................................... subsp. microspadix
22a. Geonoma ferruginea subsp. ferruginea

Geonoma versiformis Wendland ex Spruce (1871: 109). Type: COSTA RICA. Cartago: near Turrialba, no date, H.
Inflorescences rachillae 10(525), 9.7(6.415.0) cm long.

Distribution and habitat:From 9171055N and 83088529W in Costa Rica (Cordilleras
Tilarn, Central, Talamanca, and Guanacaste) at 904(4001500) m elevation in lowland to montane tropical
Several specimens have unusually high numbers of pinnae, 915 versus the more usual 35. Specimens
from Volcan Arenal (Russell 683, 893, Lent 3335) have larger leaves and inflorescences than the others.
22b. Geonoma ferruginea subsp. microspadix (Wendland ex Spruce) Henderson, comb. & stat. nov.
Basionym: Geonoma microspadix Wendland ex Spruce (1871: 110). Type: COSTA RICA. Heredia: Sarapiqu valley,
Inflorescences rachillae 22(1138), 5.5( 3.88.0) cm long.

Distribution and habitat:From 10111028N and 83548412W in Costa Rica (Sarapiqu valley
and adjacent areas) at 520(100950) m elevation in lowland tropical rainforest (Fig. 18).
22c. Geonoma ferruginea subsp. nicaraguensis Henderson, subsp. nov. (Appendix IV, Plate 33)
A Geonoma ferruginea subsp. microspadix rhachillis parvioribus differt.
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HENDERSON
Type: NICARAGUA. Zelaya: Cerro La Pimienta, ca. 1345N, 8459W, ca. 900980 m, 13 April 1979, J. Pipoly 5112
(holotype NY!, isotype MO, n.v.).
Inflorescences rachillae 17(930), 7.5( 5.59.5) cm long.

Distribution and habitat:From 12151347N and 84598552W in Nicaragua at 956(3501500)
m elevation in lowland to montane tropical rainforest (Fig. 18).
23. Geonoma fosteri Henderson, sp. nov. (Appendix IV, Plate 34)
A speciebus affinibus prophyllis brevibus inaequaliter apiculatis atque rachide breviore, crusta fructuum fibris
subepidermalibus brevibus numerosis apicem convergentibus tuberculata, operculo carens differt.
Type: ECUADOR. Sucumbios: Sinangoe Station, Shishicho Ridge, Alto Aguarico drainage, above (south of) Ro
Cofanes, west of Puerto Libre, NW of Lumbaqui, 0012N, 7731 W, 13001450 m, 14 August 2001, R. Aguinda,
N. Pitman & R. Foster 1315 (holotype F!, isotype QCNE, n.v.).
Plants height no data; stems 1.5 m tall, 0.7 cm in diameter, cane-like; internodes 1.0 cm long, yellowish and
smooth. Leaves irregularly pinnate, not plicate, bases of blades running diagonally into the rachis; sheaths 6.5
cm long; petioles 11.0 cm long, drying green or yellowish; rachis 20.5 cm long, 3.1(2.14.1) mm in diameter;
veins raised and rectangular in cross-section adaxially; pinnae 3 per side of rachis; basal pinna 19.5 cm long,
1.5 cm wide, forming an angle of 63(5868) with the rachis; apical pinna 16.5 cm long, 8.5 cm wide, forming
an angle 28 with the rachis. Inflorescences branched 3 orders; prophylls and peduncular bracts not ribbed
with elongate, unbranched fibers; prophylls 5.2 cm long, short, asymmetrically apiculate, the margins curved
around the stem, the surfaces flat with dense, felty, brown tomentum, prophyll equal to and early deciduous
with the peduncular bract, the surfaces not ridged, without unequally wide ridges; peduncular bracts no data,
inserted 0.4 cm above the prophyll; peduncles 5.7 cm long, 3.4(2.93.8) mm in diameter; rachillae 10.5 cm
long, 1.0(0.91.1) mm in diameter, the surfaces without spiky, fibrous projections or ridges, drying brown,
with faint to pronounced, short, transverse ridges, filiform with extended narrowed sections between the
flower pits; flower pits alternately arranged (sometimes distorted by twisting and contracting of rachillae),
crenulate or shallowly lobed at the apex, those of non-fertilized pistillate flowers not projecting and persistent
after anthesis; fruits size no data, the bases without a prominent stipe, the apices not conical, the surfaces not
splitting at maturity, without fibers emerging, bumpy from the numerous, subepidermal, tangential, short
fibers present, these coming to a point at fruit apices; locular epidermis without operculum.
Distribution and habitat:From 012019N and 77257731W on eastern Andean slopes in
Ecuador at 1592(13751810) m elevation in montane rainforest (Fig. 19).
Taxonomic notes:Geonoma fosteri is similar to a group of species within the G. lanata clade. It differs
from G. bernalii, G. dindoensis, G. lanata, and G. venosa in having short, asymmetrically apiculate prophylls
with the margins curved around the stem and the surfaces flat with dense, felty, brown tomentum; from G.
tenuissima in its fruit surfaces bumpy from the numerous, subepidermal, tangential, short fibers present; and
from G. operculata by its locular epidermis without an operculum. Only two specimens are known. These
share the same character states as G. braunii, but the peduncular bract is unknown in G. fosteri. Given the large
geographic distance between the two, they are kept separate pending more complete material.
Subspecific variation: No trait varies within this species.
75
FIGURE 19. Distribution maps of Geonoma fosteri, G. frontinensis, G. galeanoae, and G. gentryi.
24. Geonoma frontinensis Burret (1930a: 170). Type: COLOMBIA. Antioquia: Frontino, 14001700 m,
September 1871, F. Lehmann 7323 (holotype B, destroyed, isotype K!).
Geonoma mucronata Burret (1930a: 171). Type: COLOMBIA. Antioquia: Dos Quebradas, 1500 m, 7 January 1880, W.
Kalbreyer 1334 (holotype B, destroyed). Neotype (designated by Bernal et al. 1989): COLOMBIA. Antioquia:
Carretera San Carlos-Granada, Dos Quebradas, 4.5 km E of San Carlos, 1200 m, 20 September 1987, R. Bernal & L.
Tobn 1383 (neotype COL!, isoneotypes MO!, NY!).
Plants 1.1(0.52.0) m tall; stems 0.2(0.20.3) m tall, solitary or clustered; internodes no data. Leaves 6(48)
per stem, undivided or irregularly pinnate, not plicate, bases of blades running diagonally into the rachis;
sheaths 30.0 cm long; petioles 68.8(26.0110.0) cm long, drying green or yellowish; rachis 40.0(22.565.0)
26(2034) with the rachis. Inflorescences unbranched or branched 1 order; prophylls and peduncular bracts
not ribbed with elongate, unbranched fibers, persistent; prophylls 19.8(13.426.2) cm long, not short and
asymmetrically apiculate, the surfaces not ridged, without unequally wide ridges; peduncular bracts
20.2(14.326.5) cm long, well-developed, inserted 9.7(4.017.0) cm above the prophyll; peduncles
76
HENDERSON
arranged, densely hairy internally proximally and distally; proximal lips without a central notch before
anthesis, not recurved after anthesis, hood-shaped at anthesis, sometimes splitting post-anthesis; proximal lips
drying the same color as the rachillae; distal lips absent; staminate and pistillate petals not emergent, not
and curled over at anthesis; non-fertilized pistillate flowers persistent after anthesis; staminodial tubes
the apices not conical, the surfaces not splitting at maturity, without fibers emerging, bumpy from the
numerous, subepidermal, tangential, short fibers present, these coming to a point at fruit apices; locular
epidermis without operculum, sculpted, usually also with a raised, meridional ridge, without pores.
Distribution and habitat:From 112714N and 73587700W on eastern slopes of the Western,
Central, and Eastern Cordilleras in Colombia at 1560(10002030) m elevation in montane rainforest (Fig. 19).
Taxonomic notes:See notes under Geonoma euspatha.
inflorescence branching). There is geographic discontinuity, but too few specimens to test for differences
between areas.
25. Geonoma galeanoae Henderson, sp. nov. (Appendix IV, Plates 3537)
A speciebus affinibus crusta prophylli cristis indivisis parallelis aequalibus atque proximis ferens, defectus operculi
differt.
Type: COLOMBIA. Antioquia: Mun. San Luis, 16 km. S.O. de las partidas a San Luis, sobre la va Medelln-Bogot,
Vereda la Josefina, 600N 7450W, 800 m, 25 June 1987, R. Callejas, J. Betancur, A. Arbelaez & H. Correa 4208
(holotype NY!, isotype HUA, n.v.).
Plants 3.0 m tall; stems 2.0 m tall, 1.7 cm in diameter, solitary, cane-like; internodes 2.0 cm long, yellowish
and smooth. Leaves 12 per stem, irregularly pinnate, not plicate, bases of blades running diagonally into the
rachis; sheaths 24.0(18.030.0) cm long; petioles 37.0(36.038.0) cm long, drying green or yellowish; rachis
54.3(43.565.0) cm long, 4.9(4.45.5) mm in diameter; veins raised and rectangular in cross-section
adaxially; pinnae 7(410) per side of rachis; basal pinna 38.0 cm long, 9.2(5.515.5) cm wide, forming an
angle of 42(3549) with the rachis; apical pinna 32.8(28.537.0) cm long, 19.3(16.522.0) cm wide, forming
an angle of 32(3132) with the rachis. Inflorescences branched 2 orders; prophylls and peduncular bracts not
ribbed with elongate, unbranched fibers, deciduous; prophylls 19.5 cm long, prophylls not short and
asymmetrically apiculate, the surfaces ridged with close, equal, parallel, non-dividing ridges, scarcely
tomentose between the ridges, without unequally wide ridges; peduncular bracts 16.0 cm long, welldeveloped, inserted 0.3(0.20.4) cm above the prophyll; peduncles 7.4(4.810.3) cm long, 7.0(5.68.3) mm in
diameter; rachillae 20, 14.6(13.216.0) cm long, 2.6 mm in diameter, the surfaces without spiky, fibrous
central notch before anthesis, not recurved after anthesis, hood-shaped at anthesis, sometimes splitting postanthesis; proximal and distal lips drying the same color as the rachillae, not joined to form a raised cupule, the
proximal lip margins overlapping the distal lip margins; distal lips well-developed; staminate and pistillate
petals not emergent, not valvate throughout; staminate flowers deciduous after anthesis; stamens 6; thecae
diverging at anthesis, inserted almost directly onto the filament apices, the connectives bifid but scarcely
developed; anthers short and curled over at anthesis; non-fertilized pistillate flowers deciduous after anthesis;
staminodial tubes crenulate or shallowly lobed at the apex, those of non-fertilized pistillate flowers not
77
projecting and persistent after anthesis; fruits size no data, the bases without a prominent stipe, the apices not
conical, the surfaces not splitting at maturity, without fibers emerging, bumpy from the numerous,
without operculum.
Distribution and habitat:From 553615N and 74507500W on the eastern slopes of the
Central Cordillera in Colombia at 830(800860) m elevation in lowland rainforest (Fig. 19).
Taxonomic notes:Geonoma galeanoae is similar to a group of species (G. baculifera, G.
calyptrogynoidea, G. concinna, G. conncinnoidea, G. congesta) in having the prophyll surfaces ridged with
close, equal, parallel, non-dividing ridges. It differs from these in its locular epidermis without an operculum.
Only three specimens are known.
Subspecific variation: No trait varies within this species, nor is there geographic disjunction.
26. Geonoma gentryi Henderson, sp. nov. (Appendix IV, Plate 38)
A speciebus affinibus prophyllis haud brevibus necnon inaequaliter apiculatis differt.
Type: COLOMBIA. Choc: 2 km S of Las Animas on road to Istmina, 150 m, 13 August 1976, A. Gentry & M. Fallen
17623 (holotype MO!).
Plants 1.5 m tall; stems height no data, 0.8 cm in diameter, branching no data, cane-like; internodes 2.9 cm
long, yellowish and smooth. Leaves undivided, not plicate, bases of blades running diagonally into the rachis;
sheaths 6.0 cm long; petioles 12.0 cm long, drying green or yellowish; rachis 26.0 cm long, 2.4 mm in
diameter; veins not raised or slightly raised and triangular in cross-section adaxially; pinnae 1 per side of
rachis; basal pinna length and width not applicable, forming an angle of 25 with the rachis; apical pinna 11.6
cm long, width not applicable; forming an angle of 28 with the rachis . Inflorescences branched 2 orders;
prophylls and peduncular bracts not ribbed with elongate, unbranched fibers, flattened, deciduous or
persistent; prophylls length no data, not short and asymmetrically apiculate, surfaces no data; peduncular
bracts length no data, well-developed, inserted 2.4 cm above the prophyll; peduncles 18.7 cm, 3.1 mm in
diameter; rachillae 12, 5.3 cm long, 2.0 mm in diameter, the surfaces without spiky, fibrous projections or
ridges, drying brown, with faint to pronounced, short, transverse ridges, not filiform and not narrowed
between the flower pits; flower pits spirally arranged, glabrous internally; proximal lips without a central
notch before anthesis, not recurved after anthesis, not hood-shaped; proximal and distal lips drying the same
color as the rachillae, joined to form a raised cupule, the margins not overlapping; distal lips well-developed;
anthesis; stamens 6; thecae diverging at anthesis, inserted almost directly onto the filament apices, the
flowers deciduous after anthesis; staminodial tubes crenulate or shallowly lobed at the apex; staminodial tubes
of non-fertilized pistillate flowers not projecting and persistent after anthesis; fruits 8.6 mm long, 5.7 mm in
coming to a point at fruit apices; locular epidermis without operculum.
Distribution and habitat:At 516N and 7637W on the Pacific coast of Colombia (Choc) at 150 m
Taxonomic notes:Geonoma gentryi is a member of the G. lanata clade, within which it appears most
similar to G. spinescens. Both these species lack data for some prophyll characters, but given its wide
geographic separation and different habitat (lowland rainforest at 150 m in the Choc versus lowland or
montane rainforest at 1075(8001330) m in the Coastal Range in Venezuela), G. gentryi is recognized as a
distinct species.
Subspecific variation: No trait varies within this species, and only one specimen is known.
78
HENDERSON
27. Geonoma hollinensis Henderson, Borchsenius & Balslev (2008: 195). Type: ECUADOR. Napo: HollinLoreto road to Coca 27 km from take-off from Baeza-Tena road, 042S 7740W, 10001100 m, 28
September 1995, H. Balslev 6418 (holotype AAU!, isotype QCA, n.v.).
internodes 0.6(0.50.8) cm long, yellowish and smooth. Leaves 6(67) per stem, undivided, not plicate, bases
of blades running diagonally into the rachis; sheaths 9.3(8.510.5) cm long; petioles 8.8(6.012.5) cm long,
drying green or yellowish; rachis 25.0(23.526.7) cm long, 2.9(2.43.4) mm in diameter; veins not raised or
slightly raised and triangular in cross-section adaxially; pinnae 1 per side of rachis; basal pinna length and
width not applicable, forming an angle of 31(2638) with the rachis; apical pinna 16.8(14.519.5) cm long,
forming an angle of 27(2230) with the rachis. Inflorescences branched 2 orders; prophylls and peduncular
bracts not ribbed with elongate, unbranched fibers, flattened, deciduous or persistent; prophylls 10.0(9.9
10.0) cm long, not short and asymmetrically apiculate, the surfaces not ridged, without unequally wide ridges;
peduncular bracts 2.5 cm long, vestigial, inserted 2.2(0.93.4) cm above the prophyll; peduncles 11.1(9.5
12.7) cm long, 2.3(1.82.6) mm in diameter; rachillae 29(2334), 8.5(8.28.8) cm long, 1.6(1.32.3) mm in
diameter, the surfaces without spiky, fibrous projections or ridges, drying brown, with faint to pronounced,
short, transverse ridges, not filiform and not narrowed between the flower pits; flower pits tricussately
arranged throughout the rachillae, the groups of pits closely spaced, glabrous internally; proximal lips without
a central notch before anthesis, not recurved after anthesis, hood-shaped at anthesis, sometimes splitting postanthesis; proximal and distal lips drying the same color as the rachillae, not joined to form a raised cupule, the
proximal lip margins overlapping the distal lip margins; distal lips absent; staminate and pistillate petals not
anthesis, inserted almost directly onto the filament apices, the connectives bifid but scarcely developed;
anthers short and curled over at anthesis; staminodial tubes crenulate or shallowly lobed at the apex, those of
non-fertilized pistillate flowers not projecting, deciduous after anthesis; fruits 6.8 mm long, 5.0 mm in
diameter, the bases without a prominent stipe, the apices not ovoid and conical apices, the surfaces not
fibers present, these coming to a point at fruit apices; locular epidermis without operculum, smooth, without
pores.
Distribution and habitat:From 042052S and 77267740W in Ecuador at 1125(10501200)
m elevation in montane rainforest on eastern Andean slopes (Fig. 20).
Taxonomic notes:Geonoma hollinensis is one of only two species of Geonoma with staminate flowers
with three stamens; G. triandra is the other. Lacking staminate flowers, it can also be recognized by its
vestigial peduncular bracts and tricussately arranged flower pits.
Subspecific variation:There is no variation in any trait amongst the four specimens known.
28. Geonoma hugonis Grayum & de Nevers in de Nevers & Grayum (1998: 94). Type: PANAMA. Chiriqu:
Fortuna Dam Area, between Quebrada Los Chorros and Quebrada Hondo, to N of reservoir, in forest N of
road, 845N, 8214W, 1100 m, 20 September 1984, H. Churchill & A. Churchill 6185 (holotype MO!,
isotype CAS, n.v.).
Plants 0.8(0.31.5) m tall; stems 0.7(0.11.2) m tall, 0.6(0.40.8) cm in diameter, solitary or clustered, canelike; internodes 1.9(1.03.5) cm long, covered with reddish or brownish scales, especially in their distal part.
Leaves 8(412) per stem, undivided, not plicate, bases of blades running diagonally into the rachis; sheaths
7.2(3.514.0) cm long; petioles 7.0(1.012.0) cm long, drying green or yellowish; rachis 12.7(7.020.0) cm
long, 1.8(1.32.7) mm in diameter; veins raised and rectangular in cross-section adaxially; pinnae 1 per side
of rachis; basal pinna length and width not applicable, forming an angle of 29(2240) with the rachis; apical
pinna 10.4(5.817.2) cm long, width not applicable, forming an angle of 32(2242) with the rachis.
79
Inflorescences unbranched; prophylls and peduncular bracts ribbed with elongate, unbranched fibers, both
bracts tubular, narrow, elongate, closely sheathing the peduncle, more or less persistent; prophylls 12.8(11.0
peduncles 29.9(13.543.5) cm long, 1.3(0.92.0) mm in diameter; rachillae1, 7.1(4.612.5) cm long, 2.4(1.7
3.5) mm in diameter the surfaces without spiky, fibrous projections or ridges, drying brown or yellow-brown,
arranged, glabrous internally; proximal lips with a central notch before anthesis, often the two sides of the
notch overlapping, not recurved after anthesis, not hood-shaped at anthesis; proximal and distal lips drying the
anthesis; non-fertilized pistillate flowers deciduous after anthesis; staminodial tubes lobed, the lobes not
spreading at anthesis, not acuminate; staminodial tubes of non-fertilized pistillate flowers not projecting and
Distribution and habitat:From 830846N and 81458217W in western Panama at
1243(11001450) m elevation in montane rainforest (Fig. 20).
FIGURE 20. Distribution maps of Geonoma hollinensis and G. hugonis.
Taxonomic notes:Geonoma hugonis is a member of a group of four Central American species, part of
the G. cuneata clade, also including G. brenesii, G. monospatha, and G. epetiolata. They all have unbranched
or few-branched inflorescences and share the character state of the staminodial tubes being lobed at the apex,
but the lobes are not spreading at anthesis and are not acuminate. Geonoma hugonis differs from these species
in its internodes covered with reddish or brownish scales.
Subspecific variation:Only one trait (stem branching) varies in this species. There is geographic
disjunction but this is likely to be an artifact of insufficient collecting.
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HENDERSON
29. Geonoma interrupta (Ruz & Pavn) Martius (1823: 8). Martinezia interrupta Ruz & Pavn (1798: 296).
Type: PERU. Hunuco: between Pozuzo and Cuchero, November 1820, J. Pavn s.n. (holotype MA n.v.,
holotype image!, isotypes K!, M!, MO!).
Plants 3.7(1.08.0) m tall; stems 3.2(0.38.0) m tall, 3.7(2.65.0) cm in diameter, solitary, not cane-like or
cane-like; internodes 1.5(0.82.8) cm long, yellowish and smooth. Leaves 13(824) per stem, irregularly
mm in diameter; veins raised and rectangular in cross-section adaxially; pinnae 18(447) per side of rachis;
basal pinna 47.4(23.575.5) cm long, 5.7(0.227.0) cm wide, forming an angle of 43(2460) with the rachis;
apical pinna 40.3(23.065.5) cm long, 20.3(0.439.0) cm wide, forming an angle of 29(1440) with the
rachis. Inflorescences branched 24 orders; prophylls and peduncular bracts not ribbed with elongate,
asymmetrically apiculate, the surfaces ridged and densely tomentose with widely to closely spaced ridges, the
ridges unequally wide, often dividing from and rejoining other ridges, the prophyll margins with irregular,
spine-like projections, the prophylls usually splitting irregularly between the ridges; peduncular bracts
17.8(10.525.5) cm long, well-developed, inserted 4.1(1.08.0) cm above the prophyll; peduncle 22.7(7.0
34.0) cm long, 13.6(2.520.1) mm in diameter; rachillae 71(22120), 19.1(9.229.7) cm long, 2.3(1.33.4)
mm in diameter, the surfaces without spiky, fibrous projections or ridges, drying brown, with faint to
pronounced, short, transverse ridges, not filiform and not narrowed between the flower pits; flower pits
spirally arranged, densely hairy internally distally only (rarely some hairs on lateral) margins of the pits;
proximal lips without a central notch before anthesis, not recurved after anthesis, hood-shaped at anthesis,
sometimes splitting post-anthesis; proximal and distal lips drying the same color as the rachillae, not joined to
form a raised cupule, the proximal lip margins overlapping the distal lip margins; distal lips absent; staminate
6; thecae diverging at anthesis, inserted almost directly onto the filament apices, the connectives bifid but
scarcely developed; anthers short and curled over at anthesis; non-fertilized pistillate flowers persistent or
deciduous after anthesis; staminodial tubes crenulate or shallowly lobed at the apex; staminodial tubes of nonfertilized pistillate flowers not projecting and persistent after anthesis; fruits 5.8(4.47.5) mm long, 4.6(3.6
6.3) mm in diameter, the bases without a prominent stipe, the apices not conical, the surfaces not splitting at
maturity, without fibers emerging, bumpy from the numerous, subepidermal, tangential, short fibers present,
these coming to a point at fruit apices; locular epidermis without operculum, sculpted, usually also with a
raised, meridional ridge, without pores.
Taxonomic notes:Geonoma interrupta is a member of a group of species characterized by its lack of a
distal lip of the flower pit and flower pits hairy internally. This group, the G. interrupta clade, also includes G.
euspatha, G. frontinensis, G. pinnatifrons, and G. simplicifrons. These species have had a checkered taxonomic
history. Geonoma interrupta has been treated in a broad (e.g., Henderson et al., 1995) or narrower sense
(Wessels Boer, 1968). Geonoma interrupta differs from G. euspatha, G. frontinensis, and G. simplicifrons in its
prophyll surfaces with unequally wide ridges, and from G. pinnatifrons in its flower pits which are densely
hairy internally distally only.
Wessels Boer (1968) used rachillae hairs to distinguish G. interrupta from G. pinnatifrons. This was
refined somewhat by Hammel et al. (2003) who described G. interrupta as having hairs to ca. 0.15 mm long
with at least some branched, and G. pinnatifrons (as G. oxycarpa) as having unbranched hairs 0.10.5 mm
long. As stated in the Materials and Methods section, rachillae hairs are not used in the present study because
potential states can not be scored unequivocally and hairs are early deciduous. However, G. interrupta and G.
pinnatifrons can be distinguished by rachillae hairs if rachillae at an early stage are present.
Subspecific variation:Two traits vary within this species (stem type, pistillate flower persistence).
Excluding stem type, for which there are few data, state distributions of the remaining trait (pistillate flower
persistence) divide the specimens into two subgroups. Within each subgroup there is geographic discontinuity.
81
The first subgroup, with persistent pistillate flowers, occurs in the Andes in Colombia, and plants are
reported to be rheophytes. This subgroup is recognized as a subspecies (subsp. rivalis).
The second subgroup, with deciduous pistillate flowers, has several gaps in its distribution, and there are
several potential geographic subgroups. There is considerable variation in several variables (number of
pinnae, prophyll length, peduncular bract length, interbract distance) and combining these with geographic
division, three subgroups can be recognized: Central America and Colombia (with longer bracts and more
pinnae); Ecuador, Peru, and Bolivia (shorter bracts and fewer pinnae); and Venezuela and just reaching
adjacent Colombia (longer bracts and fewer pinnae). ANOVA shows that for pair wise comparison
probabilities, seven variables (stem height, sheath length, rachis length, number of pinnae, prophyll length,
peduncular bract length, interbract distance) differ significantly (P <0.05) between one pair of groups,
although no variable differs amongst all three groups. Based on these results and geographic discontinuity, the
three subgroups are recognized as subspecies (subsp. magnifica from Central America and Colombia, subsp.
interrupta from Ecuador, Peru, and Bolivia, and subsp. purdieana from Venezuela and adjacent Colombia).
Key to the subspecies of Geonoma interrupta
1
2
3
-
Non-fertilized pistillate flowers persistent after anthesis; rheophytes; Central Cordillera in Colombia...subsp. rivalis
Non-fertilized pistillate flowers deciduous after anthesis; non-rheophytes; widespread ............................................ 2
Pinnae 25(447) per side of rachis; Mexico, Belize, Guatemala, Honduras, Nicaragua, Costa Rica, Panama, and
Colombia..............................................................................................................................................subsp. magnifica
Pinnae 9(414) per side of rachis; Venezuela and adjacent Colombia, Ecuador, Peru, and Bolivia ............................ 3
Peduncular bracts inserted 3.0(1.28.0) cm above the prophyll; Ecuador, Peru, and Bolivia ............subsp. interrupta
Peduncular bracts inserted 7.1(6.57.5) cm above the prophyll; Venezuela and adjacent Colombia subsp. purdieana
29a. Geonoma interrupta subsp. interrupta

Leaves pinnae 9(412) per side of rachis. Inflorescences peduncular bracts inserted 3.0(1.28.0) cm above the
prophyll; non-fertilized pistillate flowers deciduous after anthesis.
Distribution and habitat:From 105N1425S and 70587930W in Ecuador, Peru, and Bolivia
at 533(501160) m elevation in lowland to montane rainforest (Fig. 21).
29b. Geonoma interrupta subsp. magnifica (Linden & Wendland) Henderson, comb. & stat. nov.
Basionym: Geonoma magnifica Linden & Wendland in Wendland (1856: 335). Type: MEXICO. Tabasco: between San
Carlos and Macsupana, no date, A. Ghiesbreght s. n. (holotype GOET, n.v.).
Geonoma dryanderae Burret (1935c: 615). Type: COLOMBIA. Valle: Central Cordillera, Ro Tulua, 1200 m, June 1935,
J. Dryander 30 (holotype B, n.v., holotype image!).
Leaves pinnae 25(447) per side of rachis. Inflorescences peduncular bracts inserted 4.0(1.57.4) cm above
the prophyll; non-fertilized pistillate flowers deciduous after anthesis.
Distribution and habitat:From 3261834N and 72369506W in Mexico, Belize, Guatemala,
Honduras, Nicaragua, Costa Rica, Panama, and Colombia at 382(01500) m elevation in lowland to montane
29c. Geonoma interrupta subsp. purdieana (Spruce) Henderson, comb. & stat. nov.
Basionym: Geonoma purdieana Spruce (1871: 109). Type: COLOMBIA. La Guajira: Ro de la Hacha, December 1844,
W. Purdie 259 (holotype K!).
Leaves pinnae 9(514) per side of rachis. Inflorescences peduncular bracts inserted 7.1(6.57.5) cm above the
prophyll; non-fertilized pistillate flowers deciduous after anthesis.
Distribution and habitat:From 714N1105S and 69167328W in Venezuela and adjacent
Colombia at 375(60775) m elevation in lowland rainforest (Fig. 21).
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HENDERSON
29d. Geonoma interrupta subsp. rivalis (Kalbreyer & Burret) Henderson, comb. & stat. nov.
Basionym: Geonoma rivalis Kalbreyer & Burret in Burret (1930a: 241). Type: COLOMBIA. Antioquia: Coco, 700900
m, 18 February 1880, W. Kalbreyer 1427 (holotype B, destroyed). Neotype (here designated): COLOMBIA.
Antioquia: Mun. San Luis, Ro Samana on the Medelln-Bogot road, 600N, 7450W, 700780 m, 23 June 1987,
R. Callejas, A. Arbelaez, H. Correa & J. Betancur 4098 (neotype NY!, isoneotypes HUA, n.v., MO!).
Leaves pinnae 31(2736) per side of rachis. Inflorescences peduncular bracts inserted 2.6(1.04.5) cm above
the prophyll; non-fertilized pistillate flowers persistent after anthesis.
Distribution:From 550722N and 74457505W on the eastern slopes of the Central Cordillera
in Colombia, at 548 (150900) m elevation in lowland rainforest (Fig. 21). Plants are reported to be
rheophytes.
There is geographic discontinuity but too few specimens to test for differences amongst areas.
FIGURE 21. Distribution maps of Geonoma interrupta subsp. interrupta, G. interrupta subsp. magnifica, G. interrupta
subsp. purdieana, and G. interrupta subsp. rivalis.
30. Geonoma lanata Henderson, Borchsenius & Balslev (2008: 195). Type: ECUADOR. Carchi: Tulcan,
Reserva Etnica Aw, Parroquia Chical, Centro Gualpi Medio, 102N 7816W, 900 m, 25 February 1993, C.
Aulestia & A. Grijalva 1200 (holotype QCNE, n.v., isotypes AAU!, MO n.v.).
Plants 1.4(0.72.0) m tall; stems 1.3(0.52.0) m tall, 0.7(0.50.9) cm in diameter, solitary or clustered, canelike; internodes 1.6(0.73.1) cm long, yellowish and smooth. Leaves 9(711) per stem, undivided, not plicate,
83
long, drying green or yellowish; rachis 15.2(10.718.7) cm long, 1.9(1.22.5) mm in diameter; veins raised
and rectangular in cross-section adaxially; pinnae 1 per side of rachis; basal pinna length and width not
applicable, forming an angle of 40(3346) with the rachis; apical pinna 13.1(10.217.0) cm long, width not
applicable, forming an angle of 39(3643) with the rachis. Inflorescences branched 1 order; prophylls and
peduncular bracts not ribbed with elongate, unbranched fibers, flattened, persistent; prophylls 10.1(9.610.5)
cm long, not short and asymmetrically apiculate, the surfaces not ridged, without unequally wide ridges;
peduncular bracts 7.8 cm long, well-developed, inserted 3.0(1.06.9) cm above the prophyll; peduncles
mm in diameter, the surfaces without spiky, fibrous projections or ridges, drying brown, with faint to
pronounced, short, transverse ridges, not filiform and not narrowed between the flower pits; flower pits
and distal lips drying the same color as the rachillae, joined laterally with no clear gap between them, often
forming a raised cupule, the margins not overlapping; distal lips well-developed; staminate and pistillate
staminodial tubes crenulate or shallowly lobed at the apex; staminodial tubes of non-fertilized pistillate
flowers not projecting and persistent after anthesis; fruits 7.8(7.09.5) mm long, 6.1(5.46.8) mm in diameter,
the bases without a prominent stipe, the apices not conical, the surfaces not splitting at maturity, without fibers
Distribution and habitat:From 115N337S and 78147949W in Ecuador on western Andean
slopes at 746(2001800) m elevation in lowland or montane rainforest (Fig. 22).
Taxonomic notes:Specimens of this species were identified by Skov (1989) as Geonoma aff.
pauciflora, and were considered by Borchsenius et al. (1998) as a western Andean form of G. leptospadix.
Geonoma lanata is not closely related to either species, and belongs to a group of 12 species, the G. lanata
clade. It differs from these in its alternately arranged flower pits and bumpy fruits with numerous,
subepidermal, tangential, short fibers present, these coming to a point at fruit apices.
Subspecific variation:Only one trait (stem branching) varies in this species. There is geographic
disjunction but this may be an artifact of insufficient collecting. Most specimens are from the northern part of
the range, and only four from the southern parttoo few to test for differences. Galeano & Bernal (2010)
report that this species also occurs in southwestern Colombia.
31. Geonoma laxiflora Martius (1823: 12). Lectotype (designated by Wessels Boer 1968): BRAZIL.
Amazonas: Rio Japur, no date, C. Martius s.n. (lectotype M!).
Geonoma laxiflora var. depauperata Trail (1876: 326). Type: BRAZIL. Amazonas: Anana, N bank of Rio Solimes, 6
September 1874, J. Trail 1024/CXVI (holotype K!).
internodes 1.7(1.03.0) cm long, yellowish and smooth. Leaves 7(510) per stem, undivided or rarely
long; petioles 9.1(4.013.5) cm long, drying green or yellowish; rachis 20.6(11.531.5) cm long, 2.9(1.14.6)
mm in diameter; veins not raised or slightly raised and triangular in cross-section adaxially; pinnae 1(13) per
side of rachis; basal pinna length and width not applicable, forming an angle of 27(1435) with the rachis;
apical pinna 22.0(13.031.5) cm long, width not applicable, forming an angle of 23(1831). Inflorescences
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HENDERSON
deciduous; prophylls 3.9(2.35.3) cm long, short, asymmetrically apiculate, the margins curved around the
stem, the surfaces flat with dense, felty, brown tomentum, prophyll equal to and early deciduous with the
peduncular bract, the surfaces not ridged, without unequally wide ridges; peduncular bracts 1.7(1.03.0) cm
long, well-developed, inserted 0.2(0.10.4) cm above the prophyll; peduncles 3.8(2.25.5) cm long, 3.0(1.5
without spiky, fibrous projections or ridges, drying brown, with faint to pronounced, short, transverse ridges,
not filiform and not narrowed between the flower pits; flower pits usually spirally arranged, sometimes
decussately or tricussately, then the groups not closely spaced nor consistently arranged throughout the
rachillae, glabrous internally; proximal lips without a central notch before anthesis, not recurved after
anthesis, hood-shaped at anthesis, sometimes splitting post-anthesis; proximal and distal lips drying the same
lobed at the apex; staminodial tubes of non-fertilized pistillate flowers not projecting and persistent after
apices not conical, the surfaces not splitting at maturity, without fibers emerging, not bumpy, not apiculate;
Colombia, Brazil, Ecuador, Peru, and Bolivia at 161(80320) m elevation usually along river margins in
Taxonomic notes:This species is commonly associated with both G. leptospadix and G. deversa (e.g.,
Wessels Boer 1968) to which it is closely related. It differs from these in its prophyll surfaces which are flat
and have dense, felty, brown tomentum.
Subspecific variation:No trait except leaf division varies within this species. One specimen (of 34) has
a pinnate leaf.
32. Geonoma lehmannii Burret (1930a: 180). Type: COLOMBIA. Antioquia: between Abejoral and La Ceja,
18002200 m, no date, F. Lehmann 4630 (holotype B, destroyed, isotype K!).
undivided or irregularly pinnate, not plicate or plicate, bases of blades running diagonally into the rachis;
51.5) cm long, 3.5(1.47.6) mm in diameter; veins raised and rectangular in cross-section adaxially or not
raised or slightly raised and triangular in cross-section adaxially; pinnae 3(19) per side of rachis; basal pinna
unbranched; prophylls and peduncular bracts ribbed with elongate, unbranched fibers, both bracts tubular,
narrow, elongate, closely sheathing the peduncle, more or less persistent; prophylls 23.4(10.041.5) cm long,
not short and asymmetrically apiculate, the surfaces not ridged, without unequally wide ridges; peduncular
bracts 23.3(8.749.0) cm long, well-developed, inserted 17.5(4.039.0) cm long,; peduncles 44.5(12.588.5)
cm long, 2.3(1.34.2) mm in diameter; rachillae 1, 15.5(5.529.0) cm long, 3.8(2.26.5) mm in diameter, the
surfaces without spiky, fibrous projections or ridges, drying brown or yellow-brown, without short, transverse
ridges, not filiform and not narrowed between the flower pits; flower pits usually spirally arranged, glabrous
internally; proximal lips apiculate and lobed before anthesis, tearing in the center after anthesis, not recurved
after anthesis, not hood-shaped; proximal and distal lips drying the same color as the rachillae, not joined to
85
form a raised cupule, the proximal lip margins overlapping the distal lip margins; distal lips well-developed;
flowers persistent after anthesis; staminodial tubes crenulate or shallowly lobed at the apex, those of nonfertilized pistillate flowers not projecting and persistent after anthesis; fruits 8.1(6.610.7) mm long, 6.3(5.2
7.6) mm in diameter, the bases with a prominent, asymmetric stipe, the apices not conical, the surfaces not
fibers present, these coming to a point at fruit apices; locular epidermis without operculum, sculpted, usually
also with a raised, meridional ridge, without pores.
FIGURE 22. Distribution maps of Geonoma lanata, G. laxiflora, G. lehmannii subsp. lehmannii, and G. lehmannii subsp.
corrugata.
Taxonomic notes:Geonoma lehmannii is the first species dealt with here in a group of high elevation,
Andean species, the G. undata clade. This group also includes G. orbignyana, G. talamancana, G. trigona, and
G. undata. These species have been treated differently by both Wessels Boer (1968) and Henderson et al.
(1995). They are closely related and three of themG. lehmannii, G. orbignyana, and G. undataare difficult
to distinguish from one another, and extremely complex internally. Geonoma lehmannii differs from other
species in this group, except G. talamancana, in its tubular, narrow, elongate, closely sheathing, more or less
persistent prophylls and peduncular bracts which are ribbed with elongate, unbranched fibers. It differs from
G. talamancana in its well-developed peduncular bract.
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HENDERSON
Subspecific variation:Four traits vary within this species (stem branching, stem type, leaf division,
leaf plication). There is geographic discontinuity, and Central American specimens are isolated from South
American ones. All Central American specimens have plicate leaves, whereas few South American specimens
do. Specimens from Central America differ significantly from South American ones in eight variables (rachis
width, basal pinna angle, apical pinna length, apical pinna angle, prophyll length, peduncular bract length,
peduncle length, rachilla width)(t-test, P <0.05). Based on these results and geographic discontinuity, the two
subgroups are recognized as subspecies (subspp. corrugata, lehmannii).
Key to the subspecies of G. lehmannii
1
-
Peduncles 38.7(12.563.0) cm long; South America (Venezuela, Colombia, Ecuador, and Peru) ... subsp. lehmannii
Peduncles 62.9(19.088.5) cm long; western Panama .......................................................................subsp. corrugata
32a. Geonoma lehmannii subsp. lehmannii

Geonoma helminthostachys Burret (1930a: 176). Type: COLOMBIA. Antioquia: Western Cordillera, above
Ciudad Antioquia, 18002400 m, August 1891, F. Lehmann 7233 (holotype B, destroyed, isotypes F!, K!).
Geonoma acutangula Burret (1930a: 177). Type: COLOMBIA. Antioquia: Plateado, 2150 m, 6 April 1880, W. Kalbreyer
1570 (holotype B, destroyed). Neotype (designated by Bernal et al. 1989): COLOMBIA. Antioquia: Mun. Salgar,
camino de ascenso al cerro Plateado, ca. 2200 m, 4 November 1985, P. Franco et al. 2353 (neotype COL!,
isoneotype AAU, n.v.).
Geonoma parvifrons Burret (1930a: 178). Type: ECUADOR. Loja: Loja to Zamora, 15002000 m, no date, F. Lehmann
5288 (holotype B, destroyed, isotype K!).
Leaves seldom plicate. Inflorescences prophylls 20.8(10.035.0) cm long; peduncles 38.7(12.563.0) cm

long.
Distribution and habitat:From 953N1300S and 69567903W in the Andes of South America
in Venezuela, Colombia, Ecuador, and Peru at 2100(12002900) m elevation in montane rainforest (Fig. 22).
Geonoma lehmannii subsp. lehmannii is widespread and variable, and occurs in several, disjunct
populations. In Venezuela, most specimens occur in one area in Portuguesa and Trujillo, with an outlier in
Lara. They have leaves with 3(35) pinnae per side of the rachis. They are geographically isolated and differ
from the closest population in northern Colombia in nine variables.
In northern Colombia, specimens occur in the Central and Western Cordilleras only. Leaves are usually
pinnate with 4(18) pinnae per side of the rachis, although rarely they are undivided and plicate. The types of
G. lehmannii, G. helminthostachys, and G. acutangula are from this area. In Antioquia, specimens from the
Central Cordillera tend to be smaller and those from the Western Cordillera larger, although there are
exceptions. One specimen (Callejas 2138) has two inflorescences inserted at the same node. Specimens from
southern Colombia (Caquet, Huila, Putumayo) are geographically isolated but similar to those from the
Central Cordillera in Antioquia.
To the south, specimens are from scattered areas in southern Ecuador and northern Peru, central Peru, and
southern Peru. They have leaves with 3(17) pinnae per side of the. The type of G. parvifrons is from southern
Ecuador. Four specimens from the Cordillera del Condor in Ecuador (Croat 98964, Neill 14982, 15024,
Quizhpe 2222) have undivided, plicate leaves, but other specimens from the same region have non-plicate
leaves. Most specimens from Cajamarca in Peru have slender peduncles not covered by the bracts. In San
Martn, one specimen (Smith 4475) has undivided leaves with more or less parallel sides.
32b. Geonoma lehmannii subsp. corrugata Henderson, subsp. nov. (Appendix IV, Plate 39)
Geonomae lehmannii subsp. lehmannii pedunculis longioribus differt.
Type: PANAMA. Chiriqu: Cerro Pate de Macho, 849N, 8224W, 2150 m, 31 December 1985, G. de Nevers & S.
Charnley 6684 (holotype NY!, isotype MO!).
87
Leaves plicate. Inflorescences prophylls 30.8(16.541.5) cm long; peduncles 62.9(19.088.5) cm long.

Distribution and habitat:From 829852N and 81058235W in western Panama at
1715(11002500) m elevation in montane rainforest (Fig. 22). The outlying specimens are likely to be an
artifact of insufficient collecting.
There is geographical variation in this subspecies (unlike subsp. lehmannii). Regression shows there are
significant positive associations between elevation and one plant, five leaf and two inflorescence variables.
Squared multiple R for the regression of stem height on elevation is 0.38, rachis width 0.19, number of pinnae
0.22, basal pinna length 0.38, basal pinna angle 0.29, apical pinna angle 0.34, peduncular bract length 0.67,
and peduncle length 0.43. Values of all these variables increase with increasing elevation. Plants at higher
elevations have taller stems, leaves with more pinnae and wider angles, and longer inflorescences. de Nevers
and Grayum (1998) considered that there were two morphotypes of this taxon (as G. jussieuana); one with
deciduous leaf bases and narrow, undivided leaves occurring in tall forest at lower elevations, and the other
with persistent leaf bases and pinnate leaves occurring on wind-swept ridges in low forest at higher elevations.
33. Geonoma leptospadix Trail (1876: 327). Type: BRAZIL. Amazonas: Tonantins, 24 November 1874, J.
Trail 963/CLXXII (holotype K!, isotypes F!, NY!, P!).
Geonoma saramaccana Bailey (1948: 104). Type: SURINAM. Saramacca River, 9 July 1944, B. Maguire 24095
(holotype NY!).
Plants 1.5(0.53.0) m tall; stems 0.8(0.42.0) m tall, 0.7(0.50.9) cm in diameter, solitary or clustered, canelike; internodes 0.9(0.51.6) cm long, yellowish and smooth. Leaves 13(1017) per stem, undivided or rarely
long; petioles 6.6(2.513.0) cm long, drying green or yellowish; rachis 32.0(21.843.0) cm long, 2.7(1.54.3)
side of rachis; basal pinna length and width not applicable, forming an angle 20(1028) with the rachis;
apical pinna 11.4(6.315.3) cm long, width not applicable, forming an angle of 26(2035) with the rachis.
flattened, persistent; prophylls 5.9(4.310.4) cm long, not short and asymmetrically apiculate, the surfaces not
ridged, without unequally wide ridges; peduncular bracts 5.0(3.76.4) cm long, well-developed, inserted
central notch before anthesis, not recurved after anthesis, hood-shaped at anthesis, sometimes splitting postanthesis; proximal and distal lips drying the same color as the rachillae, not joined to form a raised cupule, the
proximal lip margins overlapping the distal lip margins; distal lips well-developed; staminate and pistillate
staminodial tubes crenulate or shallowly lobed at the apex, when lobed the lobes not spreading at anthesis nor
acuminate, those of non-fertilized pistillate flowers not projecting and persistent after anthesis; fruits 8.0(7.0
9.4) mm long, 7.2(6.07.8) mm in diameter, the bases without a prominent stipe, the apices not conical, the
surfaces not splitting at maturity, without fibers emerging, not bumpy and not apiculate; locular epidermis
Distribution and habitat:From 643N1304S and 46267718W in the Amazon region of
Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil, Peru, Ecuador, and Bolivia at 361(125850)
m elevation in lowland rainforest (Fig. 23).
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HENDERSON
Taxonomic notes:Geonoma leptospadix appears similar to both G. deversa and G. laxiflora. It differs
from G. deversa in its spirally arranged flower pits and from G. laxiflora in its prophylls which are not short
and asymmetrically apiculate.
Subspecific variation: No trait varies within this species except for stem branching and leaf division.
One specimen (of 47) has a pinnate leaf.
between longitude and two stem, six leaf and four inflorescence variables. Squared multiple R for the
regression of plant height on longitude is 0.15, stem diameter 0.43, petiole length 0.24, rachis length 0.37,
rachis width 0.23, basal pinna angle 0.08, apical pinna length 0.29, apical pinna angle 0.25, prophyll length
0.45, peduncle length 0.43, peduncle width 0.14, and rachillae length 0.38. From east to west, plants become
taller with thinner stems, petioles longer, and basal and apical angles wider. All other variables decrease and
inflorescences in particular become smaller.
See under Geonoma deversa for hybrids between that species and G. leptospadix.
FIGURE 23. Distribution maps of Geonoma leptospadix and G. longipedunculata.
34. Geonoma longipedunculata Burret (1930c: 8). Type: COLOMBIA. Caquet: Getuch, Ro Orteguaza, 21
July 1926, G. Woronow & S. Juzepczuk 6157 (holotype LE, n.v.).
Plants 1.5(0.83.0) m tall; stems 0.3(0.11.0) m tall, 1.7(1.32.1) cm in diameter, solitary, not cane-like;
internodes 0.4(0.30.4) cm long, not scaly. Leaves 10(613) per stem, irregularly pinnate, not plicate, bases of
blades running diagonally into the rachis; sheaths 13.4(8.020.0) cm long; petioles 61.0(43.577.5) cm long,
drying green or yellowish; rachis 46.1(26.064.0) cm long, 3.8(2.27.0) mm; veins raised and rectangular in
28.0) cm wide, forming an angle of 30(1835) with the rachis. Inflorescences branched 12 orders; prophylls
and peduncular bracts ribbed with elongate, unbranched fibers, both bracts tubular, narrow, elongate, closely
glabrous internally; proximal lips without a central notch before anthesis, not recurved after anthesis, hood-
89
shaped at anthesis, sometimes splitting post-anthesis; proximal and distal lips drying the same color as the
lips a scarcely raised rim; staminate and pistillate petals not emergent, not valvate throughout; staminate
apex, those of non-fertilized pistillate flowers not projecting and persistent after anthesis; fruits 6.0(5.96.1)
mm long, 4.9(4.65.2) mm in diameter, the bases without a prominent stipe, the apices not conical, the
surfaces not splitting at maturity, without fibers emerging, bumpy from the numerous, subepidermal,
tangential, short fibers present, these coming to a point at fruit apices; locular epidermis without operculum,
smooth, without pores.
Distribution and habitat:From 400N425S and 73507838W in the western Amazon and subAndean regions of Colombia, Ecuador, and Peru at 410(2001050) m elevation in lowland rainforest (Fig.
23).
Taxonomic notes:Geonoma longipedunculata was included as a synonym of G. dicranospadix (here
treated as a synonym of G. frontinensis) by Wessels Boer (1968). However, it is unrelated to G. frontinensis,
but similar to G. brongniartii, G. poeppigiana, and G. sanmartinensis. It differs from these in its rachillae with
faint to pronounced, short, transverse ridges.
Subspecific variation: No trait varies within this species. There is no geographic discontinuity except
for outlying specimens in Colombia and Peru.
35. Geonoma longivaginata Wendland ex Spruce (1871: 109). Type: COSTA RICA. Heredia: Sarapiqu,
Plants 3.0(1.08.0) m tall; stems 3.0(1.05.0) m tall, 1.1(0.62.1) cm in diameter, solitary or clustered, canelike; internodes 3.0(1.27.0) cm long, yellowish and smooth. Leaves 9(610) per stem, irregularly pinnate, not
58.0) cm long, drying green or yellowish; rachis 52.5(16.5101.0) cm long, 4.1(1.48.0) mm in diameter;
veins raised and rectangular in cross-section adaxially or not raised or slightly raised and triangular in crosssection adaxially; pinnae 6(213) per side of rachis; basal pinna 23.8(8.550.0) cm long, 4.1(0.734.5) cm
wide, forming an angle of 64(3192) with the rachis; apical pinna 16.6(6.731.5) cm long, 12.7(5.530.0)
cm wide, forming an angle of 38(2750) with the rachis. Inflorescences branched 12 orders; prophylls and
peduncular bracts not ribbed with elongate, unbranched fibers, flattened, deciduous; prophylls 9.0(3.316.0)
cm long, short, asymmetrically apiculate, the margins curved around the stem, the surfaces flat with dense,
felty, brown tomentum, prophyll equal to and early deciduous with the peduncular bract, the surfaces not
0.3(0.10.6) cm above the prophyll; peduncles 8.8(3.816.0) cm long, 4.9(2.39.2) mm in diameter; rachillae
6(218), 23.3(10.042.0) cm long, 3.2(1.95.3) mm in diameter, the surfaces without spiky, fibrous
projections or ridges, drying brown, with short, transverse ridges, not filiform and not narrowed between the
flower pits; flower pits spirally arranged, glabrous internally; proximal lips without a central notch before
deciduous after anthesis; stamens 6; thecae diverging at anthesis, inserted onto bifid and well-developed, nonjointed connectives; anthers short and curled over at anthesis; non-fertilized pistillate flowers deciduous after
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HENDERSON
Taxonomic notes:Geonoma longivaginata is a member of a group of three species within the G. stricta
clade, the other two being G. divisa and G. ferruginea. It differs from G. divisa in its crenulate or shallowly
lobed staminodial tube; and from G. ferruginea in its rachillae with short, transverse ridges.
Subspecific variation:Two traits vary within this species (stem branching, adaxial veins). The species
is distributed in Panama and Costa Rica and just reaches Nicaragua. It is divided into several disjunct
populations. The state distribution of one trait (adaxial veins) together with geography suggest several
subgroups.
Specimens with raised adaxial veins occur in Panama (San Blas) and are treated as a separate subgroup.
Specimens with non-raised adaxial veins occur in five areas: Atlantic slope in Costa Rica and adjacent
Nicaragua and Panama; Pacific slope in Costa Rica; El Cop, Llano Grande, and Cerro Tife in Panama; El
Valle in Panama; and the Santa Rita Ridge in Panama. Specimens from the Atlantic and Pacific slopes in
Costa Rica do not differ significantly from each other in any quantitative variable, and these are treated as one
subgroup. There are thus five potential subgroups, but there are only three specimens of one of these, from the
Santa Rita Ridge in Panama, too few to test for differences. Four potential subgroups are tested for
differences.
ANOVA shows that for pair wise comparison probabilities, all variables except leaf number and fruit
length differ significantly (P <0.05) between at least one pair of subgroups, although no variable differs
amongst all four subgroups. Based on these results and geography, the four subgroups are recognized as
subspecies. Specimens from Santa Rita Ridge in Panama are included with those from the Atlantic and Pacific
slopes in Costa Rica (see below) (subspp. copensis, longivaginata, sanblasensis, vallensis).
Key to the subspecies of G. longivaginata
1
2
3
-
Veins raised and rectangular in cross-section adaxially; Panama (San Blas) ..................................subsp. sanblasensis
Veins not raised or slightly raised and triangular in cross-section adaxially; Nicaragua, Costa Rica, and Panama..... 2
Rachis 67.2(42.0101.0) cm long; rachillae 7(418), 25.7(10.042.0) cm long...........................subsp. longivaginata
Rachis 24.9(16.537.5) cm long; rachillae 3(25), 17.5(14.025.2) cm long.............................................................. 3
Peduncles 5.2(3.87.5) cm long; El Cop, Llano Grande, Cerro Tife................................................... subsp. copensis
Peduncles 9.2(7.412.1) cm long; El Valle............................................................................................ subsp. vallensis
35a. Geonoma longivaginata subsp. longivaginata

Leaves veins not raised or slightly raised and triangular in cross-section adaxially; rachis 67.2(42.0101.0) cm
long. Inflorescences peduncles 9.5(5.016.0) cm long; rachillae 7(418), 25.7(10.042.0) cm long.
Distribution and habitat:From 8401130N and 79408425W in Nicaragua, Costa Rica
(Atlantic and Pacific slopes) and Panama at 185(51000) m elevation in lowland tropical rainforest (Fig. 24).
The outlying specimens from the Santa Rita Ridge in Panama (de Nevers 10649, Hammel 14498, Porter
4741) have fewer pinnae, wider basal pinnae, and fewer rachillae than the others, but there are too few
specimens to test for differences. Three specimens (Cooper 493, de Nevers 6864, Lewis2162) from western
Panama are smaller than others and resemble Geonoma deversa subsp. deversa (although this subspecies does
not occur in the area). They may be hybrids and are not included in the above description and analysis.
35b. Geonoma longivaginata subsp. copensis Henderson, subsp. nov. (Appendix IV, Plates 40 & 41)
A subspeciebus aliis venis haud prominentibus atque pedunculis brevioribus differt.
Type: PANAMA. Cocl: continental divide above El Cop, 838N, 8039W, 650750 m, 27 November 1985, G. de
Nevers, A. Henderson, H. Herrera, G. McPherson & L. Brako 6392 (holotype PMA!, isotypes NY!, MO!).
91
long. Inflorescences peduncles 5.2(3.87.5) cm long; rachillae 3(24), 17.0(14.020.5) cm long.
Distribution and habitat:From 837847N and 80288039W in central Panama (Cerro Tife, El
Cop, Llano Grande) at 721(2001200) m elevation in lowland to montane tropical rainforest (Fig. 24).
35c. Geonoma longivaginata subsp. sanblasensis Henderson, subsp. nov. (Appendix IV, Plates 4244)
A subspeciebus aliis venis prominentibus differt.
Type: PANAMA. San Blas: El Llano-Cart road, km 1719, 919N, 7855W, 19 June 1986, G. de Nevers & H. Herrera
7957 (holotype NY!, isotype MO, n.v.).
Leaves veins raised and rectangular in cross-section adaxially; rachis 52.8(42.064.0) cm long. Inflorescences
peduncles 11.4(8.014.0) cm long; rachillae 6(48), 27.7(22.337.0) cm long.
Distribution and habitat:From 919S924N and 78487908W in San Blas, Panama at 322(80
One specimen (de Nevers 4959excluded from the above analyses and descriptions) is considerably
smaller than the others and) may be a hybrid with Geonoma deversa.
FIGURE 24. Distribution maps of Geonoma longivaginata subsp. longivaginata, G. longivaginata subsp. copensis, G.
longivatinata subsp. sanblasensis, G. longivatinata subsp. vallensis, and G. macrostachys.
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HENDERSON
35d. Geonoma longivaginata subsp. vallensis Henderson, subsp. nov. (Appendix IV, Plate 45)
A subspeciebus aliis venis haud prominentibus atque pedunculis longioribus differt.
Type: PANAMA. Cocl: El Valle de Antn, El Monasterio, near Cerro Gaital, 27 November 1995, A. Henderson & R.
Bernal 2039 (holotype PMA!, isotype NY!).
long. Inflorescences peduncles 9.2(7.412.1) cm long rachillae 4(35), 18.8(15.425.2) cm long.
Distribution and habitat:From 837840N and 80058012W in central Panama (El Valle) at
917(880950) m elevation in lowland tropical rainforest (Fig. 24).
36. Geonoma macrostachys Martius (1823: 19). Taenianthera macrostachys (Martius) Burret (1930a: 268).
Type: BRAZIL. Amazonas: Rio Japur, no date, C. Martius s.n. (holotype M!).
Geonoma acaulis Martius (1823: 18). Taenianthera acaulis (Martius) Burret (1930a: 269). Geonoma macrostachys var.
acaulis (Martius) Henderson (1995: 274). Type: COLOMBIA. Amazonas: Rio Negro ad montem Cupati [Ro
Caquet, Cerro Yupat], no date, C. Martius s.n. (holotype M!).
Geonoma tamandua Trail (1876: 323). Taenianthera tamandua (Trail) Burret (1930a: 268). Type: BRAZIL. Amazonas:
Rio Javari, So Antonio de Boa Vista 4 December 1874, J. Trail 976/CLXXXIII (holotype K!, isotypes GH!, P!).
Geonoma acaulis subsp. tapajotensis Trail (1876: 324). Geonoma tapajotensis (Trail) Drude (1882: 508). Taenianthera
tapajotensis (Trail) Burret (1930a: 269). Type: BRAZIL. Par: Aramanahy, Rio Tapajs, 10 January 1874, J. Trail
1017/IX (holotype K!).
Taenianthera oligosticha Burret (1931a: 201). Type: PERU. Loreto: Ro Nanay, May-June 1929, L. Williams 737
(holotype F!).
Geonoma atrovirens Borchsenius & Balslev in Borchsenius et al. (2001: 342). Type: ECUADOR. Napo: Jatun Satcha
Biological Field Station, SE of Mishualli, ca. 400 m, 2 October 1995, H. Balslev et al. 6430 (holotype AAU!,
isotype QCA, n.v.), synon. nov.
Geonoma supracostata Svenning in Borchsenius et al. (2001: 344). Type: ECUADOR. Napo: E of Yasun Scientific
Station, Yasun National Park, 040S, 7623W, 23 March 1995, J.-C. Svenning 148 (holotype AAU!), synon. nov.
Geonoma ecuadoriensis Henderson, Borchsenius & Balslev (2008: 192). Type: ECUADOR. Napo: carretera HollinLoreto-Coca, km 40, entre Ro Guaman y Ro Pucuno, 040S 7738W, 1200 m, 11 December 1987, D. Neill, W.
Palacios & C. Cern 8073 (holotype NY!, isotypes AAU!, MO n.v.), synon. nov.
cane-like; internodes 0.2(0.10.4) cm long, not scaly. Leaves 9(315) per stem, undivided or irregularly
mm in diameter; veins raised and rectangular in cross-section adaxially or not raised or slightly raised and
8.8(1.428.0) cm wide, forming an angle of 31(877) with the rachis. Inflorescences unbranched; prophylls
59.2(19.2128.5) cm long, 2.9(0.76.8) mm in diameter; rachillae 1, 14.1(4.231.0) cm long, 5.87(1.712.3)
notch overlapping, not recurved after anthesis, not hood-shaped; proximal and distal lips drying the same
93
margins; distal lips well-developed; staminate and pistillate petals emergent, valvate throughout or not
emergent, not valvate throughout; staminate flowers deciduous after anthesis; stamens 6; thecae diverging or
not diverging at anthesis, inserted onto poorly to well-developed, non-split, jointed connectives, connectives
when well-developed alternately long and short; anthers short at anthesis, remaining straight and parallel;
non-fertilized pistillate flowers deciduous after anthesis; staminodial tubes lobed at the apex, the lobes
spreading at anthesis, acuminate, those of non-fertilized pistillate flowers not projecting and persistent after
subepidermal, tangential, short fibers present, these coming to a point at fruit apices; locular epidermis with
operculum, smooth or sculpted and then usually also with a raised, meridional ridge, without pores.
Distribution and habitat:From 245N1750S and 55007830W in the western Amazon region
in Venezuela, Colombia, Ecuador, Peru, and Brazil at 443(751800) m elevation in lowland or montane
Taxonomic notes:Geonoma macrostachys is the most variable species in the whole genus, and most
difficult taxonomically. It has been dealt with differently by both Wessels Boer (1968) and Henderson et al.
(1995). However, no satisfactory treatment is possible based on herbarium specimens because variation is so
complex and there is little geographic disjunction. This is discussed in some detail in the section on
Infraspecific Variation. In this treatment a broad approach is taken and only one species recognized with
various morphotypes, as discussed below.
Subspecific variation:Five traits vary within this species (stem branching, leaf division, adaxial veins,
petals, locular epidermis sculpting). Excluding two of these (stem branching, leaf division), and one for which
there are few data (locular epidermis sculpting), the state combinations of the other two traits (adaxial veins,
petals) divide the species into three subgroups (raised veins/non-exerted petals; non-raised veins/non-exerted
petals; non-raised veins/exerted petals). There is no geographic discontinuity between any of these trait
subgroups. Geonoma macrostachys is distributed widely in the central and western Amazon region. These
three subgroups are not consistent. Within each there is much quantitative variation such that there are no
clearly defined subgroups. Furthermore, there is unexplained variation in staminate flowers.
All specimens with staminate flowers are scored as having the thecae inserted onto poorly to welldeveloped, non-split, jointed connectives, when well-developed the connectives alternately long and short.
However, there appear to be two different kinds of connective, poorly-developed and well-developed, and
these two can occur together at the same locality. For example, in Ecuador (Napo), on eastern Andean slopes
at 1200 m near the Ro Guaman, Neill 8073 has poorly-developed connectives and occurs near to, and is
apparently identical with Neill 8622, which has well-developed connectives. In Peru (Madre de Dios) at low
elevations in Amazon forests near Puerto Maldonado, Gentry 68666 has poorly-developed connectives, and is
apparently identical with Timana 1588 which has well-developed connectives. Lacking staminate flowers, it
is not possible to distinguish two kinds of connective. This problem cannot be solved with the material
available. Although G. macrostachys is extremely variable, a number of morphotypes can be distinguished.
In the western Amazon region in Colombia, Ecuador, Peru, and Brazil, with an outlying population in
southern Peru, there is a morphotype (macrostachys) with usually undivided leaves, non-raised veins, and
1(14) pinnae with narrow basal angles of 10(340). Staminate flowers have well-developed connectives,
jointed at the apices of the filament, and alternately long and short. Specimens of this morphotype lacking
staminate flowers and having undivided leaves are difficult to distinguish from other morphotypes (e.g.,
atrovirens). One specimen (Daz 7058) from Peru has an exceptionally long interbract distance of 22.6 cm,
and the prophyll is longer than the peduncular bract. The mean interbract distance of macrostachys without
this specimen is 0.6 cm and the prophyll is always shorter than the peduncular bract. Specimens from
Amazonas in Peru are particularly variable. Some specimens have wider leaves and almost sigmoid venation
and others have unusually large leaves. The type of G. macrostachys is of this morphotype.
In the western and central Amazon region in Venezuela, Colombia, Ecuador, Peru, and Brazil there is a
morphotype (acaulis) having pinnate leaves, non-raised adaxial veins, 5(39) pinnae with wide basal angles
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HENDERSON
of 61(4090), and staminate flowers with poorly developed connectives. All the specimens assigned to this
morphotype have staminate flowers, but there are similar specimens, lacking staminate flowers, that are
assigned to the tapajotensis morphotype (see below). One acaulis specimen (Balick 1140) from northwest of
Iquitos has pinnae with raised veins. This occurs in an area without other acaulis but in which G. brongniartii
is abundant.
The types of G. acaulis and G. ecuadoriensis are of the acaulis morphotype. The original description and
illustration of G. acaulis by Martius (1823) showed staminate flowers with poorly-developed connectives.
However, the illustration in Drude (1882), presumably using a different specimen (possibly a Trail specimen)
than that of Martius (1823), shows the staminate flowers with well-developed, non-split connectives, these
jointed at the apices of the filament. Because of this, Geonoma acaulis was subsequently interpreted as having
Taenianthera-type staminate flowers with well-developed connectives (Wessels Boer, 1968; Henderson,
1995; Borchsenius et al., 2001).
In the western and central Amazon region in Colombia, Ecuador, Peru, Bolivia, and Brazil, with an
outlying population in Bolivia, there is a morphotype (tapajotensis) that is similar to the acaulis morphotype
in its pinnate leaves with non-raised veins and 4(210) pinnae with wide basal angles of 58(27100), but has
well-developed connectives. The types of G. acaulis subsp. tapajotensis and Taenianthera oligosticha are of
this morphotype.
Some specimens (large-size morphotype), similar to the tapajotensis morphotype, from central Peru and
Brazil (Acre), are larger in size but overlap with normalsized specimens. They have pinnate leaves with nonraised veins, 4(39) pinnae with wide basal angles of 51(2077), and well-developed connectives. They are
extremely variable and some, apart from their non-raised veins, resemble the large-raised morphotype (see
below). These large-size specimens occur sympatrically with large size specimens of G. brongniartii.
In the western Amazon region of Ecuador and adjacent Peru and Colombia there is a morphotype
(atrovirens) with undivided leaves with non-raised veins with narrow basal angles of 7(510). On specimen
labels the leaves are reported to be dark green or blackgreen. Connectives are almost intermediate between
well-developed and poorly-developed ones. However, two, outlying specimens from Colombia (Galeano
1887, 2096) have poorly-developed connectives. Without staminate flowers it is difficult to distinguish this
morphotype from the morphotypes macrostachys or tamandua. The type of G. atrovirens is of this
morphotype.
In the western Amazon region of Colombia, Ecuador, Peru, and Brazil there is a morphotype (tamandua)
with exceptionally long rachis and undivided leaves with non-raised veins with narrow basal angles of 7(3
17). The leaves are described on labels as dark green, blue green, almost black, nearly black, or dark
black-green. In leaf color this morphotype resembles the atrovirens morphotype, but has well-developed
connectives. Specimens come from two areasEcuador and adjacent Colombia, and Peru and Brazil. There is
no difference between these two populations except that specimens from Ecuador and adjacent Colombia
have wider rachillae. The type of G. tamandua is of this morphotype.
In the western Amazon region of Ecuador and Peru there is a morphotype (supracostata) having
undivided or pinnate leaves with raised adaxial veins and 2(14) pinnae with wide basal angles of 26(1048).
Staminate flowers have well-developed connectives. The type of G. supracostata is of this morphotype.
In the western Amazon region of Ecuador and Peru there is a morphotype (large-raised) with usually
divided leaves with raised adaxial veins and 7(116) pinnae and narrow basal pinna angles of 10(912).
Staminate flowers have well-developed connectives. Specimens of this morphotype lacking staminate flowers
and/or fruits are difficult to distinguish from specimens of G. multisecta and G. schizocarpa.
In the Peruvian Amazon and adjacent parts of Colombia there is a morphotype (grandiflora) with nonraised adaxial veins, 4(37) pinnae per side of the rachis, and wide basal angles of 66(4490). The staminate
and pistillate petals are emergent and valvate throughout at anthesis, and they tend to be larger than those of
other specimens. The proximal lip of the flower pit is also more pronounced. One specimen (Gentry 54536)
has 11 stamens.
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37. Geonoma maxima (Poiteau) Kunth (1841: 229). Gynestum maximum Poiteau (1822: 388). Type:
FRENCH GUIANA. Without locality, no date, A. Poiteau s.n. (holotype P!).
irregularly pinnate, if regularly pinnate the pinnae with 1 main vein only (rarely with several lateral veins), not
plicate or plicate, bases of blades running diagonally into the rachis; sheaths 12.6(4.030.0) cm long; petioles
fibers, flattened, deciduous or persistent; prophylls 10.6(3.721.8) cm long, not short and asymmetrically
apiculate, the surfaces not ridged, without unequally wide ridges; peduncular bracts 8.7(4.713.0) cm long,
well-developed, inserted 0.6(0.23.0) cm above the prophyll; peduncles 8.2(3.219.5) cm long, 4.9(1.510.0)
mm in diameter; rachillae 19(450), 11.4(4.824.3) cm long, 2.7(0.76.0) mm in diameter, the surfaces
proximal lips with a central notch before anthesis, often the two sides of the notch overlapping, not recurved
after anthesis, not hood-shaped; proximal and distal lips drying the same color as the rachillae, not joined to
form a raised cupule, the proximal lip margins overlapping the distal lip margins; distal lips well-developed;
anthesis; stamens 6; thecae diverging at anthesis, inserted directly onto the apiculate filament apices; anthers
not short and curled at anthesis, usually elongate, spiraled and twisted or sometimes remaining straight; nonfertilized pistillate flowers deciduous after anthesis; staminodial tubes lobed at the apex, the lobes spreading at
anthesis, acuminate, those of non-fertilized pistillate flowers not projecting and persistent after anthesis; fruits
11.8(6.918.2) mm long, 9.0(5.513.0) mm in diameter, apices not conical, the surfaces not splitting at
maturity, without fibers emerging, not bumpy, not apiculate; locular epidermis without operculum, smooth,
without pores.
Taxonomic notes:Geonoma maxima is a member of the G. macrostachys clade, differing from all other
species in its locular epidermis without an operculum. It also has more branched inflorescences (450
rachillae) compared to the rest of the clade (19 rachillae). It is another variable species which has been
treated differently by Wessels Boer (1968) and Henderson et al. (1995). Unlike other species complexes, for
example G. macrostachys, G. maxima exhibits rather welldefined geographic disjunction and morphological
variation, allowing for recognition of various subspecies, as explained below.
Subspecific variation:Four traits vary within this species (stem branching, leaf division, leaf plication,
pistillate flower persistence). There are very few data for stem branching, and this and leaf division are
excluded from the following analyses. Leaf plication and pistillate flower persistence divide the specimens
into three groups. However, leaf plication is difficult to score in this species and specimens with persistent
pistillate flowers may be of hybrid origin (see below). Furthermore, there is no geographic separation between
these groups. Geonoma maxima is widespread across the Amazon region, and there are isolated populations in
the Magdalena valley and Pacific coast of Colombia and eastern Andean slopes in Venezuela.
Apart from traits, one attribute (leaf division) divides the specimens into two groups. Leaves of one group
are regularly pinnate and the pinnae (at least those from the middle part of the leaf) have several main veins,
usually three to five. In the second group, all specimens have either undivided leaves, irregularly pinnate
leaves (sometimes with 1-veined pinnae present), or leaves with 1-veined pinnae (at least those from the
middle part of the leaf).
The first groupwith regularly pinnate leaves with 35-veined pinnaecan be divided into four separate
subgroups based partly on geography and partly on variables. ANOVA shows that for pair wise comparison
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HENDERSON
probabilities, 18 variables (plant height, stem diameter, internode length, petiole length, rachis length, rachis
width, basal pinna length, basal pinna width, apical pinna length, apical pinna angle, prophyll length,
interbract distance, peduncle length, peduncle width, rachilla length, rachilla width, number of rachillae, fruit
diameter) differ significantly (P <0.05) between at least one pair of subgroups, although no variable differs
amongst all four subgroups. Based on these results and geography, these four subgroups are recognized as
subspecies (subspp. hexasticha, maxima, multiramosa, sigmoidea).
The second groupwith leaves undivided, irregularly pinnate (sometimes with 1-veined pinnae present),
or regularly pinnate with 1-veined pinnaecan be divided into six separate subgroups based partly on
geography, partly on variables, and on one trait (leaf plication). ANOVA shows that for pair wise comparison
probabilities, 22 variables (plant height, stem diameter, leaf number, petiole length, rachis length, rachis
width, number of pinnae, basal pinna length, basal pinna width, basal pinna angle, apical pinna length, apical
pinna width, apical pinna angle, prophyll length, interbract distance, peduncle length, peduncle width, rachilla
length, rachilla width, number of rachillae, fruit length, fruit diameter) differ significantly (P <0.05) between
at least one pair of subgroups, although no variable differs amongst all six subgroups. Based on these results
and geography, these six subgroups are recognized as subspecies (subspp. ambigua, chelidonura,
camptoneura, compta, dispersa, spixiana).
Key to the subspecies of G. maxima
1
2
3
4
5
6
7
8
9
-
Prophylls 5.6(3.77.0) cm long; north of the Amazon region on the Pacific coast and Magdalena valley in Colombia,
and eastern Andean slopes in Venezuela................................................................................................ subsp. dispersa
Prophylls 10.9(4.021.8) cm long; Amazon region...................................................................................................... 2
Leaves regularly pinnate with 35-veined pinnae (except for basal and apical ones); rachis 68.4(25.5120.0) cm
long; pinnae 19(431) per side of rachis ..................................................................................................................... 3
Leaves undivided, irregularly pinnate (sometimes with 1-veined pinnae present), or regularly pinnate with 1-veined
pinnae; rachis 36.6(7.398.0) cm long; pinnae 5(129) per side of rachis................................................................... 6
Rachillae 36(2147); Amazonian Ecuador and adjacent Colombia (Caquet, Putumayo) and Peru (Loreto) ..............
........................................................................................................................................................subsp. multiramosa
Rachillae 21(650); Amazonian Colombia, Venezuela, Peru, Brazil, Bolivia, and the Guianas ................................. 4
Rachillae 28(1642), 1.0(0.71.2) mm in diameter; Amazonian Colombia (Amazonas, Caquet).. subsp. sigmoidea
Rachillae 21(650), 2.5(1.53.6) mm in diameter; Amazonian Colombia (Guainia), Venezuela, Brazil, Bolivia, and
the Guianas................................................................................................................................................................... 5
Rachillae 13(625); southern Venezuela and adjacent Colombia (Guainia) and Brazil (Amazonas) ............................
............................................................................................................................................................subsp. hexasticha
Rachillae 31(950); central and eastern Amazon region of Brazil and the Guianas.............................. subsp. maxima
Leaves regularly pinnate with 1-veined pinnae; rachis 44.5(11.087.0) cm long; basal pinna 0.3(0.10.5) cm wide;
mostly south or west of the Amazon in Colombia, Brazil, Peru, and Bolivia ........................................ subsp. compta
Leaves undivided or irregularly pinnate (sometimes with 1-veined pinnae present); rachis 35.6(7.398.0) cm long;
basal pinna 6.1(0.234.0) cm wide; widespread .......................................................................................................... 7
Leaves plicate; basal pinna forming an angle of 13(420) with the rachis; centralwestern Amazon region of Brazil
and adjacent Colombia .......................................................................................................................... subsp. spixiana
Leaves not plicate; basal pinna forming an angle of 41(1087) with the rachis ........................................................ 8
Rachis 51.9(38.070.0) cm long; western Amazonian Brazil (Acre), Peru, and Bolivia ..............subsp. camptoneura
Rachis 28.4(7.363.0) cm long; central-western Amazon region of Brazil and adjacent Colombia, Peru, and Bolivia,
and the Guianas ............................................................................................................................................................ 9
Rachis 40.0(13.059.0) cm long; Guyana and adjacent Venezuela and Brazil ..................................... subsp. ambigua
Rachis 25.0(7.363.0) cm long; central-western Amazon region of Venezuela, Colombia, Brazil, Peru, and Bolivia .
..........................................................................................................................................................subsp. chelidonura
37a. Geonoma maxima subsp. maxima

Geonoma multiflora Martius (1823: 7). Lectotype (designated by Wessels Boer 1968): BRAZIL. Par: without locality,
no date, C. Martius s.n. (lectotype M!).
Geonoma paraensis Spruce (1871: 112). Type: BRAZIL. Par: near Belm, no date, R. Spruce 69 (holotype K!).
97
Leaves regularly pinnate with 35-veined pinnae (except for basal and apical ones), not plicate; rachis
77.9(49.5120.0) cm long; pinnae 18(431) per side of rachis; basal pinna 0.7(0.22.5) cm wide, forming an
angle of 57(3480) with the rachis. Inflorescences rachillae 31(950).
Distribution and habitat:From 510N558S and 46306011W in central and eastern Amazon
region of Brazil, Suriname, French Guiana, with outliers in Guyana and Brazil at 288(50700) m elevation in
A specimen from French Guiana (de Granville 16838) differs from the others in its narrower pinnae.
Another specimen from French Guiana (de Granville 13389) has only four pinnae per side of the rachis. Both
these specimens approach subsp. ambigua in their leaf morphology. The geographically isolated specimens
(Henderson 649, 664, 1055, 1075, 1163, Moore 9534, Prance 2239) from near Manaus in Brazil have longer
rachis, more pinnae, and narrower apical pinnae with narrower angles. However, there are too few specimens
to test these differences, and it is not clear if the gap between them and other specimens is an artifact caused
by incomplete collecting.
FIGURE 25. Distribution maps of Geonoma maxima subsp. maxima, G. maxima subsp. ambigua, G. maxima subsp.
camptoneura, and G. maxima subsp. chelidonura.
37b. Geonoma maxima subsp. ambigua (Spruce) Henderson, comb. & stat. nov.
Basionym: Geonoma ambigua Spruce (1871: 111). Geonoma maxima var. ambigua (Spruce) Henderson (1995: 278).
Type: GUYANA. Without locality, no date, C. Appun 566 (holotype K!).
Geonoma schomburgkiana Spruce (1871: 111). Type: GUYANA. Without locality, 1837, R. Schomburgk 705 (holotype
K!, excluding leaf).
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HENDERSON
Leaves irregularly pinnate (sometimes with 1-veined pinnae present), not plicate; rachis 40.0(13.059.0) cm
long; pinnae 4(212) per side of rachis; basal pinna 7.5(1.922.0) cm wide, forming an angle of 40(2060)
with the rachis. Inflorescences rachillae 18(539).
Distribution and habitat:From 155834N and 57526255W in Guyana and adjacent
Venezuela and Brazil at 337(35900) m elevation in lowland rainforest in a variety of habitats, including
flooded and non-flooded areas (Fig. 25).
A specimen from Guyana (McDowell 4479) has more pinnae than others and approaches subsp. maxima
in its leaf morphology. Although subsp. ambigua has fewer pinnae per side of the rachis than subsp. maxima,
it usually has a few, 1-veined pinnae present.
37c. Geonoma maxima subsp. camptoneura (Burret) Henderson, comb. & stat. nov.
Basionym: Geonoma camptoneura Burret (1931a: 201). Type: PERU. Loreto: Ro Huallaga, Yurimaguas, March 1930,
L. Williams 7836 (holotype B, destroyed, isotype F!, excluding leaf)[The type specimen appears to be a mixture, the
leaf may belong to another species, possibly G. deversa].
Leaves undivided or irregularly pinnate (sometimes with 1-veined pinnae present), not plicate; rachis
Distribution and habitat:From 3501345S and 68097822W in the southwestern Amazon
region of Peru and adjacent areas of Brazil and Bolivia at 334(135800) m elevation in lowland rainforest
(Fig. 25).
37d. Geonoma maxima subsp. chelidonura (Spruce) Henderson, comb. & stat. nov.
Basionym: Geonoma chelidonura Spruce (1871: 111). Geonoma maxima var. chelidonura (Spruce) Henderson (1995:
279). Type: BRAZIL. Amazonas: Rio Uaups, November 1852, R. Spruce 73 (holotype K!).
Geonoma tuberculata Spruce (1871: 112). Geonoma spruceana subsp. intermedia var. tuberculata Trail (1876: 329).
Type: BRAZIL. Amazonas: Rio Negro, no date, R. Spruce 18 (holotype K!).
Geonoma densiflora Spruce (1871: 112). Type: BRAZIL. Amazonas: So Gabriel, near Gamas sitio, March 1852, R.
Spruce 30 (holotype K!).
Geonoma personata Spruce (1871: 112). Type: BRAZIL. Amazonas: Serra of So Gabriel, June 1852, R. Spruce 34
(holotype K!).
Geonoma densiflora var. monticola Spruce (1871: 118). Type: BRAZIL. Amazonas: So Gabriel, June 1852, R. Spruce
33 (holotype K!).
Geonoma spruceana Trail (1876: 328). Type: BRAZIL. Par: Lago Juruty, 3 April 1874, J. Trail 1002/XXIV (holotype
K!, isotype BM!).
Geonoma spruceana subsp. spruceana var. heptasticha Trail (1876: 329). Type: BRAZIL. Amazonas: Rio Negro at
Assutuba, 6 July 1874, J. Trail 1007/XCIII (holotype K!, isotype BM!).
Geonoma spruceana subsp. intermedia var. major Trail (1876: 330). Type: BRAZIL. Amazonas: Rio Solimes, Coary,
16 October 1874, J. Trail 984/CXLIV (holotype K! isotypes BM!, NY!).
Geonoma juruana Dammer (1907: 119). Type: BRAZIL. Acre: Rio Juru, Juru-mirim, August 1901, E. Ule 5744
(holotype B, destroyed, isotypes F!, MG!).
Geonoma longisecta Burret (1930a: 257). Type: PERU. Loreto: Iquitos, 4 May 1925, G. Tessmann 5087 (holotype B,
destroyed, isotype, NY!).
Geonoma parvisecta Burret (1930b: 1018). Type: BRAZIL. Amazonas: Rio Negro, So Pedro do Uaups, 23 September
1928, P. Luetzelburg 22278 (holotype B, destroyed, isotypes M!, NY!).
Leaves undivided or irregularly pinnate (sometimes with 1-veined pinnae present), not plicate; rachis
Distribution and habitat:From 203N1145S and 55007550W in the centralwestern Amazon
region of Venezuela, Colombia, Brazil, Peru, and Bolivia at 160(48525) m elevation in lowland rainforest
(Fig. 25).
This is the most variable subspecies with several local morphotypes.
99
Some specimens (large fruit morphotype) in the western part of the range in Peru and Colombia,
especially from around Iquitos in Peru, have larger fruits (mean fruit length 14.0 mm, mean fruit diameter
11.0 mm) than those of other specimens (mean fruit length 11.0 mm, mean fruit diameter 8.2 mm).
Specimens (parvisecta morphotype) from scattered areas in Colombia, Venezuela, Peru, and Brazil,
especially in the upper Rio Negro region, often in black water river areas on sandy soils (campina, catinga),
are smaller than others and often have undivided leaves. The types of Geonoma chelidonura and G. parvisecta
are of this morphotype.
Most specimens (e.g., Henderson 1517, Trail 989, 991, 998, 1002, 1005, 1007)(intermediate morphotype)
from the eastern part of the range in Brazil, occurring in flooded areas near the main Amazon river (vrzea),
have 1-veined pinnae interspersed between the wider basal and apical pinnae, and in some cases resemble the
sympatric subsp. compta.
A few specimens from widely scattered localities (Balick 942, Gruezmacher 45, Kuhlmann 1237, Le Fiell
4, Moore 8428, Pipoly 15638, Prance 7596, Rimachi 10965, 11275, Rudas 3101)(densiflora morphotype) in
Peru, Colombia, Venezuela, and Brazil (including an unmapped specimen, Kuhlmann 1237, from Tocantins,
Par) have thinner textured, sigmoid pinnae, few, thicker rachillae, and persistent pistillate flowers. The types
of G. densiflora, G. densiflora var. monticola, and G. personata have these kinds of leaves and inflorescences.
The specimens often occur sympatrically with subsp. chelidonura and subsp. compta and may be of hybrid
origin.
37e. Geonoma maxima subsp. compta (Trail) Henderson, comb. & stat. nov.
Basionym: Geonoma spruceana subsp. intermedia var. compta Trail (1876: 329). Type: BRAZIL. Amazonas: Barcellos,
30 June 1874, J. Trail 997/LXXXIV (holotype K!).
Geonoma spruceana subsp. intermedia var. intermedia Trail (1876: 329). Type: BRAZIL. Amazonas: Lago Cerrado, Rio
Juru, 30 October 1874, J. Trail 989/CXLVII (holotype K!).
Leaves regularly pinnate with 1-veined pinnae, not plicate; rachis 44.5(11.087.0) cm long; pinnae 18(629)
per side of rachis; basal pinna 0.3(0.10.5) cm wide, forming an angle of 58(3877) with the rachis.
Inflorescences rachillae 19(440).
Distribution and habitat:From 111N1110S and 56007740W in the central and western
Amazon region in Colombia, Brazil, Peru, and Bolivia at 193(122400) m elevation in lowland rainforest
(Fig. 26).
Some specimens from the Colombian Amazon (e.g., Galeano 1973, Garca-Barriga 15005, Zarucchi
1712) have wider pinnae than the others, more like those of subsp. hexasticha. One specimen from Amazonas
in Peru (Daz 7131) appears intermediate between subsp. compta and subsp. camptoneura.
37f. Geonoma maxima subsp. dispersa Henderson, subsp. nov. (Appendix IV, Plates 4648)
A subspeciebus aliis prophyllo breviore differt.
Type: COLOMBIA. Crdoba: Mun. Tierralta, entre los ros Esmeraldas y Sin, 2 km arriba de la confluencia, 200 m, 26
July 1986, R. Bernal, G. Galeano & D. Restrepo 1148 (holotype, COL!).
Leaves irregularly pinnate (sometimes with 1-veined pinnae present), not plicate; rachis 53.0(34.080.0) cm
long; pinnae 8(710) per side of rachis; basal pinna 4.7(2.07.5) cm wide, forming an angle of 49(4160)
with the rachis. Inflorescences rachillae 21(1236).
Distribution and habitat:From 549745N and 71477730W in the northern Choc and
Magdalena valley of Colombia and eastern Andean slopes in Venezuela at 245(15700) m elevation in
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HENDERSON
FIGURE 26. Distribution maps of Geonoma maxima subsp. compta, G. maxima subsp. dispersa, G. maxima subsp.
hexasticha, and G. maxima subsp. multiramosa.
37g. Geonoma maxima subsp. hexasticha (Spruce) Henderson, comb. & stat. nov.
Basionym: Geonoma hexasticha Spruce (1871: 110). Type: BRAZIL. Amazonas: near Gamas sitio, So Gabriel, March
1852, R. Spruce 29 (holotype K!, isotypes NY!, P!).
Geonoma negrensis Spruce (1871: 113). Type: VENEZUELA. Amazonas: Ro Negro, San Carlos, September 1853, R.
Spruce 70 (holotype K!).
Distribution and habitat:From 349N070S and 65076800W in southern Venezuela and
adjacent Colombia (Guainia) and Brazil (Amazonas) at 214(651000) m elevation in lowland rainforest
usually near streams or rivers on flooded or non-flooded soils (Fig. 26).
37h. Geonoma maxima subsp. multiramosa Henderson, subsp. nov. (Appendix IV, Plates 49 & 50)
A subspeciebus aliis rachillis magis numerosis differt.
Type: ECUADOR. Napo: Aangu, south bank of Ro Napo, 95 km downstream from Coca, 032S, 7623W, 300 m, 28
July 1985, H. Balslev, A. Barfod, A. Henderson, F. Skov & A. Argello 60731 (holotype NY!. isotype AAU!).
101
Distribution and habitat:From 203N235S and 75277720W in Amazonian Ecuador and
adjacent Colombia (Caquet, Putumayo) and Peru (Loreto) at 283(200450) m elevation in lowland rainforest
(Fig. 26).
37i. Geonoma maxima subsp. sigmoidea Henderson, subsp. nov. (Appendix IV, Plate 51)
A subspeciebus aliis angulo pinnarum basalibus amplus differt.
Type: COLOMBIA. Amazonas: corregimiento de Araracuara, carretera a Puerto Arturo, sitio Gucheros, ca. 360 m,
18 September 1987, G. Galeano & J. Huitoto 1279 (holotype COL!, isotype NY!).
in Colombia (Amazonas, Caquet) at 270(200360) m elevation in lowland rainforest (Fig. 27).
One specimen (Galeano 2080) has larger leaves and exceptionally large fruits (13.3 cm long and 11.5 cm
diameter), and may be a hybrid with a morphotype of subsp. chelidonura.
FIGURE 27. Distribution maps of Geonoma maxima subsp. sigmoidea and G. maxima subsp. spixiana.
37j. Geonoma maxima subsp. spixiana (Martius) Henderson, comb. & stat. nov.
Basionym: Geonoma spixiana Martius (1823: 15). Geonoma maxima var. spixiana (Martius) Henderson (1995: 281).
Type: BRAZIL. Amazonas: Rio Japur, no date, C. Martius s. n. (holotype M!).
Leaves undivided or irregularly pinnate (sometimes with 1-veined pinnae present), plicate; rachis 70.7(44.5
98.0) cm long; pinnae 2(14) per side of rachis; basal pinna 27.8(20.034.0) cm wide, forming an angle of
13(420) with the rachis. Inflorescences rachillae 14(830).
Distribution and habitat:From 118710S and 60007200W in the centralwestern Amazon
region of Brazil and Colombia at 276(230300) m elevation in lowland rainforest (Fig. 27).
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HENDERSON
38. Geonoma monospatha de Nevers in de Nevers & Grayum (1998: 98). Type: PANAMA. Veraguas: Cerro
Tute, just W of Santa F, 840N, 8105W, 8001000 m, 27 February 1995, G. de Nevers, A. Henderson, G.
Galeano & R. Bernal 10556 (holotype PMA n.v., isotypes CAS n.v. COL! K! MO! NY!).
Plants 1.1(1.01.5) m tall; stems height no data, 0.5(0.30.7) cm in diameter, solitary or clustered, cane-like;
internodes 1.4(0.44.0) cm long, yellowish and smooth. Leaves undivided or irregularly pinnate, not plicate,
long, drying green or yellowish; rachis 15.2(9.023.2) cm long, 1.6(0.92.2) mm in diameter; veins not raised
or slightly raised and triangular in cross-section adaxially; pinnae 2(14) per side of rachis; basal pinna
unbranched or branched 1 order; prophylls and peduncular bracts not ribbed with elongate, unbranched fibers,
flattened, more or less persistent; prophylls 5.1(2.27.4) cm long, not short and asymmetrically apiculate, the
surfaces not ridged, without unequally wide ridges; peduncular bracts 0.1(0.10.2) cm, long, vestigial,
inserted 1.7(0.92.8) cm above the prophyll; peduncles 5.4(3.87.0) cm long, 1.6(1.22.0) mm in diameter;
narrowed between the flower pits; flower pits spirally arranged, glabrous internally; proximal lips with a
central notch before anthesis, often the two sides of the notch overlapping, not recurved after anthesis, not
cupule, the proximal lip margins overlapping the distal lip margins; distal lips well-developed; staminate and
pistillate petals not emergent, not valvate throughout; staminate flowers deciduous after anthesis; stamens 6;
thecae diverging at anthesis, inserted almost directly onto the filament apices, the connectives bifid but
scarcely developed; anthers short and curled over at anthesis; non-fertilized pistillate flowers persistent after
anthesis; staminodial tubes lobed, the lobes not spreading at anthesis, not acuminate, those of non-fertilized
pistillate flowers not projecting and persistent after anthesis; fruits 6.7(6.17.2) mm long, 5.2(4.95.5) mm in
coming to a point at fruit apices; locular epidermis without operculum, smooth, without pores.
Distribution and habitat:From 831942N and 80268412W in Costa Rica and Panama at
1020(3001900) m elevation in lowland to montane rainforest (Fig. 28).
Taxonomic notes:Geonoma monospatha is a member of the G. cuneata clade, within which it belongs
to a group of four Central American species, including G. brenesii, G. hugonis, and G. epetiolata. They all
have unbranched or few-branched inflorescences and share the character state of the staminodial tubes being
lobed at the apex, but the lobes are not spreading at anthesis and are not acuminate. Geonoma monospatha
differs from other species in this group by its vestigial peduncular bract and its flattened prophylls and
peduncular bracts which are not ribbed with elongate, unbranched fibers.
Subspecific variation:Three traits (stem branching, leaf division, inflorescence branching) vary within
this species. Specimens come from several separate areas but there are too few specimens to test for
differences, and the gaps may be artifacts caused by incomplete collecting. The Costa Rican specimens have
smaller leaves and inflorescences and come from higher mean elevations (1750 m versus 837 m).
39. Geonoma mooreana de Nevers & Grayum (1995: 354). Type: PANAMA. Veraguas: vicinity of Santa F,
along road between Santa F and Calovbora, 1.7 mi. past Alto Piedra School, 1.5 mi. beyond Quebrada
Cosilla (previously referred to as Ro Primero Braso), 833N, 8108W, 570 m, 13 July 1994, T. Croat &
Guanghua Zhu 76826 (holotype MO!, isotypes CAS n.v., PMA!).
1.7(1.02.8) cm long, yellowish and smooth. Leaves 9 per stem, regularly pinnate and the pinnae with 1 main
103
vein only, not plicate, bases of blades running diagonally into the rachis; sheaths 17.1(11.522.0) cm long;
mm in diameter veins raised and rectangular in cross-section adaxially; pinnae 33(2649) per side of rachis;
basal pinna 25.4(14.038.0) cm long, 0.5(0.30.5) cm wide, forming an angle of 68(4789) cm with the
rachis; apical pinna 13.7(10.016.7) cm long, 1.2(0.52.8) cm wide, forming an angle of 26(1540) with the
unbranched fibers, flattened, deciduous or persistent; prophylls 12.8(8.120.9) cm long, short, asymmetrically
apiculate, the margins curved around the stem, the surfaces flat with dense, felty, brown tomentum, prophyll
equal to and early deciduous with the peduncular bract, the surfaces not ridged, without unequally wide
ridges; peduncular bracts 12.2(8.019.0) cm long, well-developed, inserted 0.4(0.30.7) cm above the
prophyll; peduncles 11.5(7.516.0) cm long, 6.0(4.28.5) mm in diameter; rachillae 108(78136), 13.2(11.0
15.0) cm long, 0.8(0.31.0) mm in diameter, the surfaces without spiky, fibrous projections or ridges, drying
brown, with faint to pronounced, short, transverse ridges, filiform with extended narrowed sections between
the flower pits; flower pits alternately arranged (sometimes distorted by twisting and contracting of rachillae),
epidermis without operculum, smooth, without pores.
Distribution and habitat:From 831920N and 80378235W in western Panama at 607(180
Taxonomic notes:Geonoma mooreana is closely related to G. scoparia from Costa Rica. It differs from
this species in its yellowish, smooth internodes.
Subspecific variation: No trait varies within this species. Specimens come from three separate areas
but there are too few specimens to test for differences, and the gaps may be artifacts caused by insufficient
collecting.
40. Geonoma multisecta (Burret) Burret (1940a: 24). Taenianthera multisecta Burret (1930c: 13). Type:
COLOMBIA. Caquet: Sucre, 10 July 1926, G. Woronow & J. Juzepczuk 5862 (holotype LE!).
Geonoma polyandra Skov (1994: 39). Type: ECUADOR. Napo: Aangu, 032N, 7623W, 300 m, 19 June 1985, H.
Balslev, A. Barfod, & F. Skov 60536 (holotype AAU!, isotypes COL n.v., K!, NY!, QCA n.v.), synon. nov.
Plants 2.7(1.53.5) m tall; stems 0.8(0.22.3) m tall, in diameter no data, solitary; internodes length no data,
not scaly. Leaves 11(916) per stem, irregularly pinnate, not plicate, bases of blades running diagonally into
the rachis; sheaths 22.0(11.038.0) cm long; petioles 41.2(13.077.0) cm long, drying green or yellowish;
rachis 74.6(55.594.0) cm long, 4.3(2.98.0) mm in diameter; veins raised and rectangular in cross-section
adaxially; pinnae13(318) per side of rachis; basal pinna 29.0(21.054.0) cm long, 2.5(0.218.5) cm wide,
forming an angle of 56(2074) with the rachis; apical pinna 20.6(15.529.5) cm long, 5.7(2.017.0) cm
the surfaces not ridged, without unequally wide ridges; peduncular bracts 29.3(25.039.5) cm long, well-
104
HENDERSON
developed, inserted 0.8(0.22.5) cm above the prophyll; peduncles 60.8(31.099.5) cm long, 3.7(2.35.1) mm
not narrowed between the flower pits; flower pits spirally arranged, glabrous internally; proximal lips with a
pistillate petals not emergent, not valvate throughout; staminate flowers deciduous after anthesis; stamens
more than 6; thecae diverging or not diverging at anthesis, inserted onto poorly to well-developed, non-split,
jointed connectives, connectives when well-developed alternately long and short; anthers short at anthesis,
remaining straight and parallel; non-fertilized pistillate flowers deciduous after anthesis; staminodial tubes
lobed at the apex, the lobes spreading at anthesis, acuminate, those of non-fertilized pistillate flowers not
emerging, not bumpy, not apiculate; locular epidermis with operculum, smooth, without pores.
of Colombia and Ecuador at 338(2001150) m elevation in lowland or) montane rainforest (Fig. 28).
FIGURE 28. Distribution maps of Geonoma monospatha, G. mooreana, G. multisecta, and G. occidentalis.
Taxonomic notes:Skov (1994), in describing Geonoma polyandra, overlooked the earlier name of G.
multisecta. This was understandable given that the type specimen of G. multisecta lacked staminate flowers.
105
However, this specimen, with its veins raised and rectangular in cross-section adaxially and leaf and
inflorescence morphology certainly represents the earliest name for this species. Geonoma multisecta is a
member of a group of related species including G. macrostachys, G. paradoxa, G. poiteauana, and G.
schizocarpa, and is the only one of which to have more than six stamens.
Subspecific variation: No trait varies within this species, nor is there any geographic disjunction.
41. Geonoma occidentalis (Henderson) Henderson, comb. & stat. nov.

Basionym: Geonoma brevispatha var. occidentale Henderson (1995: 264). Type: PERU. Madre de Dios: Ro Tambopata,
Explorers Inn at junction with Ro La Torre, 1250S, 6917W, 3 November 1991, A. Henderson & F. Chvez 1633
(holotype USM!, isotypes CUZ!, NY!).
irregularly pinnate, not plicate, bases of blades running diagonally into the rachis; sheaths 15.0 cm long;
side of rachis; basal pinna 23.6(17.532.0) m long, 2.8(1.06.0) cm wide, forming an angle of 60(3390)
with the rachis; apical pinna 14.4(11.517.5) cm long, 14.8(8.823.4) cm wide, forming an angle of 36(30
42) with the rachis. Inflorescences branched 23 orders; prophylls and peduncular bracts not ribbed with
elongate, unbranched fibers, flattened, deciduous or persistent; prophylls 9.7(6.111.0) cm long, short,
asymmetrically apiculate, the margins curved around the stem, the surfaces flat with dense, felty, brown
tomentum, prophyll equal to and early deciduous with the peduncular bract, the surfaces not ridged, without
unequally wide ridges; peduncular bracts 9.5 cm long, well-developed, inserted 0.3(0.10.5) cm above the
prophyll; peduncles 6.4(3.39.5) cm long, 6.8(3.810.2) mm in diameter; rachillae 24(1333), 25.0(15.0
brown, with faint to pronounced, short, transverse ridges, not filiform and not narrowed between the flower
pits; flower pits decussately arranged throughout the rachillae, the groups of pits closely spaced, glabrous
anthesis, sometimes splitting post-anthesis; proximal and distal lips drying the same color as the rachillae,
joined laterally with no clear gap between them, often forming a raised cupule, the margins not overlapping;
Distribution and habitat:From 4291750S and 62537335W in the western Amazon region in
Brazil, Peru, and Bolivia at a mean elevation of 312(150950) m in lowland rainforest (Fig. 28).
Taxonomic notes:This taxon was treated as a variety of Geonoma brevispatha by Henderson (1995). In
the present study, this name is treated as an excluded name, and the taxon to which Henderson was referring is
called Geonoma pohliana subsp. weddelliana. The similarity between the two taxa is superficial and there are
many differences between them, notably in the prophylls (not available to Henderson, 1995) and in the fruits.
Subspecific variation:Two traits vary within this species (stem branching, stem type). There is no
geographic disjunctionthe gap in Bolivia is likely to be an artifact of insufficient collecting. There are three
outlying specimens from Brazil (Acre, Rondnia) and Peru (Loreto).
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HENDERSON
42. Geonoma oldemanii de Granville (1975: 553). Type: FRENCH GUIANA. St. Georges dOyapock,
Crique Cabaret, 1 October 1973, J.-J. de Granville 1992 (holotype CAY n.v., isotypes P!, US!).
internodes 1.7(0.82.2) cm long, yellowish and smooth. Leaves 14(1219) per stem, undivided, not plicate,
bases of blades recurved against the rachis; sheaths 14.0(10.017.5) cm long; petioles 9.1(1.018.0) cm long,
drying green or yellowish; rachis 125.9(107.5149.0) cm long, 6.5(4.39.7) mm in diameter; veins not raised
or slightly raised and triangular in cross-section adaxially; pinnae 1 per side of rachis; basal pinna length and
width not applicable, forming an angle of 9(415) with the rachis; apical pinna 26.8(19.038.0) cm long,
width not applicable, forming an angle of 16(1218) with the rachis. Inflorescences unbranched; prophylls
glabrous internally; proximal lips with a central notch before anthesis, often the two sides of the notch
overlapping, not recurved after anthesis, not hood-shaped; proximal and distal lips drying the same color as
the rachillae, not joined to form a raised cupule, the proximal lip margins overlapping the distal lip margins;
flowers deciduous after anthesis; stamens 6; thecae diverging at anthesis, inserted directly onto the apiculate
filament apices; anthers not short and curled at anthesis, usually elongate, spiraled and twisted or sometimes
remaining straight; non-fertilized pistillate flowers deciduous after anthesis; staminodial tubes lobed at the
apex, the lobes spreading at anthesis, acuminate, those of non-fertilized pistillate flowers not projecting and
prominent stipe, the apices not conical, the surfaces not splitting at maturity, without fibers emerging, not
bumpy, not apiculate; locular epidermis with operculum, smooth, with pores.
Distribution and habitat:From 015S503N and 50305630W in the eastern Amazon region in
French Guiana and Brazil at 51(20100) m elevation in coastal, terra firme forest or lowland rainforest (Fig.
29).
Taxonomic notes:Geonoma oldemanii is related to G. triglochin and G. umbraculiformis. It differs
from these in its prophylls and peduncular bracts which are ribbed with elongate, unbranched fibers, and both
bracts are tubular, narrow, elongate, closely sheath the peduncle, and are more or less persistent
Subspecific variation: No trait varies within this species. There is geographic discontinuity, and
specimens come from three areasFrench Guiana, Maraj, and the Rio Trombetas, the last two in Brazil.
There are too few specimens to test for differences, and the gaps are likely to be artifacts of insufficient
collecting.
One specimen from French Guiana (Roncal 396not included in the above description) is much smaller
than other specimens, and resembles G. stricta subsp. pliniana, with which it occurs.
43. Geonoma oligoclona Trail (1876: 325). Type: BRAZIL. Amazonas: Barreira Branca, Rio Juta, 31
January 1875, J. Trail 1019/CCI (holotype K!, isotypes GH!, P!).
internodes 2.3(1.14.1) cm long, covered with reddish or brownish scales, especially in their distal part.
107
36.5) cm long, 2.5(1.43.7) mm in diameter; veins raised and rectangular in cross-section adaxially; pinnae 3
per side of rachis; basal pinna 17.7(6.223.5) cm long, 2.0(1.04.5) cm wide, forming an angle of 72(4397)
with the rachis; apical pinna 16.6(5.521.5) cm long, 11.2(5.016.7) cm wide, forming an angle of 35(2242)
with the rachis. Inflorescences branched 1 order; prophylls and peduncular bracts not ribbed with elongate,
unbranched fibers, flattened, deciduous; prophylls 7.0(4.210.0) cm long, short, asymmetrically apiculate, the
margins curved around the stem, the surfaces flat with dense, felty, brown tomentum, prophyll equal to and
early deciduous with the peduncular bract, the surfaces not ridged, without unequally wide ridges; peduncular
bracts 5.6(4.07.0) cm long, well-developed, inserted 0.4(0.20.5) cm above the prophyll; peduncles 6.0(4.0
diameter, the surfaces with spiky, fibrous projections or ridges, drying brown or yellow-brown, without short,
transverse ridges, not filiform and not narrowed between the flower pits; flower pits spirally arranged,
thecae diverging at anthesis, inserted onto bifid and well-developed, non-jointed connectives; anthers short
and curled over at anthesis; non-fertilized pistillate flowers deciduous after anthesis; staminodial tubes lobed
at the apex, the lobes not spreading at anthesis, not acuminate, those of non-fertilized pistillate flowers not
emerging; fruit surfaces not bumpy, not apiculate; locular epidermis without operculum, smooth, without
pores.
Distribution and habitat:From 457N322S and 63017050W in the centralwestern Amazon
region of Colombia, Venezuela, and Brazil (including an unmapped specimen, Kuhlmann 1236, from
Tocantins, Par) at 206(100250) m elevation in lowland rainforest (Fig. 29).
Taxonomic notes:This species was considered by Wessels Boer (1968) to be related to Geonoma
deversa. In fact, the two are not related, and G. oligoclona forms a clade with G. aspidiifolia and G.
santanderensis. All three species share internodes covered with reddish or brownish scales, rachillae surfaces
with spiky, fibrous projections or ridges, and staminodial tubes lobed at the apex with the lobes not spreading
at anthesis and not acuminate. Geonoma oligoclona differs from G. aspidiifolia and G. santanderensis in its
prophyll which has the margins curved around the stem and the surfaces flat with dense, felty, brown
tomentum.
Subspecific variation: No trait varies within this species. The specimens come from scattered
localities, but this is probably an artifact of insufficient collecting. One specimen (Galeano 1869) is much
smaller than the others and is reported to come from a white sand savanna area.
44. Geonoma operculata Henderson, sp. nov. (Appendix IV, Plate 52)
A speciebus affinibus operculo differt.
Type: VENEZUELA. Miranda: Cordillera de la Costa, al noreste de Guatire, Fila Juan Torres-Fila Las Perdices, por el
ro Guayabal hacia el pueblo Guayabal, 1031N, 6620W, 700900 m, 1922 February 1993, W. Meier 3401
(holotype NY!, isotype VEN n.v.).
Plants 2.0 m tall; stems height and in diameter no data; internodes yellowish and smooth. Leaves irregularly
pinnate, not plicate, bases of blades running diagonally into the rachis; sheaths no data; petioles drying green
or yellowish; rachis 23.0 cm long; veins not raised or slightly raised and triangular in cross-section adaxially;
pinnae 3 per side of rachis; basal pinna length, width and angle no data; apical pinna 12.5 cm long, 9.0 cm
wide, forming an angle of 34 with the rachis. Inflorescences branched 2 orders; prophylls and peduncular
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HENDERSON
bracts not ribbed with elongate, unbranched fibers, flattened, deciduous or persistent; prophylls length 4 cm
long, short, asymmetrically apiculate, the margins curved around the stem, the surfaces flat with dense, felty,
brown tomentum, prophyll equal to and early deciduous with the peduncular bract, the surfaces not ridged,
without unequally wide ridges; peduncular bracts no data; peduncles 12.0 cm long, 3.2 mm in diameter;
rachillae 37, 15.0 cm long, 0.8 mm in diameter, the surfaces without spiky, fibrous projections or ridges,
drying brown, with faint to pronounced, short, transverse ridges, filiform with extended narrowed sections
between the flower pits; flower pits alternately arranged (sometimes distorted by twisting and contracting of
rachillae), glabrous internally; proximal lips without a central notch before anthesis, not recurved after
anthesis, not hood-shaped; proximal and distal lips drying the same color as the rachillae, joined to form a
raised cupule, the margins not overlapping; distal lips well-developed; staminate and pistillate petals not
anthers short and curled over at anthesis; non-fertilized pistillate flowers no data; staminodial tubes crenulate
or shallowly lobed at the apex, persistence no data; fruits 7.5 mm long, 5.8 mm in diameter, the bases without
a prominent stipe, the apices not conical, the surfaces not splitting at maturity, without fibers emerging,
bumpy from the numerous, subepidermal, tangential, short fibers present, these coming to a point at fruit
apices; locular epidermis with operculum, smooth, without pores.
Distribution and habitat:At 1031N and 6620W in the Coastal Cordillera in Venezuela (Miranda)
at 800 m elevation in lowland rainforest (Fig. 29).
FIGURE 29. Distribution maps of Geonoma oldemanii, G. oligoclona, G. operculata, and G. orbignyana subsp.
orbignyana.
109
Taxonomic notes:This species is based on a single specimen. This is very similar to specimens of G.
braunii, and in fact it was identified as such by Stauffer (1997, as G. spinescens var. braunii). However, the
fruits of the specimen (in a separate packet, see Appendix IV, Plate 52) have an operculum, and because of
this the specimen is here recognized as a separate species. It is the only species outside of the Geonoma
congesta clade to have fruits with an operculum. Apart from the operculum, the specimen bears no
resemblance to species in the G. congesta clade but appears similar to specimens of G. braunii.
Subspecific variation: No trait varies within this species and only one specimen is known.
45. Geonoma orbignyana Martius (1843: 22). Type: BOLIVIA. Cochabamba: Yuracares, no date, A.
dOrbigny 44 (holotype P n.v.).
cane-like or cane-like; internodes 1.0(0.23.8) cm long, yellowish and smooth, or, if short and congested, not
scaly. Leaves 10(420) per stem, undivided or irregularly pinnate, sometimes regularly pinnate and the pinnae
with 1 main vein only, not plicate or plicate, bases of blades running diagonally into the rachis; sheaths
long 3.5(1.28.2) mm in diameter; veins raised and rectangular in cross-section adaxially; pinnae 5(126) per
side of rachis; basal pinna 30.1(13.059.5) cm, long, 3.0(0.115.5) cm wide, forming an angle of 44(795)
with the rachis; apical pinna 20.6(7.747.5) cm long, 7.1(0.321.3) cm wide, forming an angle of 25(643)
with the rachis. Inflorescences unbranched or branched 12 orders; prophylls and peduncular bracts not ribbed
with elongate, unbranched fibers, flattened (if tubular, narrow, and elongate then not ribbed), deciduous or
39.0) cm above the prophyll; peduncles 30.9(6.088.5) cm long, 3.5(1.311.1) mm in diameter; rachillae 5(1
28), 15.3(5.031.0) cm long, 3.5(1.86.6) mm in diameter, the surfaces without spiky, fibrous projections or
between the flower pits; flower pits usually spirally arranged, sometimes decussately or tricussately, then the
lips apiculate and lobed before anthesis, tearing in the center after anthesis, not recurved after anthesis, not
bases with a prominent, asymmetric stipe, the apices not conical, the surfaces not splitting at maturity, without
fibers emerging, bumpy from the numerous, subepidermal, tangential, short fibers present, these coming to a
point at fruit apices; locular epidermis without operculum, smooth, without pores.
Taxonomic notes:Geonoma orbignyana is a member of a group of high elevation, Andean species, the
G. undata clade, which also includes G. lehmannii, G. talamancana, G. trigona, and G. undata. These species
have been treated differently by both Wessels Boer (1968) and Henderson et al. (1995). They are closely
related and three of themG. lehmannii, G. orbignyana, and G. undata are difficult to distinguish from one
another, and extremely complex internally. Geonoma orbignyana differs from G. lehmannii and G.
talamancana in its prophylls and peduncular bracts which are flattened and not ribbed with elongate,
unbranched fibers; from G. talamancana in its well-developed peduncular bract; from G. trigona in its welldeveloped distal lips; and from G. undata in its prophyll surfaces which are not ridged and without unequally
wide ridges.
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HENDERSON
Geonoma jussieuana is treated here as a synonym of G. orbignyana subsp. orbignyana (contra both
Wessels Boer, 1968 and Henderson et al., 1995). The type specimen, with its unbranched inflorescence,
comes from a Bolivian population of plants with both unbranched and branched inflorescences (sometimes on
the same specimen). In bract structure specimens of this population resemble others of G. orbignyana.
Geonoma lehmannii subsp. lehmannii, superficially similar to this population in its unbranched
inflorescences, does not reach Bolivia and has its southernmost population in central Peru.
Subspecific variation:Five traits vary within this species (stem branching, stem type, leaf division, leaf
plication, inflorescence branching). Excluding stem branching and leaf division and one trait for which there
are few data (stem type), the state distributions of the remaining two traits (leaf plication, inflorescence
branching) do not divide the specimens into consistent subgroups which are geographically separated. Both
leaf plication and inflorescence branching appear inconsistent. Leaf plication is difficult to score in this
species, and branched and unbranched inflorescences can be found on the same specimen. There is, however,
geographic disjunction and there is a gap in eastern Panama between Central American and South American
specimens.
Central American specimens differ from South American ones in 12 variables (rachis width, number of
pinnae, basal pinna length, basal pinna width, basal pinna angle, apical pinna length, apical pinna width,
apical pinna angle, peduncular bract length, interbract distance, peduncle length, number of rachillae)(t-test, P
<0.05). Based on this and geographic separation, the two subgroups are recognized as subspecies (subspp.
hoffmanniana, orbignyana).
Key to the subspecies of G. orbignyana
1
-
Peduncles 25.6(6.059.5) cm long; South America (Venezuela, Colombia, Ecuador, Peru, and Bolivia) ....................
.......................................................................................................................................................... subsp. orbignyana
Peduncles 32.4(20.956.0) cm long; Central America (Nicaragua, Costa Rica, Panama)............subsp. hoffmanniana
45a. Geonoma orbignyana subsp. orbignyana

Geonoma jussieuana Martius (1843: 24). Type: BOLIVIA. Cochabamba: Serra de Cochabamba, Cumbrecilla, no date,
A. dOrbigny 45 (holotype P!), synon. nov.
Geonoma lindeniana Wendland (1856: 337). Type: VENEZUELA. Tchira: Capacho, no date, H. Funck & L. Schlim s.
n. (holotype BR n.v., isotype K!).
Geonoma pumila Linden & Wendland (1856: 338). Type: COLOMBIA. Magdalena: Santa Marta, no date, H. Funck &
L. Schlim s. n. (holotype BR n.v., isotype K!).
Geonoma linearifolia Karsten (1856: 411). Type: COLOMBIA. Cundinamarca: Servit, Bogot, no date, H. Karsten s. n.
(holotype LE!).
Geonoma ramosa Engel (1865: 684). Type: VENEZUELA. Tchira: San Cristbal, no date, F. Engel s. n. (holotype B,
destroyed, isotype LE!).
Geonoma margyraffia Engel (1865: 685). Type: VENEZUELA. Trujillo: Villa Bocon, no date, F. Engel s. n. (holotype
B, destroyed, isotype LE!).
Geonoma goniocarpa Burret (1930a: 185). Type: COLOMBIA. Antioquia: Murr, ca. 1850 m, no date, W. Kalbreyer s. n.
(holotype B, destroyed). Neotype (selected by Bernal et al. 1989): COLOMBIA. Antioquia: Mun. Frontino,
Corregimiento de Murr, carretera Nutibara-La Blanquita, camino a Charrascal, ca. 2100 m, 7 January 1982, G.
Galeano & R. Bernal 485 (neotype COL!, isoneotype HUA n.v.).
Geonoma microclada Burret (1930a: 190). Type: COLOMBIA. Cauca [Caldas]: Montaa del Oro, Supia, 20002300 m,
no date, F. Lehmann 7322 (holotype B, destroyed, isotype K!).
Geonoma lepidota Burret (1930a: 191). Type: COLOMBIA. Antioquia: Ro Dolores, Santa Rosa, 16002000 m,
December 1891, F. Lehmann 7321 (holotype B, destroyed, isotype K!).
Geonoma paleacea Burret (1930a: 199). Type: COLOMBIA. Antioquia: Medelln, Nare, Ro Guatap, 2500 m, 25
February 1880, W. Kalbreyer 1478 (holotype B, destroyed). Neotype (selected by Bernal et al. 1989): COLOMBIA.
Antioquia: 5 km al este de Guatap, 2500 m, 17 February 1987, R. Bernal & L. Tobn 1377 (neotype COL!,
isoneotype HUA n.v.).
Geonoma pachydicrana Burret (1930a: 206). Type: BOLIVIA. Cochabamba: vicinity of Cochabamba, 1891, M. Bang
877 (holotype B, destroyed, isotypes BM!, F!, NY!, MO!, US!).
111
Geonoma aulacophylla Burret (1930a: 216). Type: COLOMBIA. Antioquia: Alto San Jos, 31003160 m, 30 April
1880, W. Kalbreyer 1607 (holotype B, destroyed). Neotype (selected by Bernal et al. 1989): COLOMBIA.
Antioquia: Cerro San Jos, ca. 10 km al noreste de Santa Rosa de Osos, 26002900 m, 78 January 1985, R. Bernal
& G. Galeano 845 (neotype COL!).
Geonoma plicata Burret (1930a: 217). Type: COLOMBIA. Antioquia: Alto San Jos, 2950 m, 30 April 1880, W.
Kalbreyer 1607b (holotype B, destroyed). Neotype (selected by Bernal et al. 1989): COLOMBIA. Antioquia: Cerro
San Jos, ca. 10 km al noreste de Santa Rosa de Osos, 26002900 m, 78 January 1985, R. Bernal & G. Galeano 843
(neotype COL, isoneotypes AAU!, HUA, NY!).
Geonoma wilsonii Galeano & Bernal (2002: 282). Type: COLOMBIA. Caquet: Mun. Florencia, Florencia-Suaza road,
km 35, vereda Las Brisas, 144N, 7544W, 16001700 m, 8 August 2001, R. Bernal & W. Malagn 2900
(holotype COL!, isotypes AAU n.v., COAH n.v., HUA n.v., MO n.v., NY!), synon. nov.
Inflorescences peduncular bracts 17.4(3.035.5) cm long; peduncles 25.8(6.059.5) cm long.

Distribution and habitat:From 1106N1747S and 64147945W in the Andes of South
America in Venezuela, Colombia, Ecuador, Peru, and Bolivia at 1966(7752850) m elevation in montane
This subspecies is widely distributed and extremely variable. There is geographical variation, although
much less than in the sympatric Geonoma undata. Regression shows there are significant associations
between elevation and six leaf and three inflorescence variables. Squared multiple R for the regression of leaf
number on elevation is 0.15, number of pinnae 0.03, basal pinna width 0.04, basal pinna angle 0.08, apical
pinna width 0.08, apical pinna angle 0.10, prophyll length 0.08, interbract distance 0.15, and peduncle length
0.06. Plants at higher elevations have fewer leaves with fewer pinnae, wider basal and apical pinnae with
narrower angles, and longer prophylls, interbract distances, and peduncles.
Specimens from the Venezuelan Andes (lindeniana morphotype) have leaves with 6(314) pinnae per
side of the rachis and inflorescences with 7(414) rachillae. The types of G. lindeniana, G. margyraffia, and G.
ramosa are from this region. Specimens occur in three areas. Those from Yaracuy have slender inflorescences
branched to one order, few rachillae, and fruits which are obviously apiculate. Specimens from Trujillo are
similar, except that one (Dorr 7315) has inflorescences branched to two orders, and the fruits are less
obviously apiculate. Specimens from Tchira have stouter inflorescences with shorter peduncles, shorter
interbract distances, and more, wider rachillae with a distinctive, thinner, sterile basal part. Several
specimens from Cesar and Norte de Santander in Colombia are similar.
Specimens from Colombia in the Sierra Nevada de Santa Marta (pumila morphotype) have smaller leaves
with 2(23) pinnae per side of the rachis and slender inflorescences with 5(39) rachillae. The type of G.
pumila is from this area.
Specimens from the central part of the Eastern Cordillera in Colombia (Boyac, Cundinamarca, Meta,
Norte de Santander, Santander)(linearifolia morphotype) have mostly regularly pinnate leaves with 16(326)
pinnae per side of the rachis and branched, rarely unbranched inflorescences with 5(112) rachillae. The type
of G. linearifolia is from this area. One specimen from Cudinamarca (Grant 9177) has larger leaves and a
large stout inflorescence, much larger than other specimens. Several specimens (Betancur 6220, Bernal 1342,
3512, 3513, Betancur 5714, Snchez Vega 6696) from the Eastern Cordillera are larger than others and appear
intermediate between this morphotype and the weberbaueri morphotype of G. undata subsp. undata. These
may be hybrids and are excluded from the above descriptions and analyses.
A few other specimens (Bernal 2900the type of G. wilsonii, Bernal 2901, Malagon 26) from Caquet in
the southern part of the Eastern Cordillera (wilsonii morphotype) are much reduced in size.
Specimens from the Central and Western Cordilleras in Colombia (plicata morphotype) have leaves with
4(114) pinnae per side of the rachis and stout, often elongate inflorescences with 9(324) rachillae.
Specimens from the Cerro San Jos and adjacent areas in Antioquia have plicate leaves. The types of
Geonoma plicata, G. paleacea, G. goniocarpa, G. aulacophylla, G. lepidota, and G. microclada are from this
area. Specimens from northern Ecuador are similar.
On the eastern Andean slopes of Ecuador on the Cordillera de Huacamayos (baeza morphotype)
specimens have leaves with 4(36) pinnae per side of the rachis and slender inflorescences with 5(19)
rachillae with the peduncular bract inserted well above the prophyll and exerted from it.
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HENDERSON
Specimens from southern Ecuador and northern Peru, and continuing south to Bolivia (southern
morphotype), are very variable. In northern Peru, there are three distinct groups of specimens from San Martn
occurring in the same area. One group (Gentry 45513, Smith 4590) has regularly pinnate leaves and
inflorescences branched to two orders; the second (Smith 4842) has regularly pinnate leaves and two, thick
rachillae; and the third (Gentry 45312, 45403, 45512, 45538) with undivided leaves and few, thin rachillae.
There are two very distinct groups from the Cerro del Sira in Hunuco. One has finely pinnate leaves and
small inflorescences and occurs at lower elevations (Dudley 13064, Rainer 133288, 1330188, 2214988,
2314988, Wolfe 12335); the second (Rainer 2513988) has irregularly pinnate leaves and larger inflorescences,
and occurs at higher elevations.
Specimens from southern part of Peru (Cuzco, Pasco, Puno) have wider rachillae. In Bolivia, some
specimens have wide apical pinna and short, thick, densely tomentose rachillae, e.g., the type of G.
pachydicrana. Other specimens have narrow and widely spaced pinnae, unbranched or branched
inflorescences (sometimes on the same specimen), the bracts cover the peduncle, and glabrous rachillae. The
types of G. orbignyana and G. jussieuana have this kind of inflorescence.
45b. Geonoma orbignyana subsp. hoffmanniana (Wendland ex Spruce) Henderson, comb. & stat. nov.
Basionym: Geonoma hoffmanniana Wendland ex Spruce (1871: 106). Type: COSTA RICA. Heredia: Volcn de Barba,
no date, H. Wendland s.n. (holotype K!).
Geonoma molinae Glassman (1964: 7). Type: NICARAGUA. Matagalpa: Santa Mara de Ostuma, between Matagalpa
and Jinotega, 13001500 m, 8 January 1963, L. Williams, A. Molina, & R. Williams 23507 (holotype F!).
Inflorescences peduncular bracts 20.6(10.727.5) cm long; peduncles 32.4(20.956.0) cm long.

Distribution and habitat:From 8521302N and 82338620W in Nicaragua, Costa Rica, and
Panama at 2008(14003000) m elevation in montane rainforest (Fig. 30).
This subspecies occurs in three separate areas; Nicaragua, the central part of Costa Rica, and eastern
Costa Rica/western Panama.
There are six specimens from Nicaragua and these are small in size. There are no differences in any
quantitative variable between these specimens and those of central Costa Rica, although they do occur at
lower mean elevations (1475 m versus 2030 m).
In central Costa Rica specimens occur on three separate Cordilleras; Pacific slope on Tilarn
(Monteverde), Atlantic slope on Central (Barva); and Pacific and Atlantic slope on Central. Specimens from
Tilarn (Monteverde) have unbranched inflorescences, as does one specimen from Central. Specimens from
Barva and the Pacific and Atlantic slopes of Central are small in size and similar to those from Nicaragua.
In eastern Costa Rica and western Panama, on the Talamanca, some specimens are also small (Davidse
26197, Fletes 1, Gamboa 708) but the others are the largest of any area, and occur at higher elevations. These
specimens occur sympatrically with large specimens of G. undata subsp. edulis. Hammel et al. (2003)
considered that larger specimens of subsp. hoffmanniana (as G. hoffmanniana) and sympatric subsp. edulis (as
G. edulis) were virtually indistinguishable.
There is geographical variation in this subspecies. Regression shows there are significant associations
between elevation and one plant, three leaf, and one inflorescence variable. Squared multiple R for the
regression of stem height on elevation is 0.32, rachis width 0.24, basal pinna length 0.43, apical pinna length
0.33, and peduncle width 0.17. Values of these variables increase with increasing elevation. Stems become
taller, rachis wider, basal and apical pinnae longer and peduncles wider with increasing elevation.
46. Geonoma paradoxa Burret (1934a: 1040). Type: COLOMBIA. Cauca: Coteje and Santa Mara on Ro
Timbiqu, 200600 m, 1898, F. Lehmann 8957 (holotype B, destroyed, isotypes K!, NY!).
Plants 0.8(0.71.0) m tall; stems 0.2(0.10.3) cm tall, 1.3(1.21.3) cm in diameter, solitary, not cane-like;
internodes 0.4(0.30.4) cm long, not scaly. Leaves 9(612) per stem, undivided or irregularly pinnate, not
113
raised and rectangular in cross-section adaxially; pinnae 1(13) per side of rachis; basal pinna 37.5 cm long,
11.0 cm wide, forming an angle of 28(1940) with the rachis; apical pinna 13.5(11.517.2) cm long, 11.0 cm
the surfaces not ridged, without unequally wide ridges; peduncular bracts 8.8(5.512.0) cm long, welldeveloped, inserted 1.5(0.72.2) cm above the prophyll; peduncles 58.8(27.5100.0) cm long, 2.0(1.33.7)
mm in diameter; rachillae 1, 11.3(7.816.0) cm long, 3.6(2.74.6) mm in diameter, the surfaces without spiky,
cupule, the proximal lip margins overlapping the distal lip margins; distal lips a scarcely raised rim; staminate
6; thecae diverging or not diverging at anthesis, inserted onto poorly to well-developed, non-split, jointed
connectives, connectives when well-developed alternately long and short; anthers short at anthesis, remaining
straight and parallel; non-fertilized pistillate flowers deciduous after anthesis; staminodial tubes lobed at the
apex, the lobes spreading at anthesis, acuminate, those of non-fertilized pistillate flowers not projecting and
prominent stipe, the apices not conical, the surfaces not splitting at maturity, without fibers emerging, not
bumpy, not apiculate; locular epidermis with operculum, smooth, without pores.
Distribution and habitat:From 115359N and 76587840W on the Pacific coast of Colombia
and Ecuador at 118(10400) m elevation in lowland rainforest (Fig. 30).
Taxonomic notes:Geonoma paradoxa is a member of the G. macrostachys clade, within which it is
related to a group of species which includes G. macrostachys, G. multisecta, G. poiteauana, and G.
schizocarpa. It differs from these in its distal lips which are scarcely raised rims. Despite having a relatively
narrow distribution it is confusingly variable, as explained below.
Subspecific variation:One trait (leaf division) varies within this species. All specimens but two have
undivided leaves. The specimens are clearly divisible into small and large size subgroups. Although there are
only six large size specimens and these have missing data, of the eight variables with enough data, large size
specimens differ from small size ones in seven variables (rachis length, rachis width, basal pinna angle, apical
pinna angle, peduncle width, rachilla length, rachilla width)(t-test, P <0.05). All large size specimens are from
low elevations (1060 m) near the sea in the Ro Naya-Ro Yurumangui region of Valle. Small size specimens
occur at higher elevations (25400 m) from near Buenaventura in Valle to northwestern Ecuador.
There is variation in connectives. Of the nine (of 16) specimens with staminate flowers, three small size
specimens from the northern most part of the range have well-developed connectives. Two, from the most
southern part of the range, have poorly developed connectives, and one, from the central part of the range has
aberrant flowers in which one theca of a pair is developed and the other not. The large size specimens also
appear to have aberrant flowers. Without more specimens it is not possible to understand the variation within
this species, and no subspecies are recognized.
47. Geonoma pauciflora Martius (1823: 12). Type: BRAZIL. State unknown: in sylvis ad fluv. Amazonum,
no date, C. Martius s.n. (holotype M!).
Geonoma conduruensis Lorenzi (2010: 226). Type: BRAZIL. Bahia: Itacar, Fazenda So Miguel, km 43 rodovia IlhusItacar, 1423S, 3903W, 70 m, 24 June 2008, H. Lorenzi, L. Noblick, & A. Guimares 6498 (holotype HPL n.v.,
isotypes CEPEC n.v., ESA n.v., NY n.v., RB n.v., SP n.v.), synon. nov.
114
HENDERSON
FIGURE 30. Distribution maps of Geonoma orbignyana subsp. hoffmanniana, G. paradoxa, G. pauciflora, and G.
peruviana.
4.5) mm in diameter; veins not raised or slightly raised and triangular in cross-section adaxially; pinnae 2(1
6) per side of rachis; basal pinna 17.5(10.526.0) cm long, 3.0(1.46.0) cm wide, forming an angle of 36(14
66) with the rachis; apical pinna 11.8(7.018.5) cm long, 8.3(5.018.0) cm wide, forming an angle of 30(17
45) with the rachis. Inflorescences unbranched or branched 1 order; prophylls and peduncular bracts not
ribbed with elongate, unbranched fibers, flattened (if tubular, narrow, and elongate then not ribbed), deciduous
or persistent; prophylls 13.5(7.019.0) cm long, not short and asymmetrically apiculate, the surfaces not
narrowed between the flower pits; flower pits usually spirally arranged, sometimes decussately or tricussately,
then the groups not closely spaced nor consistently arranged throughout the rachillae, glabrous internally;
and distal lips drying the same color as the rachillae, not joined to form a raised cupule, the proximal lip
115
margins overlapping the distal lip margins; distal lips well-developed; staminate and pistillate petals not
anthers short and curled over at anthesis; non-fertilized pistillate flowers deciduous after anthesis; staminodial
persistent after anthesis; fruits 9.6(7.312.1) mm long, 7.4(5.68.5) mm in diameter, the bases with a
prominent, asymmetric stipe, the apices conical with rounded apices, the surfaces not splitting at maturity,
without fibers emerging, not bumpy, not apiculate; locular epidermis without operculum, sculpted, usually
also with a raised, meridional ridge; locular epidermis without pores.
Distribution and habitat:From 8241704S and 35044123W in the Atlantic Coastal Forest of
Brazil from Pernambuco to Bahia, with an inland outlier in Bahia, at 291(20770) m elevation in lowland
Taxonomic notes:The type locality is unknown. The label of the type specimen (in sylvis ad fluv.
Amazonum) differs from the description (in....Provinciarum Piauhiensis et Maraguaniensis). Geonoma
pauciflora is a member of a group of species from the Atlantic Coastal Forest and adjacent Cerrado of Brazil
(the G. schottiana clade, also including G. elegans, G. pohliana, and G. schottiana). Although the group is
well-supported, all constituent species are extremely variable internally. Geonoma pauciflora differs from G.
elegans in its flattened prophylls and peduncular bracts which are not ribbed with elongate, unbranched fibers;
from G. pohliana in its prophyll surfaces not ridged and without unequally wide ridges; and from G.
schottiana in its undivided or irregularly pinnate leaves.
inflorescence branching). There is little geographic discontinuity, and the specimens are widely distributed in
the northern part of the Atlantic Coastal Forest of Brazil. Leaving aside stem branching and leaf division, the
remaining traitinflorescence branchingdoes not divide the specimens into consistent groups. However,
Geonoma pauciflora is variable in leaf division and inflorescence branching, and three morphotypes may be
recognized.
The first morphotype has pinnate (rarely undivided) leaves and branched (rarely unbranched)
inflorescences (pinnate-branched morphotype). This morphotype occurs in two separate areas; Pernambuco
and Alagoas, and central and southern Bahia. The gap between these two areas is likely to be based on
unsuitable habitat (W. Thomas, pers. comm.). However, there are too few specimens from Pernambuco and
Alagoas to test for differences between the areas. Specimens of this morphotype from the most southerly part
of the range in Bahia (e.g., Gentry 49920, Thomas 12020) tend to have smaller leaves and inflorescences, and
may be hybrids between Geonoma pauciflora and G. elegans (which see).
The second morphotype has undivided leaves and branched inflorescences, and occurs in central Bahia
with an outlier further inland (undivided-branched morphotype). The type of Geonoma conduruensis is of this
morphotype.
The third morphotype has undivided leaves and unbranched (rarely branched) inflorescences, and occurs
in two localities in central Bahia (undivided-unbranched morphotype).
48. Geonoma peruviana Henderson, sp. nov. (Appendix IV, Plate 53)
A speciebus affinibus internodis squamis brenneis crebre tectis differt.
Type: PERU. Hunuco: Prov. Leoncio Prado, Distrito Hermilio Valdizan, cerca La Divisoria, 15001600 m, 25 June
1976, J. Schunke 9416 (holotype MO!).
Plants 1.0(0.91.2) m tall; stems height no data, 1.0(0.81.2) cm in diameter, branching no data, not cane-like
or cane-like; internodes 0.4(0.30.5) cm long, covered with dense, brown scales. Leaves irregularly pinnate,
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HENDERSON
not plicate, bases of blades running diagonally into the rachis; sheaths no data; petioles drying green or
yellowish; rachis no data; veins not raised or slightly raised and triangular in cross-section adaxially; pinnae 3
per side of rachis; basal pinna length, width, and angle no data; apical pinna 12.5 cm long, 15.0 cm wide,
forming an angle of 40 with the rachis. Inflorescences branched 2 orders; prophylls and peduncular bracts not
ribbed with elongate, unbranched fibers, flattened (if tubular, narrow, and elongate then not ribbed), deciduous
or persistent; prophylls 8.5 cm long, not short and asymmetrically apiculate, the surfaces not ridged, without
unequally wide ridges; peduncular bracts length no data, well-developed, no data on insertion; peduncles 17.0
cm long, 2.3 mm in diameter; rachillae 10(910), 7.4(6.58.2) cm long, 1.2(1.11.3) mm in diameter, the
anthesis, sometimes splitting post-anthesis; proximal and distal lips drying the same color as the rachillae, not
joined to form a raised cupule, the proximal lip margins overlapping the distal lip margins; distal lips welldeveloped; staminate and pistillate petals not emergent, not valvate throughout; staminate flowers deciduous
flowers deciduous after anthesis; staminodial tubes crenulate or shallowly lobed at the apex, those of nonfertilized pistillate flowers not projecting and persistent after anthesis; fruits 5.4 mm long, 5.2 mm in diameter,
Distribution and habitat:At 900S and 7555W in Peru (Hunuco) on eastern Andean slopes at
1550 m elevation in montane rainforest (Fig. 30).
Taxonomic notes:Only two specimens of this species are known. It differs from related species in its
internodes covered with dense, brown scales.
49. Geonoma pinnatifrons Willdenow (1805: 593). Type: VENEZUELA. "Caracas, Buenavista", no date, F.
Bredemeyer 20 (holotype B n.v., holotype image!, isotype M!).
long; petioles 54.9(10.0125.0) cm long, drying green or yellowish; rachis 89.1(36.0163.0) cm long,
6.6(1.914.5) mm in diameter; veins raised and rectangular in cross-section adaxially; pinnae 9(239) per side
of rachis; basal pinna 42.3(21.565.0) cm long, 4.7(0.522.5) cm wide, forming an angle of 47(2070) with
the rachis; apical pinna 33.6(16.061.0) cm long, 17.3(3.544.5) cm wide, forming an angle of 29(2137)
unbranched fibers, flattened (if tubular, narrow, and elongate then not ribbed), deciduous or persistent;
prophylls 15.6(7.025.0) cm long, not short and asymmetrically apiculate, the surfaces ridged and densely
tomentose with widely to closely spaced ridges, unequally wide, often dividing from and rejoining other
ridges, the prophyll margins with irregular, spine-like projections, the prophylls usually splitting irregularly
between the ridges; peduncular bracts 16.8(10.027.5) cm long, well-developed, inserted 2.6(0.67.4) cm
above the prophyll; peduncles 24.2(10.042.0) cm long, 8.1(2.421.2) mm in diameter; rachillae 18(445),
14.2(7.028.3) cm long, 2.3(1.13.8) mm in diameter, the surfaces without spiky, fibrous projections or
ridges, drying brown, with faint to pronounced, short, transverse ridges, not filiform and not narrowed
between the flower pits; flower pits spirally arranged, densely hairy internally proximally and distally;
proximal lips without a central notch before anthesis, not recurved after anthesis, hood-shaped at anthesis,
117
sometimes splitting post-anthesis; proximal and distal lips drying the same color as the rachillae, not joined to
form a raised cupule, the proximal lip margins overlapping the distal lip margins; proximal lips hood-shaped;
distal lips absent; staminate and pistillate petals not emergent, not valvate throughout; staminate flowers
pistillate flowers persistent or deciduous after anthesis; staminodial tubes crenulate or shallowly lobed at the
smooth or locular epidermis sculpted and then usually also with a raised, meridional ridge, without pores.
Taxonomic notes:Geonoma pinnatifrons is a member of a group of related species characterized by its
lack of a distal lip of the flower pit and flower pits hairy internally. This group, the G. interrupta clade, also
includes G. euspatha, G. frontinensis, G. interrupta, and G. simplicifrons. These species have had a checkered
taxonomic history. Geonoma pinnatifrons was included under G. interrupta by Wessels Boer (1965), but later
reinstated (Wessels Boer, 1968). Both species are complicated internally. Geonoma pinnatifrons differs from
G. euspatha, G. frontinensis, and G. simplicifrons in its prophylls surfaces which are ridged and densely
tomentose with widely to closely spaced ridges, unequally wide, often dividing from and rejoining other
ridges; and from G. interrupta in its flower pits which are densely hairy internally proximally and distally (see
also Taxonomic notes under G. interrupta).
Subspecific variation:Three traits vary within this species (stem branching, stem type, locular
epidermis sculpting). There are few data for any of these traits. The species occurs widely in Central America
and northern South America. Six peripherally isolated areas (Lesser Antilles; Trinidad, Tobago, and the Paria
Peninsula of Venezuela; Sierra Nevada de Santa Marta in Colombia; Hispaniola; Pacific coast of Colombia;
Pacific coast of Mexico and Guatemala) contain subgroups that differ significantly from their nearest
neighbors in four to ten variables, and these are recognized as subspecies (subspp. martinicensis, vaga,
platybothros, oxycarpa, ramosissima, membranacea). The remaining specimens can be divided into three
subgroups based on geography and rachis and peduncle widthnorthern South America (Venezuela,
Colombia, and Ecuador, and just reaching Panama); eastern Panama; and the rest of Central America.
However, there is not complete geographic separation between these. ANOVA shows that for pairwise
comparison probabilities, 17 variables (plant height, stem height, leaf number, petiole length, rachis length,
rachis width, number of pinnae, basal pinna length, basal pinna width, apical pinna length, apical pinna width,
prophyll length, interbract distance, peduncle length, peduncle width, rachilla length, fruit diameter) differ
significantly (P <0.05) between one pair of subgroups, although no variables differs amongst all subgroups.
Based on this and geographic disjunction, these subgroups are recognized as subspecies (subspp. pinnatifrons,
binervia, mexicana).
Key to the subspecies of Geonoma pinnatifrons
1
2
3
4
-
Rachillae 1.6(1.12.3) mm in diameter; Pacific Coast and western slope of the Western Cordillera in Colombia .......
.........................................................................................................................................................subsp. ramosissima
Rachillae 2.3(1.13.8) mm in diameter; widespread .................................................................................................... 2
Peduncles 13.0(9.320.2) mm in diameter; eastern Panama and adjacent Colombia with outliers in western Panama,
Costa Rica, Nicaragua and Guatemala ...................................................................................................subsp. binervia
Peduncles 8.2(2.420.2) mm in diameter; widespread................................................................................................. 3
Peduncles 16.5(13.618.8) cm long; fruits 8.5(7.49.3) mm long, 6.3(5.27.0) mm in diameter; Pacific coast of
Mexico and Guatemala ................................................................................................................. subsp. membranacea
Peduncles 25.9(11.041.0) cm long; fruits 6.2(4.48.8) mm long, 4.8(3.66.3) mm in diameter; widespread........... 4
Pinnae 11(339) per side of rachis; Colombia, Venezuela, and Ecuador, and just reaching eastern Panama ................
..........................................................................................................................................................subsp. pinnatifrons
Pinnae 4(29) per side of rachis; Lesser Antilles, Central America, Colombia (Sierra Nevada de Santa Marta), Ven-
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HENDERSON
5
6
7
8
-
ezuela (Pennsula de Paria), Trinidad and Tobago, and Haiti ...................................................................................... 5

Rachillae 18.7(12.224.0) cm long; Lesser Antilles (Guadeloupe, Dominica, Martinique, and St. Lucia) ..................
....................................................................................................................................................... subsp. martinicensis
Rachillae 12.3(7.018.9) cm long; Central America, Colombia (Sierra Nevada de Santa Marta), Venezuela (Pennsula de Paria), Trinidad and Tobago, and Haiti ............................................................................................................... 6
Peduncular bracts inserted 1.5(0.72.5) cm above the prophylls; Venezuela (Pennsula de Paria), Trinidad and
Tobago.......................................................................................................................................................... subsp. vaga
Peduncular bracts inserted 3.8(1.87.4) cm above the prophylls; Central America, Central America, Colombia
(Sierra Nevada de Santa Marta), and Haiti ................................................................................................................... 7
Peduncles 4.0(2.85.6) mm in diameter; Colombia (Sierra Nevada de Santa Marta).....................subsp. platybothros
Peduncles 7.0(3.810.7) mm in diameter; Central America (Mexico, Guatemala, Belize, Honduras, Nicaragua,
Costa Rica, western Panama) and Haiti ........................................................................................................................ 8
Peduncles 28.4(19.037.0) cm long; Central America (Mexico, Guatemala, Belize, Honduras, Nicaragua, Costa
Rica, and western Panama) ..................................................................................................................subsp. mexicana
Peduncles 22.8(19.026.5) cm long; Haiti ........................................................................................... subsp. oxycarpa
49a. Geonoma pinnatifrons subsp. pinnatifrons

Geonoma pulchra Engel (1865: 686). Type: COLOMBIA. Norte de Santander: Ocaa, 20003000 m, August 1859, F.
Engel s. n. (holotype B, destroyed, isotype LE!).
Leaf pinnae 11(339) per side of rachis. Inflorescences peduncular bracts inserted 2.8(0.66.5) cm above the
prophyll; peduncles 25.2(11.041.0) cm long, 7.5(3.612.9) mm in diameter; rachillae 15.2(8.625.7) cm
long, 2.2(1.13.4) mm in diameter fruits 5.0(3.66.3) mm in diameter.
Distribution and habitat:From 243S1110N and 63257812W in Venezuela, Colombia, and
Ecuador, and just reaching eastern Panama, at 898(301900) m elevation in lowland to montane rainforest
(Fig. 31).
There is geographical variation in this subspecies. Regression shows there are significant (P <0.05)
associations between elevation and three leaf and three inflorescence variables. Squared multiple R for the
regression of petioles on elevation is 0.57, basal pinna length 0.27, basal pinna width 0.28, prophyll length
0.39, rachilla length 0.27, and fruit length 0.23. Petioles become shorter, basal pinna shorter and narrower,
prophyll and rachillae shorter, and fruits longer with increasing elevation.
Clustered-stemmed plants occur commonly in this subspecies in Venezuela, and these occur at a higher
mean elevation (1170 m) than those with solitary stems (677 m).
49b. Geonoma pinnatifrons subsp. binervia (Orsted) Henderson, comb. & stat. nov.
Basionym: Geonoma binervia rsted (1859: 33). Type: NICARAGUA. Ro San Juan: Ro San Juan, 18451848, A.
rsted 6564 (holotype C!).
long, 2.3(1.73.4) mm in diameter; fruits 4.3(3.75.1) mm in diameter.
Distribution and habitat:From 7191604N and 77059015W in eastern Panama and adjacent
Colombia with outliers in western Panama, Costa Rica, Nicaragua, and Guatemala at 282(4725) m elevation
in lowland rainforest (Fig. 31).
There is one doubtful specimen (Cook 5808), missing leaves, from the Nicoya Peninsula of Costa Rica
that is included here.
49c. Geonoma pinnatifrons subsp. martinicensis (Martius) Henderson, comb. & stat. nov.
Basionym: Geonoma martinicensis Martius (1843: 28). Type: MARTINIQUE. Palma humilis, cocifera, latifolia, major,
Plum., t. LIX, LX, LXI (holotype P, n.v.).
Geonoma dominicana Bailey (1939: 232). Type: Dominica. Morne Cola Anglais, 610732 m, 1023 August 1938, W.
Hodge 307 (holotype, BH! isotypes, GH! NY!).
119
Distribution and habitat:From 12271613N and 60566146W in the Lesser Antilles
(Guadeloupe, Dominica, Martinique, and St. Lucia) at 674(2001000) m elevation in lowland to montane
rainforest (Fig. 31). Read (1979) also included St. Vincent in the distribution of this taxon, but no specimens
from there have been seen.
49d. Geonoma pinnatifrons subsp. membranacea (Wendland ex Spruce) Henderson, comb. & stat. nov.
Basionym: Geonoma membranacea Wendland ex Spruce (1871: 106). Type: GUATEMALA. Escuintla: Volcan de
Fuego, between San Pedro and Hacienda de Sapota, January 1857, H. Wendland 7 & 8 (holotype K!).
Distribution and habitat:From 14251519N and 90039237W on slopes of volcanoes on the
Pacific coast of Mexico and Guatemala at 1235(7001650) m elevation in lowland to montane rainforest (Fig.
31).
FIGURE 31. Distribution maps of Geonoma pinnatifrons subsp. pinnatifrons, G. pinnatifrons subsp. binervia, G.
pinnatifrons subsp. martinicensis, and G. pinnatifrons subsp. membranacea.
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HENDERSON
49e. Geonoma pinnatifrons subsp. mexicana (Liebman ex Martius) Henderson, comb. & stat. nov.
Basionym: Geonoma mexicana Liebman ex Martius (1853: 316). Type: MEXICO. Oaxaca: Lobani, Chinantla, 900 m,
no date, F. Liebman 10804 (holotype C!, isotypes MO!, P!, US!).
Distribution and habitat:From 7431801N and 79559640W in Mexico, Guatemala, Belize,
Honduras, Nicaragua, Costa Rica, and western Panama (Bocas del Toro, Cocl, Herrera) at 553(501100) m
elevation in lowland or montane rainforest (Fig. 32).
FIGURE 32. Distribution maps of Geonoma pinnatifrons subsp. mexicana, G. pinnatifrons subsp. oxycarpa, G.
pinnatifrons subsp. platybothros, and G. pinnatifrons subsp. ramosissima.
Four specimens (Cook 35, 728, Doyle 10, Hernndez 1171) from scattered localities in Mexico,
Guatemala, and Costa Rica have many more pinnae than usual and the leaf appears regularly pinnate. One
possible explanation for this is that these specimens represent a mixture of inflorescences from G. pinnatifrons
subsp. mexicana and leaves from G. interrupta subsp. magnifica. The latter has many more pinnae and occurs
sympatrically with G. pinnatifrons subsp. mexicana. Leaf measurements from these four specimens are
omitted.
121
49f. Geonoma pinnatifrons subsp. oxycarpa (Martius) Henderson, comb. & stat. nov.
Basionym: Geonoma oxycarpa Martius (1843: 30). Type: HAITI. Palma humilis, cocifera, latifolia, minor, Plum., t.
LVI, LVII, LVIII (holotype P, n.v.).
Leaf pinnae 3(24) per side of rachis. Inflorescences peduncular bracts inserted no data; peduncles 22.8(19.0
26.5) cm long, 7.1(6.57.6) mm in diameter; rachillae 10.0(9.011.7) cm long, 2.7(2.52.9) mm in diameter;
fruits no data.
Distribution and habitat:From 18191945N and 72157352W in northern and southwestern
Haiti at 750 m in lowland rainforest (Fig. 32).
49g. Geonoma pinnatifrons subsp. platybothros (Burret) Henderson, comb. & stat. nov.
Basionym: Geonoma platybothros Burret (1931a: 200). Type: COLOMBIA. Magdalena: Santa Marta, 24 February 1899,
H. Smith 2340 (holotype B, destroyed, isotypes F!, K!, MO!, NY!, P!, US!).
Leaf pinnae 5(45) per side of rachis. Inflorescences peduncular bracts inserted 5.5 cm above the prophyll;
peduncles 31.5(21.042.0) cm long, 4.0(2.85.6) mm in diameter; rachillae 12.4(10.514.0) cm long, 2.2(1.7
2.7) mm in diameter; fruits 5.7(5.35.9) mm in diameter.
Distribution and habitat:From 10351110N and 73237403W on the Sierra Nevada de Santa
Marta in Colombia at 1371(3701909) m elevation in lowland or montane rainforest (Fig. 32).
49h. Geonoma pinnatifrons subsp. ramosissima (Burret) Henderson, comb. & stat. nov.
Basionym: Geonoma ramosissima Burret (1930a: 249). Type: COLOMBIA. Antioquia: Cieneguetas, 27 July 1880, W.
Kalbreyer 1892 (holotype B, destroyed). Neotype (selected by Bernal et al. 1989): COLOMBIA. Antioquia: Mun.
Frontino, Corregimiento de Murr, road from Nutibara to La Blanquita, Ro Cuevas, 950 m, 23 March 1982, R.
Bernal & G. Galeano 306 (neotype COL!, isoneotype NY!).
Distribution:From 355700N and 75547737W on the Pacific Coast and western slopes of the
Western Cordillera in Colombia, at 427(01150) m elevation in lowland to montane rainforest (Fig. 32).
There is geographic discontinuity but too few specimens to test for differences, and too few to test for
geographical variation. Three specimens (Bernal 306, Betancur 2818, Forero 7370) from higher elevations on
the western slopes of the Western Cordillera appear intermediate between this subspecies and G. interrupta
subsp. magnifica, and may represent hybrids. They have the flower pits densely hairy internally distally only,
as in G. interrupta.
49i. Geonoma pinnatifrons subsp. vaga (Grisebach & Wendland in Grisebach) Henderson, comb. & stat.
nov.
Basionym: Geonoma vaga Grisebach & Wendland in Grisebach (1864: 517). Geonoma saga Spruce (1871: 109), orth.
var. Geonoma pinnatifrons Willdenow var. vaga (Grisebach. & Wendland) Burret (1930a: 246). Lectotype (here
designated): TRINIDAD & TOBAGO. Trinidad: Mount Tamana, 28 April 1841, W. Purdie 23 (lectotype K!).
Leaf pinnae per side of rachis no data. Inflorescences peduncular bracts inserted 1.5(0.72.5) cm above the
Distribution and habitat:From 10381118N and 60346310W on the Pennsula de Paria,
Venezuela, the Northern Range, Trinidad, and Tobago at 740(400900) m elevation in lowland rainforest (Fig.
33).
Although there is geographic discontinuity there are too few specimens and too many missing data to test
for differences amongst areas.
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HENDERSON
FIGURE 33. Distribution maps of Geonoma pinnatifrons subsp. vaga, G. poeppigiana, G. pohliana subsp.
pohliana, and G. pohliana subsp. fiscellaria.
50. Geonoma poeppigiana Martius (1843: 35). Type: PERU. Loreto: Yurimaguas, February 1891, E. Poeppig
2295 (holotype M!).
cane-like; internodes 0.5(0.30.8) cm long, yellowish and smooth, or, if short and congested, not scaly. Leaves
11(416) per stem, undivided or irregularly pinnate, not plicate, bases of blades running diagonally into the
rachis; sheaths 16.9(10.025.0) cm long; petioles 49.8(30.0100.0) cm long, drying green or yellowish; rachis
adaxially or not raised or slightly raised and triangular in cross-section adaxially; pinnae 4(111) per side of
rachis; basal pinna 36.8(13.560.5) cm long, 4.9(0.515.3) cm wide, forming an angle of 40(1290) with the
rachis; apical pinna 30.0(10.345.0) cm long, 15.1(3.324.5) cm long, forming an angle of 28(2045) with
the rachis. Inflorescences unbranched or branched 1 order; prophylls and peduncular bracts not ribbed with
elongate, unbranched fibers, flattened (if tubular, narrow, and elongate then not ribbed), deciduous or
4.7) cm above the prophyll; peduncles 56.1(28.089.0) cm long, 5.6(1.911.1) mm in diameter; rachillae 4(1
10), 25.8(9.740.0) cm long, 4.2(1.97.1) mm in diameter, the surfaces without spiky, fibrous projections or
123
between the flower pits; flower pits spirally arranged, glabrous internally; proximal lips without a central
notch before anthesis, not recurved after anthesis, hood-shaped at anthesis, sometimes splitting post-anthesis;
lip margins overlapping the distal lip margins; distal lips a scarcely raised rim; staminate and pistillate petals
not emergent, not valvate throughout; staminate flowers deciduous after anthesis; stamens 6; thecae diverging
at anthesis, inserted onto bifid and well-developed, non-jointed connectives; anthers short and curled over at
lobed at the apex, those of non-fertilized pistillate flowers not projecting and persistent after anthesis; fruits
7.2(6.48.4) mm long, 5.6(4.66.6) mm in diameter, the bases without a prominent stipe, the apices not
without operculum, smooth or locular epidermis sculpted and then usually also with a raised, meridional
ridge; locular epidermis without pores.
Distribution and habitat:From 0401150S and 70097830W in the sub-Andean and western
Amazon regions of Colombia, Ecuador, Peru, and Brazil, with an outlier in southern Peru, at 288(110980) m
elevation in lowland tropical rainforest (Fig. 33).
Taxonomic notes:Wessels Boer (1968) considered that Geonoma poeppigiana had the inflorescences
covered with a distinctive brownish-gray tomentum. Some specimens have this but most do not, and it
appears to have little taxonomic significance. Geonoma poeppigiana appears most similar to G. brongniartii,
G. longipedunculata, and G. sanmartinensis. It differs from these in its thecae which diverge at anthesis and
are inserted onto bifid and well-developed, non-jointed connectives.
Subspecific variation:Five traits (stem type, leaf division, adaxial veins, inflorescence branching,
locular epidermis sculpting) vary within this species. There is geographic discontinuity and there are several
isolated populations. Leaving aside stem type and locular epidermis sculpting, for which there are few data,
and also leaf division, there is no correspondence between geography and variation in adaxial veins and
inflorescence branching. Specimens non-raised adaxial veins are from scattered localities and seem to be
associated with undivided leaves or leaves with fewer pinnae. However, this trait is difficult to score in this
species. Specimens with unbranched inflorescences occur in two areas but are intermixed with similar
specimens with branched inflorescences. Because of this inconsistency, no subspecies are recognized in this
species. However, there is much local variation.
Specimens from the Colombian and Brazilian Amazon and from the region around Iquitos in Peru
(Maynas Province) are similar. They have mostly undivided leaves or 25 pinnae per side of the rachis with
narrow basal angles and the adaxial veins are not or scarcely raised, with some exceptions (e.g., McDaniel
27585). Specimens from nearby Requena Province have more pinnae (28) and more pronounced adaxial
veins.
There is a gap in the distribution of G. poeppigiana between the region around Iquitos in Peru (Maynas
and Requena Provinces) and Ecuador and sub-Andean Peru. Despite this gap, there appears to be
geographical variation in G. poeppigiana. Regression shows there are significant associations between
longitude and eight leaf and three inflorescence variables. Squared multiple R for the regression of leaf
number on longitude is 0.50, number of pinnae 0.25, basal pinna length 0.75, basal pinna width 0.31, basal
pinna angle 0.33, apical pinna length 0.29, apical pinna width 0.49, apical pinna angle 0.52, peduncle width
0.26, rachilla length 0.27, and number of rachillae 0.21. In particular, there is a change in leaf shape, with
plants in the east having leaves with fewer pinnae (often undivided), longer and narrower basal and apical
pinnae with narrower angles. This kind of geographical variation also occurs in Geonoma brongniartii and G.
camana.
Specimens from the Ecuadorian Amazon and adjacent Peru also have more pinnae (510) per side of the
rachis and pronounced adaxial veins.
Specimens from Peru (Amazonas) are extremely variable. There are several specimens which resemble
those from the Ecuadorian Amazon, but others with unbranched inflorescences. For example, at the same
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HENDERSON
locality in Bagua Province, there are two specimens (Quipuscoa 340, Rodrguez 584) with leaves with 11,
narrow pinnae per side of the rachis and inflorescences with 45 rachillae, these 11.514.7 cm long and 3.0
3.6 mm wide; and two other specimens (Vsquez 19645, 23925) with three, broad pinnae per side of the rachis
and large, unbranched inflorescences with the rachilla 31.532.5 cm long and 6.56.6 mm wide.
Specimens from northern San Martn and adjacent Loreto in Peru are typical, although one (Moore 8530)
from Yurimaguas is considerably larger than the others, and is reported to have a stem to 4 m tall.
Specimens from southern San Martn, all from the same area near Tocache Nuevo are also very diverse.
There are several typical specimens, but one (Plowman 5937) has a undivided leaf with non-raised veins and
branched inflorescence, another (Schunke 10804) has a undivided leaf with raised veins and unbranched
inflorescence, two (Plowman 11461, Schunke 7612) have much smaller inflorescences and Schunke 7612 also
has a much smaller leaf, and others have the more common leaf type but unbranched inflorescences (Plowman
11677, Schunke 6615).
There is an outlying specimen in Madre de Dios. Three other specimens (Moreno 126, 305, Henderson
1636) from the border area between Peru and Bolivia, are not included in the above analyses or in the species
description. These have branched inflorescences and one has rachillae surfaces with faint to pronounced,
short, transverse ridges. These three specimens may represent hybrids between G. poeppigiana and another
species.
51. Geonoma pohliana Martius (1826: 142). Lectotype (here designated): BRAZIL. Rio de Janeiro: no
locality, no date, J. Pohl s.n. (lectotype M!).
Plants 2.9(1.06.0) m tall; stems 2.2(0.47.5) m tall, 1.3(0.62.9) cm in diameter, solitary or clustered, canelike; internodes 1.3(0.53.2) cm long, yellowish and smooth. Leaves 12(619) per stem, irregularly pinnate,
Inflorescences unbranched or branched 12 orders; prophylls and peduncular bracts not ribbed with elongate,
asymmetrically apiculate, the surfaces ridged and densely tomentose with widely to closely spaced ridges,
unequally wide, often dividing from and rejoining other ridges, the prophyll margins with irregular, spine-like
projections, the prophylls usually splitting irregularly between the ridges; peduncular bracts 11.8(4.022.0)
cm long, well-developed, inserted 2.3(0.212.5) cm above the prophyll; peduncles 15.9(5.034.5) cm long,
surfaces without spiky, fibrous projections or ridges, drying brown or yellow-brown, without short, transverse
ridges, filiform with extended narrowed sections between the flower pits, or not filiform and not narrowed
between the flower pits; flower pits usually spirally arranged, sometimes decussately or tricussately, then the
lips drying the same color as the rachillae or drying darker brown than the rachillae, not joined to form a
flowers deciduous after anthesis; staminodial tubes crenulate or shallowly lobed at the apex, those of nonfertilized pistillate flowers not projecting and persistent after anthesis; fruits 9.3(5.513.2) mm long, 7.2(4.7
9.9) mm in diameter, the bases with a prominent, asymmetric stipe, the apices conical with rounded apices, the
125
surfaces not splitting at maturity, without fibers emerging, not bumpy, not apiculate; locular epidermis without
operculum, sculpted, usually also with a raised, meridional ridge, without pores.
Taxonomic notes:Geonoma pohliana is a member of a group of species from the Atlantic Coastal
Forest and adjacent Cerrado (the G. schottiana clade, also including G. elegans, G. pauciflora, and G.
schottiana). Although the group is well-supported, all constituent species are extremely variable internally.
Geonoma pohliana differs from other species in the group by its prophyll surfaces which are ridged and
densely tomentose with widely to closely spaced, unequally wide ridges, these often dividing from and
rejoining other ridges. The variation within this species, both in qualitative traits, quantitative variables, and
geography allow separation into various subspecies, as described below.
Subspecific variation:Five traits vary within this species (stem branching, stem type, inflorescence
branching, rachillae narrowing, lip color). Leaving aside stem branching and stem type, for which there are
few data, the remaining three traits divide the specimens into three subgroups. However, lip color is difficult
to score in this species. There is little geographic discontinuity, and the specimens are widely distributed in the
Atlantic Coastal Forest and Cerrado regions of Brazil and just reach adjacent countries. These three subgroups
are analyzed separately.
The first subgroup, with branched inflorescences, narrowed rachillae, and concolorous lips, comprises a
single specimen from Rio de Janeiro. It is recognized as a subspecies (subsp. gastoniana).
The second subgroup, with branched inflorescences, non-narrowed rachillae, and bicolorous lips,
comprises a subgroup of smaller sized specimens and a subgroup of larger sized specimens. There are too few
smaller sized specimens to test for differences, but based on their small inflorescences with few (49)
rachillae, they are recognized as a subspecies (subsp. kuhlmannii). The large sized specimens occur in two
areasinland areas in Cerrado of Brazil and adjacent countries, and the Atlantic Coastal Forest of Brazil.
These two geographic subgroups differ from one another in 17 variables (petiole length, rachis length, number
of pinnae, basal pinna length, basal pinna width, basal pinna angle, apical pinna length, apical pinna width,
apical pinna angle, prophyll length, peduncular bract length, peduncle length, peduncle width, rachillae
length, rachillae width, fruit length, fruit diameter) (t-test, P <0.05). They also differ in their triad
arrangement. The Cerrado subgroup has mostly decussately arranged triads and the Atlantic Coastal Forest
subgroup has mostly spirally arranged triads. Based on this and geographic separation, they are recognized as
subspecies (subsp. pohliana, weddelliana).
The third subgroup comprises specimens with mostly branched inflorescences, non-narrowed rachillae,
and concolorous lips. One attribute serves to divide these specimensthe presence or absence of dense,
wooly hairs on the rachillae. There are too few specimens for quantitative analysis, but using the rachillae hair
attribute and geography, several distinct subgroups can be recognized. One subgroup from Esprito Santo,
Minas Gerais, and Rio de Janeiro has short, narrow, densely hairy rachillae and is recognized as a subspecies
(subsp. wittigiana). A second subgroup from Rio de Janeiro has long, thick, hairy rachillae, and is recognized
as a subspecies (subsp. fiscellaria). A third subgroup from Rio de Janeiro has more, narrower pinnae (1319
versus 37) and is recognized as a subspecies (subsp. trinervis). A fourth subgroup from Esprito Santo,
Minas Gerais, Rio de Janeiro, and So Paulo has few, thick, non-hairy rachillae and is recognized as a
subspecies (subsp. rodriguesii). A fifth subgroup from central Bahia has more numerous non-hairy rachillae
and is recognized as a subspecies (subsp. rubescens). A sixth subgroup from near Una in Bahia has few, hairy
rachillae and is recognized as a subspecies (subsp. unaensis). Finally, a seventh subgroup from southern Bahia
with more, hairy rachilllae is recognized as a subspecies (subsp. linharensis).
Key to the subspecies of G. pohliana
1
2
-
Rachillae filiform with extended narrowed sections between the flower pits; Rio de Janeiro.......... subsp. gastoniana
Rachillae not filiform and not or scarcely narrowed between the flower pits; widespread ......................................... 2
Proximal and distal lips drying darker brown than the rachillae .................................................................................. 3
Proximal and distal lips drying the same color as the rachillae ................................................................................... 5
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HENDERSON
3
4
Rachillae 5(49); Rio de Janeiro, Serra do Mar ................................................................................ subsp. kuhlmannii

Rachillae 12(332); widespread .................................................................................................................................. 4
Triads mostly decussately arranged; Cerrado region of central Brazil and just reaching adjacent Bolivia and Paraguay.................................................................................................................................................. subsp. weddelliana
Triads mostly spirally arranged; Atlantic Coastal Forest of Brazil from Cear, Pernambuco, and Alagoas to Rio
Grande do Sul ......................................................................................................................................supsp. pohliana
5 Pinnae 16(1323) per side of rachis; Rio de Janeiro, Serra do Mar ............................................................................ 6
Pinnae 4(27) per side of rachis; Bahia, Esprito Santo, Minas Gerais, Rio de Janeiro, and So Paulo...................... 7
6 Rachillae densely hairy....................................................................................................................... subsp. fiscellaria
Rachillae glabrous.................................................................................................................................. subsp. trinervis
7 Rachillae densely hairy................................................................................................................................................. 8
Rachillae glabrous or nearly so .................................................................................................................................. 10
8 Rachillae 1.8(1.22.6) mm in diameter ...............................................................................................subsp. wittigiana
Rachillae 4.8(3.76.8) mm in diameter ....................................................................................................................... 9
9 Rachillae 2(23); Bahia, near Una ........................................................................................................ subsp. unaensis
Rachillae 9(510); southern Bahia, Esprito Santo........................................................................... subsp. linharensis
10 Rachillae 4(26); Bahia ....................................................................................................................... subsp. rubescens
Rachillae 2(14); So Paulo, Rio de Janeiro, Minas Gerais, Esprito Santo, and southern Bahia .... subsp. rodriguesii
51a. Geonoma pohliana subsp. pohliana

Geonoma macroclona Drude (1882: 486). Type: BRAZIL. Bahia: near Ilhus, no date, M. Lhotzky s.n. (holotype BR n.v.,
isotype G, image!).
Geonoma blanchetiana Wendland ex Drude (1882: 494). Type: BRAZIL. Bahia: no locality, no date, J. Blanchet s.n.
(holotype BR!).
Geonoma luetzelburgii Burret (1930a: 235). Type: BRAZIL. Rio de Janeiro: Serra da Estrella, 1400 m, August 1915, P.
Luetzelburg 6073 (holotype M!).
Geonoma bondariana Lorenzi (2010: 221). Type: BRAZIL. Bahia: Itacar, Fazenda Petizeiro, estrada IlhusItacar,
1423S, 3902W, 61 m, 9 February 2009, H. Lorenzi, J. Jardim, A. Guimares 6619 (holotype HPL n.v., isotypes
CEPEC n.v., ESA n.v., NY n.v., RB n.v., SP n.v.), synon. nov.
Geonoma littoralis Noblick & Lorenzi in Lorenzi (2010: 226). Type: BRAZIL. Bahia: Mun. Itacar, Fazenda So
Miguel, 7 July 2009, H. Lorenzi, R. Pimenta, T. Flores & A. Guimares 6709 (holotype HPL n.v., isotypes CEPEC
n.v., ESA n.v., NY n.v., SP n.v.), synon. nov.
Geonoma meridionalis Lorenzi (2010: 240). Type: BRAZIL. Santa Catarina: Corup, estrada para Itapocu-hansa,
2625S, 4911W, 24 February 2010, H. Lorenzi, K. Soares & T. Flores 6834 (holotype HPL n.v., isotypes ESA n.v.,
NY n.v., RB n.v., SP n.v.), synon. nov.
Leaves pinnae 6(342) per side of rachis. Inflorescences rachillae 11(332), 2.9(1.14.4) mm in diameter, not
hairy, not filiform and not or scarcely narrowed between the flower pits; proximal and distal lips drying darker
brown than the rachillae; triads mostly spirally arranged.
Distribution and habitat:From 3542941S and 35525128W in the Atlantic Coastal Forest of
Brazil from Cear, Pernambuco, and Alagoas to Rio Grande do Sul at 450(71000) m elevation in brejo
vegetation or lowland tropical rainforest (Fig. 33).
An extremely variable subspecies consisting of several morphotypes. In the northern-most part of the
range there are isolated populations in Cear, and in Pernambuco and Alagoas, occurring in brejo vegetation
(northern morphotype), but too few specimens to test for differences.
Specimens from central Bahia (bahia morphotype) in coastal forest at low elevations have large, pinnate
leaves with 42 pinnae per side of the rachis (data from only one specimen). Inflorescences are large, with
wide, long peduncles and long rachillae.
Specimens from scattered localities in Bahia in restinga vegetation near sea level (littoralis morphotype)
have pinnate leaves with distinctive narrow pinnae. The type of Geonoma littoralis is of this morphotype.
Some specimens from central Bahia (e.g., Noblick 4747, Thomas 14115) are sympatric with and similar to
those of subsp. rubescens. Some specimens from southern Bahia (e.g., Noblick 4778, 4789, Harley 17853) are
sympatric with and similar to those of subsp. linharensis. The differences between these subspecies in these
areas are not clear.
127
A specimen (Amorim 4208) from central Bahia has unusual, elongate bracts, and is from a higher
elevation (750 m). It may be a hybrid with G. pauciflora.
There is a gap in the distribution of subsp. pohliana in northern Esprito Santo, and this gap is occupied by
subsp. linharensis. There are no differences between northern and southern populations of subsp. pohliana,
although northern populations occur at lower mean elevations (287 versus 624 m).
Five specimens (dos Santos s. n., Fernandes 1135, 3107, Fraga 2117, MelloSilva 1570) from the central
part of Esprito Santo in the Serra da Mantequeira appear to be intermediate in morphology between subsp.
pohliana and subsp. schottiana and may be hybrids between these subspecies.
Some specimens (e.g., Fiaschi 163, 552) from eastern So Paulo have more, narrower pinnae (mean of 8
versus 4) and appear larger than usual.
Two specimens (Lombardi 348, 1550), unplaced for morphotype, from the Rio Doce valley in Minas
Gerais have three broad pinnae and narrow inflorescence bracts.
51b. Geonoma pohliana subsp. fiscellaria (Martius ex Drude & Wendland) Henderson, comb. & stat.
nov.
Basionym: Geonoma fiscellaria Martius ex Drude (1882: 486). Type: BRAZIL. Rio de Janeiro: Retiro de Petropolis, no
date, A. Glaziou 1180 (holotype BR!, isotype P!).
Leaves pinnae 21(1923) per side of rachis. Inflorescences rachillae 8(1421), 3.7(3.04.8) mm in diameter,
hairy, not filiform and not or scarcely narrowed between the flower pits; proximal and distal lips drying the
same color as the rachillae; triads mostly spirally arranged.
Distribution and habitat:At 22002228S and 42034427W in the Atlantic Coastal Forest of
Brazil in the Serra do Mar and Serra da Mantequeira in Rio de Janeiro at 1050(9001100) m elevation in
montane tropical rainforest (Fig. 33).
51c. Geonoma pohliana subsp. gastoniana (Glaziou ex Drude) Henderson, comb. & stat. nov.
Basionym: Geonoma gastoniana Glaziou ex Drude (1882: 496). Type: BRAZIL. Rio de Janeiro: haut de Tingu, 7 June
1877, A. Glaziou 9019 (holotype P!, isotypes BR!, C n.v., FI!, K!).
Leaves pinnae 3 per side of rachis. Inflorescences rachillae 19, 0.5 mm in diameter, not hairy, filiform,
narrowed between the flower pits; proximal and distal lips drying darker brown than the rachillae; triads
mostly spirally arranged.
Distribution and habitat:At 2235S and 4328W in the Atlantic Coastal Forest of Brazil in the Serra
do Mar in Rio de Janeiro at 900 m elevation in lowland tropical rainforest (Fig. 34).
51d. Geonoma pohliana subsp. kuhlmannii (Burret) Henderson, comb. & stat. nov.
Basionym: Geonoma kuhlmannii Burret (1938b: 261). Type: BRAZIL. Rio de Janeiro: Nova Friburgo, 20 November
1922, J. Kulhmann 141 (holotype B, destroyed, isotype RB!).
hairy, not filiform and not or scarcely narrowed between the flower pits; proximal and distal lips drying darker
brown than the rachillae; triads mostly spirally arranged.
of Brazil in the Serra do Mar in Rio de Janeiro at 783(3001300) m elevation in lowland tropical rainforest
(Fig. 34).
51e. Geonoma pohliana subsp. linharensis Henderson, subsp. nov. (Appendix IV, Plate 54)
A subspeciebus aliis pinnis in quoque latere 4(27) atque rachillis 9(510) crebre pilosis differt.
128
HENDERSON
Type: BRAZIL. Esprito Santo: Reserva Florestal da CVRD, Linhares, 15 March 1989, G. Farias 258 (holotype
MBML!, isotype NY!).
densely hairy, not filiform and not narrowed between the flower pits; proximal and distal lips drying the same
color as the rachillae; flower pits mostly spirally arranged.
of Brazil in southern Bahia and northern Esprito Santo at 35(3040) m elevation in lowland rainforest (Fig.
34).
FIGURE 34. Distribution maps of Geonoma pohliana subsp. gastoniana, G. pohliana subsp. kuhlmannii, G. pohliana
subsp. linharensis, and G. pohliana subsp. rodriguesii.
51f. Geonoma pohliana subsp. rodriguesii Henderson, subsp. nov. (Appendix IV, Plate 55)
A subspeciebus aliis pinnis in quoque latere 4(27) atque rachillis 2(14) glabris differt.
Type: BRAZIL. Rio de Janeiro: Praia de Joo Gago, on Sepetiba Bay, on road to Mangaratiba, 2256S 4359W, 19
November 1966, G. Gottsberger 14191166 (holotype NY!).
hairy, not filiform and not or scarcely narrowed between the flower pits; proximal and distal lips drying the
same color as the rachillae; triads mostly spirally arranged.
129
of Brazil in Esprito Santo, Minas Gerais, Rio de Janeiro, and So Paulo at 251(50650) m elevation in
51g. Geonoma pohliana subsp. rubescens (Wendland ex Drude) Henderson, comb. & stat. nov.
Basionym: Geonoma rubescens Wendland ex Drude (1882: 491). Type: BRAZIL. Bahia: Ilhus, no date, J. Blanchet s.n.
(holotype BR!, isotype P!).
Geonoma platycaula Drude & Trial in Drude (1882: 490). Type: BRAZIL. Bahia: without locality, no date, C. Martius
s.n. (holotype M!).
hairy, not filiform and not narrowed between the flower pits; proximal and distal lips drying the same color as
the rachillae; triads mostly spirally arranged.
of Brazil in central Bahia at 212(45380) m elevation in lowland tropical rainforest (Fig. 35).
Labels of some specimens (e.g., Carvalho 6775, Noblick 4726) describe the stems, leaves, and
inflorescences as reddish-tinged. Some specimens (e.g., Fiaschi 1051), sympatric with subsp. unaensis,
exactly resemble that subspecies except for the non-hairy rachillae. The differences between the two
subspecies are not clear.
51h. Geonoma pohliana subsp. trinervis (Drude & Wendland) Henderson, comb. & stat. nov.
Basionym: Geonoma trinervis Drude & Wendland in Drude (1882: 492). Type: BRAZIL. Rio de Janeiro: Serra dos
Orgos, May 1832, L. Riedel 734 (holotype BR!, isotypes F!, K!, M!, P!).
of Brazil in the Serra do Mar in Rio de Janeiro at medium elevations in lowland rainforest (Fig. 35).
51i. Geonoma pohliana subsp. unaensis Henderson, subsp. nov. (Appendix IV, Plate 56)
A subspeciebus aliis pinnis in quoque latere 4(27) atque rachillis 2(23) crebre pilosis differt.
Type: BRAZIL. Bahia: Mun. Una, Reserva Biolgico do Mico-leo, entrada no km 46 da Rod. BA-001 Ilhus-Una,
1509S, 3905W, 812 March 1993, A. Amorim, S. SantAna, J. Jardim, E. Santos & J. Hage 1119 (holotype NY!,
isotype CEPEC n.v.).
Leaves pinnae 3(34) per side of rachis. Inflorescences rachillae 2(23), 6.3(5.56.8) mm in diameter, densely
of Brazil near Una, Bahia at low elevations in lowland rainforest (Fig. 35).
51j. Geonoma pohliana subsp. weddelliana (Wendland ex Drude) Henderson, comb. & stat. nov.
Basionym: Geonoma weddelliana Wendland ex Drude (1882: 494). Type: BRAZIL. Gois: between Gois and Cuiaba,
November-December 1844, H. Weddell 2983 (holotype P!, isotype F!).
Geonoma schottiana var. palustris Warming ex Drude (1882: 493). Geonoma warmingii Hawkes (1952: 189). Type:
BRAZIL. Minas Gerais: Lagoa Santa, no date, E. Warming 1843 (holotype C n.v., holotype image!).
Geonoma stenoschista Burret (1930a: 233). Type: BRAZIL. Minas Gerais: between Rio Parauna and Serra do Cip, 24
April 1892, A. Glaziou 20030 (holotype P!, isotype FI!).
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HENDERSON
Geonoma caudulata Loesener in Taubert (1896: 423). Type: BRAZIL. Gois: Serra dos Viadeiros, 17 October 1894, E.
Ule 226 (3150) (holotype P!).
Geonoma plurinervia Burret (1940b: 99). Type: BRAZIL. Mato Grosso: Capo Bonito, Campo Grande, 5 September
1936, W. Archer & A. Gehrt 74 (holotype B, destroyed, isotypes BH!, SP!, US!).
Geonoma telesana Lorenzi (2010: 252). Type: BRAZIL. Mato Grosso: Mun. Matinha, Serra do Roncador, 11 December
2009, H. Lorenzi, J. Teles, K. Suares & T. Flores 6789 (holotype HPL n.v., isotypes ESA n.v., SP n.v.), synon. nov.
not hairy, not filiform and not or scarcely narrowed between the flower pits; proximal and distal lips drying
darker brown than the rachillae; triads mostly decussately arranged.
Distribution and habitat:From 6002407S and 36006049W in the Cerrado region and
adjacent areas of campo rupestre of Brazil and just reaching adjacent Bolivia and Paraguay at 1015(200
1650) m elevation in wet places, usually in gallery forest (Fig. 35).
FIGURE 35. Distribution maps of Geonoma pohliana subsp. rubescens, G. pohliana subsp. trinervis, G. pohliana subsp.
unanensis, and G. pohliana subsp. weddelliana.
There is geographical variation in this subspecies. Linear regression shows there are significant
associations between elevation and eight leaf variables and four inflorescence variables. Squared multiple R
for the regression of leaf number on elevation is 0.58, rachis width 0.10, number of pinnae 0.28, basal pinna
length 0.18, basal pinna width 0.40, basal pinna angle 0.37, apical pinna width 0.31, apical pinna angle 0.19,
131
peduncle width 0.15, number of rachillae 0.12, fruit length 0.33, and fruit diameter 0.45. In particular, number
of pinnae increases and they become shorter and narrower with narrower angles with increasing elevation,
giving a distinctive, small pinnate leaf. For inflorescences, peduncles become thinner, rachillae fewer, and
fruits larger with increasing elevation. Specimens from higher elevations (10001600 m) in Minas Gerais
(Serra do Cip), Gois (Chapada dos Veadeiros), and Bahia (Pico das Almas) have these kinds of leaves and
inflorescences.
Specimens from the western and northern margins of the range have fewer, broader pinnae. Some
specimens (e.g., Noblick 3209, Sant'Ana 311, Thomas 9251) from the eastern margin of the range in Bahia
occur near to the range of subsp. pohliana, and there may be hybrids between these two subspecies in this
area.
Some specimens (Irwin 6276, 15633) from the Distrito Federal near Braslia, have pinnae more like those
of subsp. schottiana. It is unclear if this subspecies is present in the Distrito Federal, or if these specimens are
hybrids.
51k. Geonoma pohliana subsp. wittigiana (Glaziou ex Drude) Henderson, comb. & stat. nov.
Basionym: Geonoma wittigiana Glaziou ex Drude (1882: 499). Type: BRAZIL. Rio de Janeiro: Serra dos Orgos, 23
August 1872, A. Glaziou 6458 (holotype P!, isotypes C n.v., FI!, K!).
Leaves pinnae 4(36) per side of rachis. Inflorescences rachillae 7(310)1.8(1.22.6) mm in diameter, hairy,
not filiform and not or scarcely narrowed between the flower pits; proximal and distal lips drying the same
color as the rachillae; triads mostly spirally arranged.
region of Brazil in Esprito Santo, Minas Gerais, and Rio de Janeiro at 955(1751265) m elevation in lowland
or montane rainforest (Fig. 36).
Most specimens are from the Serra do Mar in Rio de Janeiro. The outliers from Minas Gerais and Esprito
Santo appear somewhat different but there are too few specimens to test for differences.
52. Geonoma poiteauana Kunth (1841: 233). Gynestum acaule Poiteau (1822: 391). Geonoma poiteana
Martius (1843: 39). Geonoma acaulis (Poiteau) Burret (1930a: 162). Geonoma macrostachys var. poiteauana
(Kunth) Henderson (1995: 277). Type: FRENCH GUIANA. Without locality, no date, A. Poiteau s.n.
(holotype P!).
Geonoma dammeri Huber (1902: 409). Taenianthera dammeri (Huber) Burret (1930c: 13). Type: BRAZIL. Par: Furo
Macujubim, 6 October 1901, M. Guedes 2241 (holotype MG!).
Geonoma chaunostachys Burret (1931c: 318). Type: VENEZUELA. Amazonas: Mount Duida, ca. 250 m, 18 November
1928, G. Tate 394 (holotype NY!).
Plants 1.2(0.52.0) m tall; stems 0.1 m tall, 2.1 cm in diameter, branching no data, not cane-like; internodes
0.2 cm long, not scaly. Leaves undivided or irregularly pinnate, not plicate, bases of blades running diagonally
into the rachis; sheaths 13.7(12.017.0) cm long; petioles 21.6(18.026.7) cm long, drying green or yellowish;
rachis 57.0(38.079.5) cm long, 4.2(3.15.9) mm in diameter; adaxial veins not raised or slightly raised and
16.9(9.531.5) cm wide, forming an angle of 18(1024) with the rachis. Inflorescences unbranched;
80.2(54.5119.5) cm long, 2.9(1.33.7) mm in diameter; rachillae 1, 13.2(8.520.0) cm long, 4.8(3.16.0)
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HENDERSON
staminate flowers deciduous after anthesis; stamens 6; thecae diverging or not diverging at anthesis, inserted
onto well-developed, non-split, jointed connectives, connectives alternately long and short; anthers short at
anthesis, remaining straight and parallel; non-fertilized pistillate flowers deciduous after anthesis; staminodial
tubes lobed at the apex, the lobes spreading at anthesis, acuminate, those of non-fertilized pistillate flowers
emerging, not bumpy, not apiculate; locular epidermis with operculum, smooth, with pores.
Distribution and habitat:From 745N800S and 47107011W in the eastern and central
Amazon region of the Guianas, Venezuela, Colombia, and Brazil, at 242(1725) m elevation in lowland
Taxonomic notes:Geonoma poiteauana was recognized by Henderson (1995) as a variety of Geonoma
macrostachys, but is here recognized at the species level. The two are closely related, G. poiteauana differing
by its fruits which are not bumpy and not apiculate. It belongs to a group of species within the G.
macrostachys clade, comprising G. macrostachys, G. multisecta, G. paradoxa, and G. schizocarpa.
Subspecific variation:Only one trait (leaf division) varies within this species.
FIGURE 36. Distribution maps of Geonoma pohliana subsp. wittigiana, G. poiteauana, G. sanmartinensis, and G.
santanderensis.
133
53. Geonoma sanmartinensis Henderson, sp. nov. (Appendix IV, Plate 57)
A speciebus affinibus crusta fructuum fibris epidermalibus brevibus numerosis apicem convergentibus differt.
Type: PERU. San Martn: Prov. Rioja, Dist. Naranjillo, sector San Augustin, Bosque de Proteccin de Alto Mayo,
548S, 7721W, 1250 m, 13 November 1996, I. Snchez Vega & M. Dillon 8658 (holotype NY!, isotypes CPUN
n.v., F!).
Plants 1.2(0.52.0) m tall; stem branching no data; stem type no data; stem height no data; stem diameter no
data; internode color no data; internode length no data. Leaves irregularly pinnate, not plicate, bases of blades
running diagonally into the rachis; sheaths no data; petioles drying green or yellowish; rachis 38.6(35.841.0)
60) with the rachis; apical pinna length no data, 12.3(9.215.0) cm wide, forming an angle of 27(2232)
with the rachis. Inflorescences unbranched; prophylls and peduncular bracts ribbed with elongate, unbranched
fibers, both bracts tubular, narrow, elongate, closely sheathing the peduncle, more or less persistent; prophylls
12.6(11.014.2) cm long, not short and asymmetrically apiculate, the surfaces not ridged, without unequally
wide ridges; peduncular bracts 23.0(16.030.0) cm long, well-developed; inserted 1.1(0.81.4) cm above the
prophyll; peduncles 63.5(63.064.0) cm long, 2.9(2.63.1) mm in diameter; rachillae 1, 21.1(18.524.7) cm
long, 3.6(3.13.9) mm in diameter, the surfaces with spiky, fibrous projections or ridges, drying brown or
yellow-brown, without short, transverse ridges, not filiform and not narrowed between the flower pits; flower
pits spirally arranged, glabrous internally; proximal lips without a central notch before anthesis, not recurved
after anthesis, hood-shaped at anthesis, sometimes splitting post-anthesis; proximal and distal lips drying the
lip margins; distal lips a scarcely raised rim; staminate and pistillate petals not emergent, not valvate
almost directly onto the filament apices, the connectives bifid but scarcely developed; anthers short and curled
over at anthesis; non-fertilized pistillate flowers deciduous after anthesis; staminodial tubes crenulate or
shallowly lobed at the apex, those of non-fertilized pistillate flowers not projecting and persistent after
anthesis; fruits size no data, the bases without a prominent stipe, the apices not conical, the surfaces not
fibers present, these coming to a point at fruit apices; locular epidermis without operculum, smooth, without
pores.
Distribution and habitat:From 545554S and 77217745W in Andean regions of Peru (San
Martn) at 1525(12501850) m elevation in montane rainforest (Fig. 36).
Taxonomic notes:Geonoma sanmartinensis appears most similar to G. brongniartii. It differs in its
fruits which are bumpy from the numerous, subepidermal, tangential, short fibers present.
54. Geonoma santanderensis Galeano & Bernal (2002: 282). Type: COLOMBIA. Santander: Suaita, San
Jos de Suaita, ca. 610N, 7327W, 17001900 m, 30 July 2001, G. Galeano, J. Betancur, N. Castao, L.
Clavijo & N. Garcia 6884 (holotype COL!, isotypes HUA n.v., NY n.v., UIS n.v.).
Plants height no data; stems 1.3(1.01.5) m tall, 0.8(0.60.9) cm in diameter, solitary or clustered, cane-like;
internodes 1.5(1.02.2) cm long, covered with reddish or brownish scales, especially in their distal part.
25.0) cm long, 1.6(1.42.2) mm in diameter; veins not raised or slightly raised and triangular in cross-section
134
HENDERSON
adaxially; pinnae 5(36) per side of rachis; basal pinna 10.7(9.711.5) cm long, 1.1(0.81.8) cm wide,
forming an angle of 60(4572) with the rachis; apical pinna 8.2(7.010.0) cm, 7.5(6.311.0) cm wide,
forming an angle of 42(3848) with the rachis. Inflorescences unbranched; prophylls and peduncular bracts
not ribbed with elongate, unbranched fibers, flattened, deciduous or persistent; prophylls 8.5(6.711.2) cm
long, not short and asymmetrically apiculate, the surfaces not ridged, without unequally wide ridges;
peduncles 8.7(6.014.5) cm long, 2.1(1.52.8) mm in diameter; rachillae 1, 8.4(5.710.0) cm long, 3.4(2.6
4.1) mm in diameter, the surfaces with spiky, fibrous projections or ridges, drying brown or yellow-brown,
without short, transverse ridges, not filiform and not narrowed between the flower pits; flower pits usually
spirally arranged, glabrous internally; proximal lips without a central notch before anthesis, not recurved after
anthesis; stamens 6; thecae diverging at anthesis, inserted onto bifid and well-developed, non-jointed
connectives; anthers short and curled over at anthesis; non-fertilized pistillate flowers deciduous after
Distribution and habitat:From 610708N and 72547327W in the Eastern Cordillera in
Colombia at 1800 m elevation in montane rainforest (Fig. 36).
Taxonomic notes:Geonoma santanderensis was compared to G. monospatha by Galeano and Bernal
(2002). However, it appears closely related to a group of species within the G. stricta clade comprising G.
aspidiifolia and G. oligoclona. All three species share internodes covered with reddish or brownish scales,
especially in their distal part, rachillae surfaces with spiky, fibrous projections or ridges, and staminodial tubes
lobed at the apex with the lobes not spreading at anthesis and not acuminate. Geonoma santanderensis differs
from G. aspidiifolia in its fruits which are bumpy from the numerous, subepidermal, tangential, short fibers
present; and from G. oligoclona in its prophyll which is not short and asymmetrically apiculate.
Subspecific variation:No traits, except for stem branching, vary within this species. The specimens
come from two separate areas, but there are too few specimens to test for differences between these.
55. Geonoma schizocarpa Henderson, sp. nov. (Appendix IV, Plates 58 & 59)
A speciebus affinibus crusta fructuum profunde scindens differt.
Type: PERU. Amazonas: al lado de Huampami, 18 July 1974, R. Kyap 1212 (holotype MO!).
Plants 2.2(1.05.0) m tall; stem height no data, 2.8 cm in diameter, branching no data, not cane-like;
internodes 0.3 cm long, not scaly. Leaves irregularly pinnate, not plicate, bases of blades running diagonally
into the rachis; sheaths 12.0 cm long; petioles drying green or yellowish; rachis 90.0(88.092.0) cm long,
7.9(6.28.7) mm in diameter; veins raised and rectangular in cross-section adaxially; pinnae 14 per side of
rachis; basal pinna 34.0 cm long, 1.2(0.81.6) cm wide, forming an angle of 67(5182) with the rachis;
apical pinna 20.0 cm long, 4.9(2.86.5) cm, forming an angle of 31(3033) with the rachis. Inflorescences
unbranched; prophylls and peduncular bracts ribbed with elongate, unbranched fibers, both bracts tubular,
narrow, elongate, closely sheathing the peduncle, more or less persistent; prophylls 14.5 cm long, not short
length no data, well-developed, inserted 0.6 cm above the prophyll; peduncles 93.0 cm long, 5.7(4.67.2) mm
135
cupule, the proximal lip margins overlapping the distal lip margins; distal lips well-developed; staminate
flowers persistent post-anthesis or deciduous post-anthesis; staminate and pistillate petals not emergent, not
valvate throughout; stamens 6; thecae not diverging at anthesis, inserted onto well-developed, non-split,
jointed connectives, alternately long and short; anthers short at anthesis, remaining straight and parallel; nonfertilized pistillate flowers deciduous after anthesis; staminodial tubes lobed at the apex, the lobes spreading at
anthesis, acuminate, those of non-fertilized pistillate flowers not projecting and persistent after anthesis; fruits
conical, the surfaces splitting deeply and longitudinally at maturity to reveal mesocarp with dense layer of
radial fibers, without fibers emerging, not bumpy, not apiculate; locular epidermis with operculum, smooth,
without pores.
Distribution and habitat:From 415430S and 78107830W in Peru (Amazonas) at 225 m
FIGURE 37. Distribution maps of Geonoma schizocarpa, G. schottiana, G. scoparia, and G. simplicifrons.
Taxonomic notes:Geonoma schizocarpa is a member of the G. macrostachys clade. Within this it is

most closely related to G. macrostachys, G. multisecta, G. paradoxa, and G. poiteauana. It differs from these in
136
HENDERSON
its fruits surfaces which split deeply and longitudinally at maturity to reveal mesocarp with a dense layer of
radial fibers.
Subspecific variation:One trait (staminate flower persistence) varies within this species. Two of 10
specimens have deciduous staminate flowers.
56. Geonoma schottiana Martius (1826: 143). Type: BRAZIL. Rio de Janeiro: no locality, no date, H. Schott
4111 (holotype M!).
Geonoma schottiana var. angustifolia Drude (1882: 492). Lectotype (here designated): BRAZIL. Rio de Janeiro:
Petropolis, 6 April 1877, A. Glaziou 9011 (lectotype BR!, duplicates K!, FI!, P!).
Geonoma schottiana var. latifolia Drude (1882: 492). Lectotype (here designated): BRAZIL. Rio de Janeiro: Restinga de
Mau, 8 June 1876, A. Glaziou 8493 (lectotype BR!, duplicate P!).
Geonoma hoehnei Burret (1930a: 231). Type: BRAZIL. So Paulo: Japuhyba, 17 April 1926, F. Hoehne & A. Gehrt
17391 (holotype SP!).
Plants 2.9(1.56.0) m tall; stems 1.7(0.33.5) m tall, in diameter, no data, solitary, cane-like; internodes no
data, yellowish and smooth. Leaves 15(724) per stem, regularly pinnate, the pinnae with 1 main vein and 2
lateral veins on either side of main vein, not plicate, bases of blades running diagonally into the rachis; sheaths
26.9(23.533.0) cm long; petioles 57.8(30.087.0) cm long, drying green or yellowish; rachis 70.4(44.0
118.0) cm long, 5.1(2.68.9) mm in diameter; veins raised and rectangular in cross-section adaxially; pinnae
26(1038) per side of rachis; basal pinna 36.8(17.559.0) cm long, 0.5(0.22.1) cm wide, forming an angle of
49(2590) with the rachis; apical pinna 22.4(12.536.5) cm long, 3.0(0.610.2) cm wide, forming an angle
of 21(734) with the rachis. Inflorescences unbranched or branched 13 orders; prophylls and peduncular
bracts not ribbed with elongate, unbranched fibers, flattened, deciduous or persistent; prophylls 26.4(16.0
peduncles 37.7(18.073.5) cm long, 5.1(1.610.6) mm in diameter; rachillae 16(169), 20.6(7.537.0) cm
long, 1.9(0.73.5) mm in diameter, the surfaces without spiky, fibrous projections or ridges, drying brown or
pits tricussately arranged, the groups not closely spaced nor consistently arranged throughout the rachillae,
hood-shaped; proximal and distal lips drying darker brown than the rachillae, not joined to form a raised
scarcely developed; anthers short and curled over at anthesis; non-fertilized pistillate flowers deciduous after
bases with a prominent, asymmetric stipe, the apices conical with rounded apices, the surfaces not splitting at
maturity, without fibers emerging, not bumpy, not apiculate; locular epidermis without operculum, smooth,
without pores.
and inland areas of southeastern Brazil (Esprito Santo, Minas Gerais, Paran, Rio de Janeiro, Rio Grande do
Sul, Santa Catarina, So Paulo) at 747(31600) m elevation in lowland to montane tropical rainforest or
gallery forest (Fig. 37).
Taxonomic notes:Geonoma schottiana is a member of a group of species from the Atlantic Coastal
Forest and adjacent Cerrado (the G. schottiana clade, also including G. elegans, G. pauciflora, and G.
pohliana). Although the group is well-supported, all constituent species are extremely variable internally.
Geonoma schottiana differs from other species in the group by its regularly pinnate leaves, the pinnae with 1
main vein and 2 lateral veins on either side of main vein.
137
Subspecific variation:Only one trait varies within this species (inflorescence branching). One
specimen (of 76) has an unbranched inflorescence. There is geographic discontinuity.
Geonoma schottiana occurs in two areas in Brazilthe Atlantic Coastal Forest at 31600 m elevation in
various habitats including restinga, lowland, and montane forest; and further inland in Minas Gerais in the
southern part of the Serra do Espinhao at 7601450 m elevation in gallery forest. Specimens from Minas
Gerais (minas morphotype) differ from those of other areas (excluding possible hybrids, see below) in 14
variables (plant height, stem height, rachis length, rachis width, number of pinnae, basal pinna length, basal
pinna angle, apical pinna length, interbract distance, peduncle length, peduncle width, rachilla length, rachilla
width, number of rachillae)(t-test, P <0.05). In particular, minas specimens have shorter interbract distances
(mean of 8.9 cm versus 18.2 cm), shorter peduncles (mean of 24.8 cm versus 39.2 cm), and more rachillae
(mean of 42 versus 17). The minas morphotype occurs sympatrically with G. pauciflora subsp. weddelliana.
Eleven specimens from the Serra do Castelo in Esprito Santo (castelo morphotype) differ from others
from the Atlantic Coastal Forest in 11 variables (plant height, stem height, rachis width, basal pinna angle,
peduncular bract length, interbract distance, peduncle length, peduncle width, rachillae length, rachillae
width, number of rachillae)(t-test, P <0.05). In particular, these specimens have shorter (mean of 15.6 cm
versus 22.2 cm), thicker (mean of 2.6 mm versus 1.8 mm), and fewer (3 versus 17) rachillae than other
specimens. One specimen (Kollmann 9579) has only one rachilla. In these variables the castelo morphotype
resembles G. elegans, with which it occurs sympatrically. There may be introgression between these two in
this area.
One specimen (Fiaschi 613) from Rio de Janeiro has leaves like those of G. pauciflora subsp. pohliana
but inflorescences like those of G. schottiana. For other potential hybrids with G. pohliana, see under that
subspecies.
57. Geonoma scoparia Grayum & de Nevers (1988: 111). Type: COSTA RICA. Puntarenas: 7 km W of
Rincn de Osa, ridge between Ro Riyito and Quebrada Banegas, 841N, 8332W, 200300 m, G. de
Nevers, B. Hammel, & M. Grayum 7757 (holotype MO!, isotypes CAS n.v., CR!).
Plants 2.5(2.03.0) m tall; stems 2.5(2.03.0) m tall, 1.1(0.81.5) cm in diameter, solitary, cane-like;
internodes 1.5(0.82.5) cm long, covered with dense, brown scales. Leaves 16(1219) per stem, irregularly
mm in diameter; veins raised and rectangular in cross-section adaxially; pinnae 3 per side of rachis; basal
fibers, flattened, deciduous; prophylls 7.0(6.57.3) cm long, short, asymmetrically apiculate, the margins
curved around the stem, the surfaces flat with dense, felty, brown tomentum, prophyll equal to and early
deciduous with the peduncular bract, the surfaces not ridged, without unequally wide ridges; peduncular
bracts 7.0 cm long, well-developed, inserted 0.3(0.20.6) cm above the prophyll; peduncles 2.8(2.53.2) cm
long, 3.3(3.13.6) mm in diameter; rachillae 112(85138), 9.1(6.013.5) cm long, 0.6(0.41.0) mm in
short, transverse ridges, filiform with extended narrowed sections between the flower pits; flower pits
and distal lips drying the same color as the rachillae, joined to form a raised cupule, the margins not
overlapping; distal lips well-developed; staminate and pistillate petals not emergent, not valvate throughout;
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HENDERSON
6.0(5.86.1) mm long, 4.7 mm in diameter, the bases without a prominent stipe, the apices not conical, the
Distribution and habitat:From 833845N and 83068333W in Costa Rica on the Osa
Peninsula and adjacent areas at 245(100350) m elevation in lowland rainforest (Fig. 37).
Taxonomic notes:Geonoma scoparia is closely related to G. mooreana, the two species differing from
related species (the G. lanata clade) in their proximal and distal lips joined to form a raised cupule. Geonoma
scoparia differs from G. mooreana in its solitary stems and internodes covered with dense, brown scales.
Subspecific variation: No trait varies within this species nor is there any geographic disjunction.
58. Geonoma simplicifrons Willdenow (1805: 594). Type: VENEZUELA. Caracas, no date, F. Bredemeyer
20 (holotype B n.v., holotype image!).
Geonoma willdenowii Klotzsch (1846: 112). Type: VENEZUELA. Carobobo: near Puerto Cabello, 1846, H. Karsten s.n.
(holotype LE n.v., isotypes BM!, G!).
Geomoma fendleriana Spruce (1871: 108). Type: VENEZUELA. Vargas: between Petaquire and the sea, 9 February
1857, A. Fendler 2467 (holotype K!, isotype NY!).
cane-like; internodes 1.0(0.71.8) cm long, yellowish and smooth. Leaves undivided or irregularly pinnate,
pinna 19.3(13.032.0) cm long, 10.9(6.014.3) cm wide, forming an angle of 37(3043). Inflorescences
branched 1 order; prophylls and peduncular bracts not ribbed with elongate, unbranched fibers, flattened (if
tubular, narrow, and elongate then not ribbed), deciduous or persistent; prophylls 14.3(8.219.3) cm long, not
short and asymmetrically apiculate, the surfaces not ridged, without unequally wide ridges; peduncular bracts
12.0 cm long, well-developed, inserted 0.9(0.32.6) cm above the prophyll; peduncles 18.1(8.026.5) cm
long, 2.9(1.74.6) mm in diameter; rachillae 3(25), 14.1(8.019.5) cm long, 2.8(2.23.8) mm in diameter,
the surfaces without spiky, fibrous projections or ridges, drying brown, with faint to pronounced, short,
transverse ridges, not filiform and not narrowed between the flower pits; flower pits spirally arranged, densely
hairy internally distally only (rarely some hairs on lateral margins of the pit); proximal lips without a central
notch before anthesis, not recurved after anthesis, hood-shaped at anthesis, sometimes splitting post-anthesis;
lip margins overlapping the distal lip margins; distal lips absent; staminate and pistillate petals not emergent,
not valvate throughout; staminate flowers deciduous after anthesis; stamens 6; thecae diverging at anthesis,
and curled over at anthesis; non-fertilized pistillate flowers persistent or deciduous after anthesis; staminodial
locular epidermis without operculum, sculpted, usually also with a raised, meridional ridge, without pores.
Distribution and habitat:From 10041037N and 65487645W on the Coastal Cordillera in
Venezuela at 785(421600) m elevation in lowland to montane rainforest (Fig. 37).
139
Taxonomic notes:Geonoma simplicifrons is a member of a group of related species, the G. interrupta

clade, characterized by its lack of a distal lip of the flower pit and flower pits hairy internally (and not related
to G. jussieuana or G. orbignyana, contra Henderson et al., 1995). This group also includes G. euspatha, G.
frontinensis, G. interrupta, and G. pinnatifrons. These species have had a checkered taxonomic history but G.
simplicifrons has usually been recognized as a distinct species. It differs from G. euspatha, G. frontinensis, and
G. pinnatifrons in its flower pits which are densely hairy internally distally only; and from G. interrupta in its
prophyll surfaces which are not ridged and without unequally wide ridges.
Subspecific variation:Only one trait (leaf division) varies within this species. Specimens from the
eastern part of the range (east of 67W) have simple leaves with longer apical pinnae, but simple leaves also
occur, rarely, in the western part of the range. Most specimens from the western part of the range have pinnate
leaves.
Fred Stauffer (pers. comm.) considers that the maximum height of plants is 1.7 m, and the figure of 3 m is
a mistake on the specimen label; that inflorescences are sometimes unbranched (25 rachillae reported here);
and that the minimum elevation of this subspecies is about 700 m, and that the lower records are mistakes on
specimen labels.
59. Geonoma spinescens Wendland ex Burret (1930a: 230). Type: VENEZUELA. Nueva Grenada, no date,
L. Schlim s.n. (holotype B, destroyed). Lectotype (designated by Stauffer 1997): Dahlgren (1959), plate 286.
Geonoma tenuis Burret (1937a: 478). Type: VENEZUELA. Aragua: Rancho Grande, 1200 m, 8 January
1937, H. Pittier 13840 (holotype B, destroyed, isotypes F!, VEN n.v.).
Plants 1.0 m tall; stems 0.5(0.30.75) m tall, in diameter no data, solitary; internodes length no data,
yellowish and smooth. Leaves undivided, not plicate, bases of blades running diagonally into the rachis;
sheaths 18.0 cm long; petioles 26.3(15.040.0) cm long, drying green or yellowish; rachis 37.3(33.540.5) cm
long, 3.6(3.03.9) mm in diameter; veins not raised or slightly raised and triangular in cross-section adaxially;
pinnae 1 per side of rachis; basal pinna length and width not applicable, forming an angle of 25(1731) with
the rachis; apical pinna 22.4(18.525.8) cm long, width not applicable, forming an angle of 30(2830).
fibers, flattened, deciduous or persistent; prophylls no data; peduncular bracts length no data, well-developed,
inserted 0.4 cm above the prophyll; peduncles 28.5(27.529.5) cm long, 2.9(2.53.4) mm in diameter;
same color as the rachillae, joined to form a raised cupule, the margins not overlapping; distal lips welldeveloped; staminate and pistillate petals not emergent, not valvate throughout; staminate flowers deciduous
flowers deciduous after anthesis; staminodial tubes crenulate or shallowly lobed at the apex, those of nonfertilized pistillate flowers not projecting and persistent after anthesis; fruits 7.1 mm long, 5.5 mm in diameter,
Distribution and habitat:From 10201024N and 67416757W in the Coastal Cordillera in
Venezuela (Aragua, Carabobo) at 1075(8001330) m elevation in lowland or montane rainforest (Fig. 38).
Taxonomic notes:Geonoma spinescens is a member of a relatively large group of similar species, the
G. lanata clade. It differs from all of these, except G. gentryi, in its spirally arranged flower pits. See notes
under G. gentryi for further discussion.
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Subspecific variation: No trait varies within this species. According to Stauffer (1997) the leaves are
rarely irregularly pinnate with multi-veined pinnae. This species is scored as having the flower pits spirally
arranged, but some specimens, especially those from Carabobo, have almost distichously arranged flower pits.
FIGURE 38. Distribution maps of Geonoma spinescens, G. stricta subsp. stricta, G. stricta subsp. antioquiensis, and G.
stricta subsp. arundinacea.
60. Geonoma stricta (Poiteau) Kunth (1841: 232). Gynestum strictum Poiteau (1822: 391). Type: FRENCH
GUIANA. Without locality, no date, A. Poiteau s.n. (holotype P!).
undivided or irregularly pinnate, not plicate, bases of blades running diagonally into the rachis; sheaths
long, 2.6(0.96.0) mm in diameter; veins raised and rectangular in cross-section adaxially or not raised or
slightly raised and triangular in cross-section adaxially; pinnae 2(112) per side of rachis; basal pinna
12.6(3.238.5) cm long, 10.3(1.523.5) cm wide, forming an angle of 34(1450) with the rachis.
Inflorescences unbranched or branched 1 order; prophylls and peduncular bracts not ribbed with elongate,
141
9.6) cm long, vestigial, the prophyll three times or more long, sometimes the peduncular bract apparently
well-developed but then soon disintegrating, inserted 1.8(0.19.5) cm above the prophyll; peduncles 5.0(0.5
diameter, the surfaces with spiky, fibrous projections or ridges, drying brown or yellow-brown, without short,
transverse ridges, not filiform and not narrowed between the flower pits; flower pits spirally arranged,
pistillate petals not emergent, not valvate throughout; staminate flowers persistent or deciduous after anthesis;
stamens 6; thecae diverging at anthesis, inserted onto bifid and well-developed, non-jointed connectives;
anthers short and curled over at anthesis; non-fertilized pistillate flowers persistent after anthesis; staminodial
prominent stipe, the apices not conical, the surfaces not splitting at maturity, without fibers emerging, ridged
from the numerous, subepidermal, meridional, elongate fibers present, these coming to a point at fruit apices;
Taxonomic notes:Geonoma stricta is a member of the G. stricta clade, within which it is most closely
related to G. aspidiifolia, G. oligoclona, and G. santanderensis. It differs from these three species in several
character states, most obviously in its yellowish and smooth internodes. It is an extremely complicated
species, treated in the past as either one species with several varieties (e.g., Henderson, 1995) or several
species (e.g., Wessels Boer, 1968). Henderson and Martins (2002), in a morphometric study of this species,
concluded that the varietal classification proposed by Henderson (1995) was not supported. However, these
authors used only quantitative variables. With more specimens, and an analysis of both quantitative variables
and qualitative traits, as well as geography, some resolution is possible, as discussed below.
Subspecific variation:Six traits (stem branching, stem type, leaf division, adaxial veins, inflorescence
branching, staminate flowers) vary within this species. For stem type, only one of 390 specimens is scored as
not cane-like, and this and stem branching and leaf division are not used in the following analyses. There is no
correspondence between geography and the three remaining traits (adaxial veins, inflorescence branching,
staminate flowers). Geonoma stricta is widespread across the Amazon region and beyond. However, there are
gaps in its distribution, and G. stricta occurs in four separate areasthe Pacific coast of Colombia (Choc), the
Central Cordillera in Colombia (Antioquia), the central and western Amazon region and adjacent sub-Andean
regions, and the Guianas and adjacent Brazil (Amap). Specimens from these four regions are analyzed
separately.
There are only three specimens from the Pacific coast of Colombia (Choc), too few to test for
differences. However, these differ from the nearest other G. stricta population in the Central Cordillera in
Colombia, in their shorter interbract distances (0.10.2 cm versus 1.21.3 cm) and narrower rachillae (3.14.5
mm versus 4.87.4 mm). They also occur at lower elevations, 97(50150) m versus 852(7001075) m. Based
on these differences, and geographic separation, these specimens are recognized as a separate subspecies
(subsp. quibdoensis).
There are only six specimens from the Central Cordillera in Colombia (Antioquia), too few to test for
differences. Based on their geographic isolation, and differences from subsp. quibdoensis, they are recognized
as a separate subspecies (subsp. antioquiensis).
In central and western Amazon regions and adjacent sub-Andean regions, Geonoma stricta is abundant,
widespread, and extremely variable. In sub-Andean regions of Peru (Amazonas, Loreto, Hunuco, Pasco,
Ucayali) there are specimens with branched inflorescences (these occur rarely in other areas). One subgroup
of these, from Hunuco and Pasco, has leaves with raised adaxial veins, and this is recognized as a subspecies
(subsp. submontana). The remaining specimens, with non-raised adaxial veins, can be divided into three
subgroups, one from Amazonas with mostly undivided leaves and pendulous inflorescences; one from
Amazonas and Loreto with pinnate leaves and erect inflorescences; and one from Hunuco, Pasco, and
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HENDERSON
Ucayali with pinnate leaves and erect inflorescences. ANOVA shows that for pair wise comparison
probabilities, 13 variables (stem diameter, internode length, petiole length, rachis length, rachis width, number
of pinnae, basal pinna width, basal pinna angle, apical pinna width, peduncle length, peduncle width, rachilla
length, rachilla width) differ significantly (P <0.05) between one pair of subgroups, and one (prophyll length)
differs amongst all three subgroups. Based on these results, these three subgroups are recognized as
subspecies (subspp. bracteata, divaricata, pendula).
The remaining specimens from the central and western Amazon region and eastern Andean slopes in
Ecuador cannot be divided into consistent groups based on traits or geography. Adaxial veins are difficult to
score in several cases; inflorescences are seldom branched, but both branched and unbranched ones can occur
on the same plant; and staminate flower persistence is also difficult to score. For these reasons, these
specimens are recognized as one subspecies (subsp. arundinacea).
In the Guianas and adjacent Brazil (Amap) there are two subgroups of specimens, one with pinnate
leaves and raised adaxial veins and the other with undivided (rarely pinnate) leaves and non-raised adaxial
veins. There are only four specimens of the subgroup with pinnate leaves and raised adaxial veins, too few to
test for differences. However, this subgroup is geographically isolated from the other, and the two are
recognized as subspecies (subsp. stricta, pliniana).
Key to the subspecies of Geonoma stricta
1 Pacific coast of Colombia (Choc) and Central Cordillera in Colombia (Antioquia) ...................................................... 2
- All other areas .................................................................................................................................................................... 3
2 Pacific coast of Colombia (Choc) .......................................................................................................subsp. quibdoensis
- Central Cordillera in Colombia (Antioquia) ........................................................................................subsp. antioquiensis
3 Eastern Andean slopes and central and western Amazon region...................................................................................... 4
- Northeastern Amazon region of the Guianas and Brazil (Amap) ................................................................................... 8
4 Inflorescences unbranched or branched; central and western Amazon region, or eastern Andean slopes in Ecuador.......
.........................................................................................................................................................subsp. arundinacea
- Inflorescences branched; eastern Andean slopes in northern (Amazonas, Loreto) and central (Hunuco, Pasco, Ucayali)
Peru ............................................................................................................................................................................... 5
5 Leaves mostly undivided; inflorescences pendulous; Amazonas ............................................................... subsp. pendula
- Leaves pinnate; inflorescences erect; Amazonas, Hunuco, Loreto, Pasco, Ucayali ....................................................... 6
6 Veins raised and rectangular in cross-section adaxially; Hunuco, Pasco ..................................................... submontana
- Veins not raised or slightly raised and triangular in cross-section adaxially; Amazonas, Hunuco, Loreto, Pasco, Ucayali .................................................................................................................................................................................. 7
7 Amazonas, Loreto ................................................................................................................................... subsp. divaricata
- Hunuco, Pasco, Ucayali ..........................................................................................................................subsp. bracteata
8 Veins raised and rectangular in cross-section adaxially; leaves pinnate .....................................................subsp. pliniana
- Veins not raised or slightly raised and triangular in cross-section adaxially; leaves undivided (rarely pinnate) ................
...................................................................................................................................................................subsp. stricta
60a. Geonoma stricta subsp. stricta

Geonoma maguirei Bailey (1948: 102). Type: SURINAME. Coppename River headwaters, 24 July 1944, B. Maguire
24166 (holotype NY!).
Leaves undivided, rarely pinnate; veins not raised or slightly raised and triangular in cross-section adaxially;
pinnae 1(13) per side of rachis. Inflorescences unbranched, rarely branched; staminate flowers deciduous
after anthesis, rarely persistent.
Distribution and habitat:From 220710N and 52505850W in the northeastern Amazon
region of Suriname and French Guiana, with an outlying specimen in Guyana, at 373(100800) m elevation in
Two specimens from French Guiana (de Granville 4803, Le Prieur s. n.) are unusual in having branched
inflorescences with 23, stouter rachillae, and one has a pinnate leaf (the only one in this subspecies). They
143
are recognized as the neglecta morphotype. One of these specimens (Le Prieur s. n.) has Geonoma neglecta
written, in Trails hand-writing, on the label. This specimen was illustrated by Wessels Boer (1968, plate V)
and determined by him as G. bartlettii. However, the type of that name was destroyed and it is treated here as
an excluded name. Two specimens from Suriname (Sastre 1567, Wessels Boer 1558), with larger leaves and
inflorescences with persistent staminate flowers appear more like those of the pycnostachys morphotype of
subsp. arundinacea. These are recognized as the large morphotype.
60b. Geonoma stricta subsp. antioquiensis Henderson, subsp. nov. (Appendix IV, Plate 60)
A subspeciebus extraamazonicis aliis bractea pedunculare altior supra prophyllo inserta differt.
Type: COLOMBIA. Antioquia: Mun. San Luis, autopista Medelln-Bogot, sector Ro Samana-Ro Claro, hacia
Aquitania, entrando por vereda Altavista, a 5 km de la autopista, 700800 m, 27 September 1986, A. Cogollo & R.
Torres 2397 (holotype COL!).
Leaves undivided; veins not raised or slightly raised and triangular in cross-section adaxially; pinnae 1 per
side of rachis. Inflorescences unbranched; staminate flowers deciduous after anthesis.
Colombia (Antioquia) at 852(7001075) m elevation in lowland rainforest (Fig. 38).
60c. Geonoma stricta subsp. arundinacea (Martius) Henderson, comb. & stat. nov.
Basionym: Geonoma arundinacea Martius (1823: 17). Lectotype (selected by Wessels Boer 1968): BRAZIL. Amazonas:
Rio Negro, no date, C. Martius s.n. (lectotype M!).
Geonoma pycnostachys Martius (1823: 16). Type: BRAZIL. Amazonas: Rio Japur, no date, C. Martius s.n. (holotype
M!).
Geonoma stricta var. trailii (Burret) Henderson (1995: 288). Geonoma trailii Burret (1930a: 178). Geonoma elegans
Martius var. amazonica Trail (1876: 324). Type: BRAZIL. Amazonas: Rio Purus, Barreiras de Mancira, 29
September 1874, J. Trail 1032/CXXXIII (holotype K!).
Geonoma piscicauda Dammer (1907: 123). Geonoma stricta var. piscicauda (Spruce) Henderson (1995: 287). Type:
BRAZIL. Acre: Rio Juru, Juru-mirim, May 1901, E. Ule 5520 (holotype B, destroyed, isotypes F!, K!, MG!).
Geonoma wittiana Dammer (1907: 124). Type: BRAZIL. Acre: Rio Juru, Juru-mirim, September 1901, E. Ule 5884
(holotype B, destroyed, isotype MG!).
Geonoma uleana Dammer (1907: 122). Type: BRAZIL. Acre: Rio Juru, Cachoeira, May 1901, E. Ule 5521 (holotype
B, destroyed, isotype MG!).
Geonoma trauniana Dammer (1907: 124). Type: BRAZIL. Acre or Amazonas: Rio Juru, Fortaleza, October 1901, E.
Ule 5946 (holotype B, destroyed, isotype MG!).
Geonoma dasystachys Burret (1930a: 251). Type: BRAZIL. Amazonas: Rio Negro, Jauapasse assu, 5 July 1874, J. Trail
981/XC (holotype K!).
Geonoma bella Burret (1935b: 304). Type: BRAZIL. Amazonas: Mun. Tef, Paranagua, 22 May 1933, B. Krukoff 4543
(holotype B, destroyed, isotypes F!, M!, MO!, NY!, US!).
Leaves undivided or pinnate; veins raised and rectangular in cross-section adaxially, or not raised or slightly
raised and triangular in cross-section adaxially; pinnae 2(112) per side of rachis. Inflorescences unbranched
or branched; staminate flowers deciduous or persistent after anthesis.
of Venezuela, Colombia, Ecuador, Peru, and Brazil and eastern Andean slopes in Ecuador, at 416(751850) m
elevation in lowland or montane rainforest (Fig. 38).
This is a widespread and extremely variable subspecies which can be divided into various morphotypes,
mostly based on leaf size and shape.
In the western Amazon basin of Colombia, Ecuador, Peru, and Brazil there is a morphotype
(arundinacea) with undivided or pinnate leaves with non-raised adaxial veins and unbranched or branched
inflorescences with deciduous staminate flowers. In some cases specimens approach those of the trailii
morphotype and are only distinguished by their non-raised adaxial veins. For example, in Yasuni National
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HENDERSON
Park in Amazonian Ecuador, there are two forms of this morphotype. One has smaller leaves and branched
inflorescences, the other larger leaves and unbranched inflorescences. The latter exactly resemble trailii in
their leaves, except for the non-raised veins. There are no specimens of trailii from Yasuni, but it occurs just to
the west of the Park. There may be introgression between arundinacea and trailii in this area. A specimen
(Vsquez 7413) from Amazonian Peru has an exceptionally long rachilla.
On eastern Andean foothills and in the western Amazon region in Colombia, Ecuador, and Peru there is a
morphotype (elevata) with small, undivided leaves with narrow basal angles and raised adaxial veins and
unbranched inflorescences with persistent staminate flowers. Inflorescences are usually pendulous.
In the western Amazon region in Colombia, Peru, and Brazil there is a morphotype (minor) with small,
undivided, rarely pinnate leaves and unbranched inflorescences with deciduous staminate flowers. Veins are
difficult to score in this morphotype.
In the western Amazon region in Colombia, Ecuador, Peru, and Brazil there is a morphotype (piscicauda)
with large, undivided leaves with narrow basal angles and raised adaxial veins. Inflorescences are unbranched
and often pendulous, and have persistent staminate flowers. The types of G. piscicauda and G. wittiana are of
this morphotype.
On eastern Andean foothills in Ecuador at 1217(8251600) m elevation there is a morphotype (puyo) with
pinnate leaves and branched or unbranched inflorescences with deciduous staminate flowers. Veins are
difficult to score in this morphotype, and there seem to be several local variants. Some specimens (Balslev
6419, Cern 6552, 7454, Harling 3762, llgaard 98478) have exceptionally long inflorescences.
A widespread morphotype (pycnostachys, Plates XX, XXI) occurs in the central and western Amazon
region in Venezuela, Colombia, Ecuador, Peru, and Brazil. It has mostly undivided leaves with the veins not
raised adaxially, and unbranched inflorescences with persistent staminate flowers. The type of G.
pycnostachys is of this morphotype. Several specimens from the central Amazon region (Campos 519, Cid
545, Henderson 662, 1043, 1066, Nee 42341, 42897) are more similar to subsp. stricta in their small leaves
than they are to other, more westerly specimens of pycnostachys. However, small-leaved pycnostachys also
occur sporadically in the western Amazon region. Specimens (Daz 7327, Kajekai 300, Rodrguez 261, 568,
Rojas 592, Vsquez 18741, 20286, 24195, 24322) from southeastern Ecuador and northwestern Peru
(Amazonas) have exceptionally large leaves, thick rachillae, and large fruits. Specimens from eastern Andean
slopes in Ecuador have pinnate leaves with few divisions. Some specimens (e.g., Lewis 10332, Vsquez
14439, and probably several others) appear to be hybrids between this and the trailii morphotype, and others
appear to be hybrids between this and the piscicauda morphotype.
In the central and western Amazon region of Colombia, Ecuador, Peru, Bolivia, and Brazil, with outliers
in the central Amazon of Brazil and in Bolivia, there is a morphotype (trailii) with pinnate leaves with raised
adaxial veins and unbranched inflorescences with deciduous staminate flowers. The types of Geonoma
elegans var. amazonica, Geonoma trauniana, Geonoma dasystachys, and Geonoma bella are of this
morphotype. The two outlying specimens in Bolivia occur in the same area as outlying specimens of the
tapajotensis morphotype of G. macrostachys. As in the pycnostachys morphotype, specimens from
southeastern Ecuador and northwestern Peru (Amazonas) have exceptionally large inflorescences.
In the southwestern Amazon region in Peru and Brazil there is a morphotype (uleana) with undivided or
pinnate leaves with non-raised adaxial veins and unbranched inflorescences with deciduous staminate
flowers. It has longer peduncular bracts3.5 (2.74.7) versus 0.5(0.14.0) cmthan other morphotypes. The
type of G. uleana is of this morphotype.
60d. Geonoma stricta subsp. bracteata Henderson, subsp. nov. (Appendix IV, Plate 61)
A subspeciebus aliis foliis pinnatis venis haud prominentibus, rachillis tenuibus, atque inflorescentiis ramosis differt.
Type: PERU. Hunuco: Prov. Pachitea, region of Pucallapa, western side of the Sira mountains, 928S, 7447W, 800
m, 10 September 1988, H. Rainer P2210988 (holotype NY!).
145
Leaves pinnate; veins not raised or slightly raised and triangular in cross-section adaxially; pinnae 3(34) per
side of rachis. Inflorescences branched; staminate flowers deciduous after anthesis.
Distribution and habitat:From 9031012S and 74477530W on eastern Andean slopes in
central Peru (Hunuco, Pasco, Ucayali) at 542(320800) m elevation in lowland rainforest (Fig. 39).
FIGURE 39. Distribution maps of Geonoma stricta subsp. bracteata, G. stricta subsp. divaricta, G. stricta subsp.
pendula, and G. stricta subsp. pliniana.
60e. Geonoma stricta subsp. divaricata Henderson, subsp. nov. (Appendix IV, Plate 62)
A subspeciebus aliis foliis pinnatis venis haud prominentibus, rachillis crassis, atque inflorescentiis ramosis differt.
Type: PERU. Amazonas: Distr. El Cenepa, comunidad de Tutino, 433S, 7812W, 500 m, 20 July 1997, R. Rojas, A.
Pea & E. Chvez 101 (holotype NY!, isotype MO n.v.).
Leaves pinnate; veins not raised or slightly raised and triangular in cross-section adaxially; pinnae 3(34) per
side of rachis. Inflorescences branched; staminate flowers deciduous after anthesis.
northern Peru (Amazonas, Loreto) at 510(170950) m elevation in lowland rainforest (Fig. 39).
60f. Geonoma stricta subsp. pendula Henderson, subsp. nov. (Appendix IV, Plate 63)
A subspeciebus aliis foliis plerumque simplicibus venis haud prominentibus atque inflorescentiis ramosis differt.
146
HENDERSON
Type: PERU. Amazonas: Condorcanqui Province, Distrito El Cenepa, Comunidad de Mamayaque, 434S, 7814W,
400 m, 9 August 1997, R. Rojas, A. Pea & E. Chvez 256 (holotype NY!, isotype MO n.v.).
Leaves undivided or pinnate; veins not raised or slightly raised and triangular in cross-section adaxially;
pinnae 1(12) per side of rachis. Inflorescences branched; staminate flowers deciduous after anthesis.
Distribution and habitat:From 429445S and 77587814W in the northwestern Amazon
region of Peru (Amazonas) at 310(250400) m elevation in lowland rainforest (Fig. 39).
60g. Geonoma stricta subsp. pliniana Henderson, subsp. nov. (Appendix IV, Plate 64)
A subspeciebus guianensibus aliis foliis pinnatis venis prominentibus atque inflorescentiis simplicibus differt.
Type: FRENCH GUIANA. D.Z. 5, route Rgina-Saint-Georges, bassin de lApprouague, 402N, 5201W, 100 m, 26
November 1995, J.-J. de Granville & G. Cremers 13148 (holotype NY!, isotype CAY n.v.).
Leaves pinnate; veins raised and rectangular in cross-section adaxially; pinnae 3 per side of rachis.
Inflorescences unbranched; staminate flowers deciduous after anthesis.
Distribution and habitat:From 345517N and 51485303W in the northeastern Amazon
region of French Guiana and Brazil (Amap) at 53(5100) m elevation in coastal, terra firme forests (Fig. 39).
60h. Geonoma stricta subsp. quibdoensis Henderson, subsp. nov. (Appendix IV, Plate 65)
A subspeciebus extraamazonicis aliis bractea pedunculare prophyllo propiore inserta differt.
Type: COLOMBIA. Choc: ca. 1015 km S of Quibd on road to Istmina (Panamerican Highway), and 810 km E on
road to petroleum exploration camp, 535N, 7637W, 90 m, 9 July 1986, M. Grayum, B. Hammel, J. Kress & G.
Brown 7645 (holotype MO!).
Leaves undivided; veins not raised or slightly raised and triangular in cross-section adaxially; pinnae 1 per
side of rachis. Inflorescences unbranched; staminate flowers deciduous after anthesis.
Distribution and habitat:From 535550N and 76357637W on the Pacific coast of Colombia
(Choc) at 96(50150) m elevation in lowland rainforest (Fig. 40).
60i. Geonoma stricta subsp. submontana Henderson, subsp. nov. (Appendix IV, Plate 66)
A subspeciebus aliis foliis pinnatis venis prominentibus atque inflorescentiis ramosis differt.
Type: PERU. Pasco: Prov. Oxapampa, W side of Cordillera de San Matias between Iscosacin and summit, 1011S,
7512W, 680850 m, 21 June 1982, D. Smith 2019 (holotype NY!, isotype MO n.v.).
Leaves undivided or pinnate; veins raised and rectangular in cross-section adaxially; pinnae 1(12) per side of
rachis. Inflorescences branched; staminate flowers deciduous after anthesis.
Distribution and habitat:From 9501018S and 75127515W on eastern Andean foothills in
central Peru (Hunuco, Pasco) at 437(350760) m elevation in lowland rainforest (Fig. 40).
61. Geonoma talamancana Grayum (1998: 324). Type: COSTA RICA. Limn: Cantn de Limn, Cordillera
de Talamanca, N flank of Fila de Matama in headwaters of Ro Boyei, 945N, 8319W, 12001300 m, 17
August 1995, M. Grayum 11033 (holotype MO!, isotypes CR!, INB!, K, NY!).
Plants 1.1(0.52.0) m tall; stems 1.1(1.01.3) m tall, 1.3 cm in diameter, solitary or clustered; internodes
147
length no data, yellowish and smooth, or, if short and congested, not scaly. Leaves 9(710) per stem,
undivided or irregularly pinnate, plicate, bases of blades running diagonally into the rachis; sheaths
18.9(15.023.5) cm long; petioles 27.4(12.041.5) cm long, drying green or yellowish; rachis 31.6(21.057.0)
43) with the rachis; apical pinna 29.3(18.639.0) cm long, 7.2(5.58.6) cm wide, forming an angle of 17(10
25) with the rachis. Inflorescences unbranched; prophylls ribbed with elongate, unbranched fibers, tubular,
narrow, elongate, closely sheathing the peduncle, more or less persistent; prophylls 29.9(16.538.0) cm long,
bracts absent; peduncles 68.5(55.581.0) cm long, 2.3(1.63.2) mm in diameter; rachillae 1, 16.2(11.523.5)
cm long, 3.5(2.54.6) mm in diameter, the surfaces without spiky, fibrous projections or ridges, drying brown
or yellow-brown, without short, transverse ridges, not filiform and not narrowed between the flower pits;
flower pits spirally arranged, glabrous internally; proximal lips apiculate and lobed before anthesis, tearing in
the center after anthesis, not recurved after anthesis, not hood-shaped; proximal and distal lips drying the
anthesis; non-fertilized pistillate flowers persistent after anthesis; staminodial tubes crenulate or shallowly
7.3(6.38.3) mm long, 5.2(4.55.8) mm in diameter, the bases with a prominent, asymmetric stipe, the apices
not conical, the surfaces not splitting at maturity, without fibers emerging, bumpy from the numerous,
Distribution and habitat:From 845947N and 82438341W in Costa Rica and just reaching
Panama at 1781(12502315) m elevation in montane rainforest (Fig. 40).
Taxonomic notes:Geonoma talamancana is a member of a group of high elevation, Andean species
(the G. undata clade, also including G. lehmannii, G. orbignyana, G. trigona, and G. undata). It is very similar
to G. lehmannii subsp. corrugata, differing only in its lack of a peduncular bract.
geographic discontinuity but too few specimens to test for differences. The specimens from the western part
of the range occur at lower elevations (mean of 1460 m versus 2054 m) and have fewer pinnae.
62. Geonoma tenuissima Moore (1982: 204). Type: ECUADOR. Los Rios: across Ro Palenque from Centro
Cientfico, 150220 m, 22 March 1977, F. Essig 350 (holotype USF n.v.).
internodes 1.6(0.82.8) cm long, yellowish and smooth. Leaves undivided, not plicate, bases of blades
running diagonally into the rachis; sheaths 5.2(4.75.5) cm long; petioles 5.3(3.06.5) cm long, drying green
or yellowish; rachis 27.0(22.034.0) cm long, 2.5(1.93.6) mm in diameter; veins not raised or slightly raised
and triangular in cross-section adaxially; pinnae 1 per side of rachis; basal pinna length and width not
applicable, forming an angle of 28(2336) with the rachis; apical pinna 13.5(11.515.8) cm long, width not
applicable, forming an angle of 30(2236) with the rachis. Inflorescences branched 34 orders; prophylls and
peduncular bracts not ribbed with elongate, unbranched fibers, flattened, deciduous; prophylls 4.3(3.45.2)
cm long, short, asymmetrically apiculate, the margins curved around the stem, the surfaces flat with dense,
felty, brown tomentum, prophyll equal to and early deciduous with the peduncular bract, the surfaces not
ridged, without unequally wide ridges; peduncular bracts 4.5 cm long, well-developed, inserted 0.2(0.10.3)
cm above the prophyll; peduncles 3.1(1.64.7) cm long, 2.9(2.33.8) mm in diameter; rachillae 74, 6.8(3.5
148
HENDERSON
brown, with faint to pronounced, short, transverse ridges, filiform with extended narrowed sections between
the flower pits; flower pits alternately arranged (sometimes distorted by twisting and contracting of rachillae),
epidermis without operculum, smooth, without pores.
Distribution and habitat:From 031038S and 79087918W in western Ecuador (Montaas de
Ila) at 597(520700) m elevation in lowland rainforest (Fig. 40).
Taxonomic notes:Geonoma tenuissima is a member of the Geonoma lanata clade. It appears most
similar to G. mooreana and G. scoparia. It differs from the former by its proximal and distal lips which are
joined to form a raised cupule; and from the latter in its yellowish and smooth internodes.
Subspecific variation: No trait varies within species, nor is there any geographic discontinuity.
FIGURE 40. Distribution maps of Geonoma stricta subsp. quibdoensis, G. stricta subsp. submontana, G. talamancana,
and G. tenuissima.
149
63. Geonoma triandra (Burret) Wessels Boer (1968: 85). Kalbreyera triandra Burret (1930a: 143). Type:
COLOMBIA. Antioquia: Murr, 1000 m, 21 July 1880, W. Kalbreyer 1829 (holotype B, destroyed). Neotype
(here designated): COLOMBIA. Antioquia: Mun. Mutat, Hacienda Mocar, carretera Mutat-Pavarando, 180
m, 1 May 1987, R. Fonnegra, F. Roldn, J. Betancur, & A. Betancur 2002 (neotype NY!).
1.3(0.72.4) cm long, yellowish and smooth. Leaves 10(811) per stem, undivided or irregularly pinnate, not
not raised or slightly raised and triangular in cross-section adaxially; pinnae 2(13) per side of rachis; basal
pinna 19.1(14.030.0) cm long, 6.8(3.814.5) cm wide, forming an angle of 36(2057) with the rachis;
apical pinna 13.9(8.518.0) cm long, 10.3(7.012.7) cm wide, forming an angle of 36(2846) with the rachis.
fibers, flattened, persistent; prophylls 13.7(5.527.0) cm long, not short and asymmetrically apiculate, the
surfaces not ridged, without unequally wide ridges; peduncular bracts 1.8(0.16.0) cm long, vestigial, inserted
narrowed between the flower pits; flower pits decussately arranged throughout the rachillae, the groups of pits
closely spaced, glabrous internally; proximal lips without a central notch before anthesis, not recurved after
anthesis, hood-shaped at anthesis, sometimes splitting post-anthesis; proximal and distal lips drying the same
Distribution and habitat:From 048805N and 76007844W in eastern Panama, western
Colombia, and northwestern Ecuador at 233(451250) m elevation in lowland or montane rainforest (Fig. 41).
Taxonomic notes:Geonoma triandra is one of only two species of Geonoma with staminate flowers
with three stamens; G. hollinensis is the other. Lacking staminate flowers, it can be recognized by its vestigial
peduncular bracts and decussately arranged flower pits.
Subspecific variation:One trait (leaf division) varies within this species. There is geographic
discontinuity and specimens come from several disjunct areas. However, there are too few specimens to test
for differences, and the gaps may be artifacts of insufficient collecting.
64. Geonoma triglochin Burret (1930c: 8). Type: COLOMBIA. Caquet: Sucre, 10 July 1926, G. Woronow &
S. Juzepczuk 5858 (holotype LE n.v., holotype image US!).
Plants 2.2(1.54.0) m tall; stems 2.3(0.28.0) m tall, 1.7 cm in diameter, solitary, cane-like; internodes 1.2 cm
long, yellowish and smooth. Leaves 17(1025) per stem, undivided or irregularly pinnate, not plicate, bases of
blades recurved against the rachis; sheaths 18.6(10.027.5) cm long; petioles 9.0(0.035.0) cm long, drying
green or yellowish; rachis 84.7(38.5160.0) cm long, 6.7(2.715.7) mm in diameter; veins not raised or
slightly raised and triangular in cross-section adaxially; pinnae 3(18) per side of rachis; basal pinna
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HENDERSON
flattened (if tubular, narrow, and elongate then not ribbed), deciduous or persistent; prophylls 17.4(7.533.0)
cm long, not short and asymmetrically apiculate, the surfaces not ridged, without unequally wide ridges;
5.1(2.710.6) mm in diameter, the surfaces without spiky, fibrous projections or ridges, drying brown or
pits spirally arranged, glabrous internally; proximal lips with a central notch before anthesis, often the two
sides of the notch overlapping, not recurved after anthesis, not hood-shaped; proximal and distal lips drying
the same color as the rachillae, not joined to form a raised cupule, the proximal lip margins overlapping the
distal lip margins; distal lips well-developed; staminate and pistillate petals not emergent, not valvate
directly onto apiculate filament apices; anthers not short and curled at anthesis, usually elongate, spiraled and
twisted or sometimes remaining straight; non-fertilized pistillate flowers deciduous after anthesis; staminodial
tubes lobed at the apex, the lobes spreading at anthesis, acuminate, those of non-fertilized pistillate flowers
not projecting and persistent after anthesis; fruits 12.9(10.016.0) mm long, 10.4(7.612.7) mm in diameter,
the bases without a prominent stipe, the surfaces not splitting at maturity, without fibers emerging, not bumpy,
not apiculate; locular epidermis with operculum, smooth, with pores.
FIGURE 41. Distribution maps of Geonoma triandra, G. triglochin, G. trigona, and G. umbraculiformis.
151
Distribution and habitat:From 0001235S and 71187838W in the western Amazon region in
Colombia, Ecuador, and Peru at 735(2001500) m elevation in lowland or montane rainforest (Fig. 41).
Taxonomic notes:Geonoma triglochin is a member of the G. macrostachys clade, within which it is
closely related to G. oldemanii and G. umbraculiformis. It differs from the first in its prophylls and peduncular
bracts which are not ribbed with elongate, unbranched fibers; and from the second in its fruit surfaces which
do not split at maturity.
Subspecific variation:No traits except for leaf division vary within this species. There is geographic
discontinuity, and specimens come from three areas, one in Ecuador and adjacent Colombia and Peru, and two
in southern Peru. However, there are too few specimens to test for differences. There is an isolated, low
elevation population in Amazonian Ecuador that differs significantly from other populations in nine variables.
However, it is not known if the gap between this population and other, sub-Andean ones is an artifact of
insufficient collecting.
between elevation and nine leaf and four inflorescence variables. Squared multiple R for the regression of
sheath length on elevation is 0.75, rachis length 0.63, rachis width 0.36, basal pinna length 0.70, basal pinna
width 0.82, basal pinna angle 0.22, apical pinna length 0.57, apical pinna width 0.33, apical pinna angle 0.51,
peduncle width 0.18, rachilla length 0.48, fruit length 0.43, and fruit diameter 0.56. In particular, with
increasing elevation there is a change in leaf shape, with pinnae becoming shorter and narrower with wider
angles. For inflorescences, peduncles become narrower, rachillae shorter, and fruits smaller with increasing
elevation.
65. Geonoma trigona (Ruz & Pavn) Gentry (1986: 161). Carludovica trigona Ruz & Pavn (1798: 293).
Ludovia trigona (Ruz & Pavn) Persoon (1807: 576). Salmia trigona (Ruz & Pavn) Willdenow (1811:
401). Type: PERU. Hunuco: no locality, no date, J. Pavn s.n. (holotype MA, destroyed, isotype FI n.v.).
Plants 2.9(1.54.0) m tall; stems 3.0 m tall, in diameter no data, solitary; internodes no data. Leaves 6 per
stem, undivided, plicate, bases of blades running diagonally into the rachis; sheaths 17.9(15.020.7) cm long;
petioles 7.1(3.09.0) cm long, drying green or yellowish; rachis 18.3(11.732.0) cm long, 6.3(4.97.6) mm, in
diameter; veins raised and rectangular in cross-section adaxially; pinnae 1 per side of rachis; basal pinna
length and width not applicable, forming an angle of 17(1027) with the rachis; apical pinna 22.9(18.026.0)
cm long, width not applicable, forming an angle of 17(1123) with the rachis. Inflorescences branched 12
orders; prophylls and peduncular bracts not ribbed with elongate, unbranched fibers, flattened (if tubular,
narrow, and elongate then not ribbed), deciduous or persistent; prophylls 36.2(27.543.0) cm long, not short
and asymmetrically apiculate, the surfaces not ridged, without unequally wide ridges; peduncular bracts 27.0
cm long, well-developed, inserted 18.7 cm above the prophyll; peduncles 46.4(38.061.0) cm long, 7.4(6.4
proximal lips apiculate and lobed before anthesis, tearing in the center after anthesis, not recurved after
raised cupule, the proximal lip margins overlapping the distal lip margins; distal lips absent; staminate and
not projecting and persistent after anthesis; fruits 9.0 mm long, 6.5 mm in diameter, the bases with a
prominent, asymmetric stipe, the apices not conical, the surfaces not splitting at maturity, without fibers
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HENDERSON
Ecuador and Peru at 2698(20903000) m elevation in pajonal or dwarf forest (Fig. 41).
Taxonomic notes:Geonoma trigona is a member of a group of high elevation, Andean species (the G.
undata clade, also including G. lehmannii, G. orbignyana, G. talamancana, and G. undata). It is distinguished
by its absent distal lips of the flower pits. The species is also notable for its strongly plicate leaves. In fact, it
was first placed in the Cyclanthaceae and was only recently transferred to Geonoma (Gentry, 1986).
Subspecific variation:No traits vary within this species. There is geographic discontinuity and
specimens come from two areas on the eastern slopes of the Andessouthern Ecuador, and central Peru.
There are too few specimens to test for differences between these two areas. See under Geonoma undata
subsp. undata for possible hybrids with that species.
66. Geonoma umbraculiformis Wessels Boer (1965: 35). Type: SURINAME. Lawa river, no date, Versteeg
322 (holotype U n.v.).
Plants 1.9(1.82.3) m tall; stems 1.5(0.72.0) m tall, 1.7 cm in diameter, cane-like; internodes 1.2 cm long,
yellowish and smooth. Leaves 16(1222) per stem, undivided or irregularly pinnate, not plicate, bases of
blades recurved against the rachis; sheaths 21.0(20.022.0) cm long; petioles 3.8(0.010.5) cm long, drying
green or yellowish; rachis 75.9(63.090.0) cm long, 5.0(3.17.9) mm in diameter; veins not raised or slightly
raised and triangular in cross-section adaxially; pinnae 3(14) per side of rachis; basal pinna 45.0(40.551.0)
cm long, 27.6(21.039.0) cm wide, forming an angle of 16(922) with the rachis; apical pinna 31.0(15.0
25.3) cm long, 18.3(15.521.5) cm wide, forming an angle of 31(2837) with the rachis. Inflorescences
branched 1 order; prophylls and peduncular bracts not ribbed with elongate, unbranched fibers, flattened (if
tubular, narrow, and elongate then not ribbed), deciduous or persistent; prophylls 13.9(13.015.0) cm long,
bracts 28.8(23.034.5) cm long, well-developed, inserted 4.4(3.05.7) cm above the prophyll; peduncles
35.4(31.040.5) cm long, 3.9(2.55.0) mm in diameter; rachillae 4(36), 17.4(12.023.5) cm long, 3.3(2.7
4.2) mm in diameter, the surfaces without spiky, fibrous projections or ridges, drying brown or yellow-brown,
staminate flowers deciduous after anthesis; stamens 6; thecae diverging at anthesis, inserted directly onto
apiculate filament apices; anthers not short and curled at anthesis, usually elongate, spiraled and twisted or
sometimes remaining straight; non-fertilized pistillate flowers deciduous after anthesis; staminodial tubes
lobed at the apex, the lobes spreading at anthesis, acuminate, those of non-fertilized pistillate flowers not
projecting and persistent after anthesis; fruits 12.6(12.013.0) mm long, 10.3(9.711.0) mm in diameter, the
bases without a prominent stipe, the surfaces not splitting at maturity, without fibers emerging, not bumpy, not
apiculate; locular epidermis with operculum, smooth, with pores.
Distribution and habitat:From 150S529N and 46105823W in the eastern Amazon region in
Guyana, French Guiana, and Brazil at 549(200750) m elevation in lowland rainforest (Fig. 41).
Taxonomic notes:Wessels Boer (1968) included Geonoma umbraculiformis as a synonym under G.
triglochin. The two are closely related but G. umbraculiformis differs from G. triglochin in its fruits surfaces
which do not split at maturity. The two species are also widely separate geographically, with G. triglochin
occurring in the western Amazon region in Colombia, Ecuador, and Peru.
153
Subspecific variation:Only one trait (leaf division) varies within this species. Specimens come from
scattered areas, but the gaps are likely to be artifacts of insufficient collecting.
67. Geonoma undata Klotzsch (1847: 452). Type: VENEZUELA. Miranda: Tovar, no date, H. Karsten 26
(holotype not known; isotype BM!).
undivided or irregularly pinnate, not plicate or plicate, bases of blades running diagonally into the rachis;
sheaths 39.4(5.097.5) cm long; petioles 30.7(0.0113.0) cm long, drying green or yellowish; rachis
adaxially or not raised or slightly raised and triangular in cross-section adaxially; pinnae 19(165) per side of
rachis; basal pinna 43.3(14.083.0) cm long, 3.2(0.327.0) cm wide, forming an angle of 48(1090) with the
rachis; apical pinna 32.7(8.066.0) cm long, 9.6(0.130.0) cm wide, forming an angle of 25(541) with the
unbranched fibers, flattened (if tubular, narrow, and elongate then not ribbed), deciduous or persistent;
prophylls 27.8(5.449.0) cm long, prophylls not short and asymmetrically apiculate, the surfaces ridged and
densely tomentose with widely to closely spaced ridges, the ridges unequally wide, often dividing from and
rejoining other ridges, the prophyll margins with irregular, spine-like projections (rarely these absent), the
prophylls usually splitting irregularly between the ridges; peduncular bracts 18.7(7.039.0) cm long, welldeveloped, inserted 2.9(0.411.0) cm long; peduncles 18.4(4.750.0) cm long, 10.5(1.534.4) mm in
diameter; rachillae 21(380), 19.7(5.054.0) cm long, 3.7(0.89.4) mm in diameter, the surfaces without
filiform and not narrowed between the flower pits; flower pits usually spirally arranged, sometimes
decussately or tricussately, then the groups not closely spaced nor consistently arranged throughout the
rachillae, glabrous internally; proximal lips apiculate and lobed before anthesis, tearing in the center after
pistillate flowers persistent after anthesis; staminodial tubes crenulate or shallowly lobed at the apex, those of
non-fertilized pistillate flowers not projecting and persistent after anthesis; fruits 9.5(4.415.4) mm long,
sculpted, usually also with a raised, meridional ridge, without pores.
Taxonomic notes:Geonoma undata is a member of a group of high elevation, Andean species (the G.
undata clade, also including G. lehmannii, G. orbignyana, G. talamancana, and G. trigona). These species
have been treated differently by both Wessels Boer (1968) and Henderson et al. (1995). They are closely
related and three of themG. lehmannii, G. orbignyana, and G. undata are difficult to distinguish from one
another, and extremely complex internally. In fact, G. undata is the second most variable species of Geonoma
(Table 2). It differs from other species in this group by its prophyll surfaces which are ridged and densely
tomentose with widely to closely spaced ridges, the ridges unequally wide, often dividing from and rejoining
other ridges.
Subspecific variation:Five traits vary within this species (stem branching, stem type, leaf division, leaf
plication, adaxial veins)(distal lip also varies but may be a result of hybridization, see under G. undata subsp.
undata). Excluding stem branching and leaf division and the trait for which there are few data (stem type), the
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HENDERSON
state distributions of the remaining two traits (leaf plication, adaxial veins) divide the specimens into three
subgroups. One of these has non-raised adaxial veins and a discrete geographical range, and based on this is
recognized as subspecies (subsp. stenothrysa). The second subgroup has plicate leaves and the third has nonplicate leaves. However, leaf plication is difficult to score in this species and does not divide the specimens
into consistent subgroups. These specimens are therefore examined on a geographical basis. There are several
geographically isolated subgroups, but usually too few specimens in each subgroup to test for differences.
These subgroups are recognized as subspecies.
There is an isolated subgroup in the Lesser Antilles (Dominica, Guadeloupe, and Martinique). There are
only four specimens, but given their geographic isolation they are recognized as a subspecies (subsp.
dussiana).
There is an isolated subgroup in the Guayana Highland region of Venezuela and adjacent Brazil and
Guyana. It differs from its nearest neighbors in Andean Venezuela in seven variables (stem height, number of
pinnae, apical pinna width, apical pinna angle, prophyll length, rachilla length, rachilla width) (t-test, P
<0.05). It is recognized as a subspecies (subsp. appuniana).
There is an isolated subgroup from the Tumuc-Humac mountains in French Guiana and Suriname. There
are only two specimens, but the flower pits tend to be decussately arranged and there are few pinnae (34)
compared with other specimens and they occur at lower elevations (600620 m). These are recognized as a
subspecies (subsp. tumucensis).
There is an isolated subgroup from Central America in Mexico, Guatemala, Honduras, Nicaragua, Costa
Rica, and western Panama. It differs from the remaining specimens in seven variables (plant height, basal
pinna width, apical pinna width, apical pinna angle, peduncle width, rachilla length, rachilla width) (t-test, P
<0.05). It is recognized as a subspecies (subsp. edulis).
There is an isolated subgroup from Cerro Tacarcuna in Panama. There are only two specimens but they
have distinctive, distantly spaced flower pits and are recognized as a subspecies (subsp. tacarcunensis).
There is an isolated subgroup from Andean Venezuela in Carabobo state. The two specimens have
distinctive, pinnate leaves with narrow, linear pinnae and are recognized as a subspecies (subsp. venezuelana).
There is a subgroup from eastern Andean slopes in Ecuador that have distinctive, narrow, linear pinnae
and are reported to be rheophytes. These are recognized as a subspecies (subsp. pulcherrima).
There is a subgroup from Andean Ecuador which has distinctive, narrow rachillae and distantly spaced
flower pits. It differs from other Ecuadorian specimens in 17 variables (plant height, stem height, internode
length, leaf number, rachis length, rachis width, number of pinnae, basal pinna length, basal pinna width, basal
pinna angle, apical pinna length, prophyll length, peduncular bract length, interbract distance, peduncle width,
rachilla length, rachilla width)(t-test, P <0.05) and is recognized as a subspecies (subsp. skovii).
The remaining specimens, from Andean regions of South America in Venezuela, Colombia, Ecuador,
Peru, and Bolivia, are extremely variable and are not divisible into subspecies and are recognized as a single
subspecies (subsp. undata).
Key to the subspecies of G. undata
1
2
3
4
-
Veins not raised or slightly raised and triangular in cross-section adaxially; Central Cordillera in Colombia (Antioquia) ................................................................................................................................................. subsp. stenothrysa
Veins raised and rectangular in cross-section adaxially; widespread .......................................................................... 2
Prophyll margins without irregular, spine-like projections; basal pinna 0.7(0.31.0) cm wide; apical pinna 0.8(0.5
1.0) cm wide; rheophytes on eastern Andean slopes in southeastern Ecuador ............................... subsp. pulcherrima
Prophyll margins with irregular, spine-like projections; basal pinna 3.3(0.427.0) cm wide; apical pinna 10.1(1.5
30.0) cm wide; non-rheophytes, widespread ............................................................................................................... 3
Lesser Antilles .......................................................................................................................................subsp. dussiana
All other areas............................................................................................................................................................... 4
Flower pits decussately arranged; TumucHumac mountains in French Guiana and Suriname ...... subsp. tumucensis
Flower pits usually spirally arranged, sometimes decussately or tricussately, then the groups not closely spaced nor
consistently arranged throughout the rachillae; all other areas .................................................................................... 5
155
5
6
7
8
9.
-
Guayana Highland region of Venezuela, Brazil, and Guyana ............................................................subsp. appuniana

All other areas............................................................................................................................................................... 6
Central America (Mexico, Guatemala, Honduras, Nicaragua, Costa Rica, western Panama) .................. subsp. edulis
South America in Andean regions from Venezuela to Bolivia, and just reaching eastern Panama............................. 7
Flower pits distantly spaced; eastern Panama ...............................................................................subsp. tacarcunensis
Flower pits rarely distantly spaced; South America in Andean regions from Venezuela to Bolivia ........................... 8
Basal pinna 0.5(0.40.5) cm wide; apical pinna 1.7 cm wide; Carabobo state, Venezuela............. subsp. venezuelana
Basal pinna 3.0(0.514.0) cm wide; apical pinna 10.6(2.030.0) cm wide; widespread ............................................. 9
Flower pits distantly spaced; eastern Andean slopes in southern Ecuador ................................................subsp. skovii
Flower pits not distantly spaced; widespread ...........................................................................................subsp. undata
67a. Geonoma undata subsp. undata

Geonoma densa Linden & Wendland (1856: 333). Type: COLOMBIA. Santander: Cachiri, no date, N. Funck & L.
Schlim s.n. (holotype BR n.v., isotype K!), synon. nov.
Geonoma margaritoides Engel (1865: 682). Type: VENEZUELA. Tchira: La Grita, no date, F. Engel s.n. (holotype B,
Geonoma barthia Engel (1865: 688). Type: VENEZUELA. Mrida: no locality, no date, F. Engel s.n. (holotype B
Geonoma iodolepis Burret (1930a: 198). Type: COLOMBIA. Antioquia: Amalfi, 18002000 m, 14 May 1880, W.
Kalbreyer 1668 (holotype B, destroyed). Neotype (designated by Bernal et al. 1989): COLOMBIA. Valle del Cauca:
La Cumbre, Western Cordillera, 17002100 m, 9 September 1922, E. Killip & T. Hanzen 11137 (neotype NY!,
isoneotype US!).
Geonoma weberbaueri Dammer ex Burret (1930a: 221). Type: PERU. Hunuco: SW of Manzn, no date, A. Weberbauer
3552 (holotype B n.v., holotype image!), synon. nov.
Geonoma megalospatha Burret (1930a: 218). Type: PERU. Hunuco: Ro Pozuzo, 19091914, A. Weberbauer 6800
(holotype B n.v., holotype image!, isotypes F!, GH!).
Leaves veins raised and rectangular in cross-section adaxially; basal pinna 3.0(0.514.0) cm wide; apical
pinna 10.6(2.030.0) cm wide. Inflorescences prophyll margins with irregular, spine-like projections; flower
pits usually spirally arranged, not distantly spaced.
Distribution and habitat:From 1056N1748S and 63288042W in Andean regions of South
America from Venezuela to Bolivia at 1964(5503370) m elevation in lowland or, more often, montane
There is considerable variation in this widespread subspecies and there are many local morphotypes.
These are difficult to distinguish from one another and are variable within themselves.
In Venezuela, Colombia, Ecuador, Peru, and Bolivia there is a widespread morphotype (margaritoides
morphotype). It has 25.4(6.454.0) cm long and 4.3(2.27.1) mm wide rachillae and 8.5(5.512.5) mm long
and 6.3(4.28.9) mm wide fruits. This morphotype is relatively uniform throughout the northern part of its
range, usually with elongate rachillae, often tricussately arranged flower pits, and small fruits (e.g., types of G.
margaritoides and G. barthia). However, there are many exceptions, notably specimens with unusually short
and thin, or unusually short and thick rachillae that occur in scattered localities.
There is geographical variation in this morphotype. Linear regression shows there are significant
associations between elevation and one plant, two leaf, and five inflorescence variables. Squared multiple R
for the regression of plant height on elevation is 0.07, basal pinna angle 0.19, apical pinna angle 0.11, prophyll
length 0.35, interbract distance 0.22, number of rachillae 0.76, rachilla width 0.05, and fruit diameter 0.19.
With increasing elevation, pinnae have narrower angles. prophylls and interbract distances are longer,
rachillae wider, and fruits larger.
Some specimens from the Coastal Range in Venezuela (undata morphotype) occur at higher elevations
and have greater interbract distances, longer peduncles, shorter rachillae, and longer and wider fruits. The
type of G. undata has this kind of inflorescence.
Some specimens from Venezuela (Andes and Peninsula Paraguan, Falcn) and Colombia are distinctive
in their shorter, thinner rachillae (densa morphotype). The type of G. densa has this kind of rachillae.
156
HENDERSON
FIGURE 42. Distribution maps of Geonoma undata subsp. undata, G. undata subsp. appunniana, G. undata subsp.
dussiana, and G. undata subsp. edulis.
Some specimens lack distal lips of the flower pits (megalospatha morphotype). These are hypothesized to
be hybrids between the margaritoides morphotype and Geonoma trigona. They share with G. trigona the
absence of an upper lip, and occur in two areas where G. trigona occurs (Ecuador, Peru). It is predicted that G.
trigona will be found in Bolivia at the third hybrid locality. The type of G. megalospatha has this kind of
flower pits.
Specimens from the Western Cordillera of the Colombian Andes (iodolepis morphotype) have small
leaves like those of G. orbignyana and inflorescences with short, thin rachillae 9.0(7.011.0) cm long and
2.6(2.13.5) mm wide. Fruits are 8.5 mm long and 6.3 mm wide. The type of G. iodolepis is of this
morphotype.
In the Venezuelan, Colombian, Ecuadorian, and Peruvian Andes, and also the Sierra Nevada de Santa
Marta in Colombia, there is an extremely variable morphotype (weberbaueri morphotype). Specimens have
plicate leaves, short, thick rachillae 19.0(9.033.0) cm long and 5.7(3.09.4) mm wide, and 12.5(8.015.4)
mm long and 8.7(5.012.0) mm wide fruits. Most specimens with fruits have large-sized fruits, but a few
specimens (Dodson 15213, Vsquez 26597) have small, globose fruits. This morphotype differs from the
margaritoides morphotype in 10 variables. See under Geonoma orbignyana subsp. orbignyana for potential
hybrids with that subspecies.
There is geographical variation in this morphotype. Linear regression shows there are significant
associations between elevation and three inflorescence variables. Squared multiple R for the regression of
157
prophyll length 0.39, interbract distance 0.26, and peduncle length 0.33. Prophylls, interbract distances, and
peduncles increase in length with increasing elevation.
On eastern Andean slopes in Ecuador, particularly from Mera and Puyo, specimens (mera-puyo
morphotype) have relatively slender inflorescences, 14.9(9.821.0) cm long and 2.1(1.52.5) mm wide
rachillae, and 6.0(5.96.0) mm long and 4.6(4.15.0) mm wide fruits.
Also on eastern Andean slopes in Ecuador, there are specimens (intermediate morphotype) which appear
intermediate between the margaritoides morphotype and Geonoma orbignyana. They have small leaves and
inflorescences, like the latter, but the bracts of the inflorescence are like those of the former. They have
16.8(8.829.3) cm long and 3.8(3.15.0) mm wide rachillae and 8.2(7.68.8) mm long and 6.1(5.86.3) mm
wide fruits.
One specimen (sira morphotype) from Peru (Hunuco) has small leaves like those of Geonoma
orbignyana, slender rachillae 13.5 cm long and 1.9 mm wide, almost decussately arranged flower pits, and 8.9
mm long and 6.3 mm wide fruits.
67b. Geonoma undata subsp. appuniana (Spruce) Henderson, comb. & stat. nov
Basionym: Geonoma appuniana Spruce (1871: 106). Type: GUYANA. Cuyuni-Mazaruni: Mount Roraima, 1864, C.
Appun 1141 (holotype K!).
Geonoma roraimae Dammer (1915: 261). Type: GUYANA. Cuyuni-Mazaruni: Mount Roraima, 18002300 m, February
1910, E. Ule 8805 (holotype B, destroyed, isotypes K!, L!, US!).
Distribution and habitat:From 046604N and 59506604W in the Guayana Highland region
of Venezuela, Brazil, and Guyana at 1752(8102700) m elevation in montane rainforest (Fig. 42).
There is geographic discontinuity and specimens occur on many, isolated mountains, but there are too few
specimens from each mountain to test for differences amongst them.
There is geographical variation in this subspecies. Linear regression shows there are significant
associations between elevation and three leaf and four inflorescence variables. Squared multiple R for the
regression of rachis width on elevation is 0.40, basal pinna angle 0.29, apical pinna angle 0.58, prophyll
length 0.23, rachilla width 0.23, fruit length 0.49, and fruit diameter 0.45. The rachis becomes thicker, pinna
angles narrower, prophylls longer, rachillae thicker, and fruits longer and wider with increasing elevation.
67c. Geonoma undata subsp. dussiana (Becc.) Henderson, comb. & stat. nov.
Basionym: Geonoma dussiana Beccari (1920: 436). Lectotype (designated by Read 1979): GUADELOUPE. Bois du
Nez-Cass, 28 February 1904, A. Duss 4198 (lectotype US!, duplicates F!, FI!, LE!, MO!, NY!).
Geonoma hodgeorum Bailey in Hodge (1942: 108). Type: DOMINICA. Morne Trois Pitons, 7631400 m, 23 February
1940, W. Hodge 1430 (holotype BH!, isotype NY!).
Leaves veins raised and rectangular in cross-section adaxially; basal and apical pinna width no data.
Inflorescences prophyll margins with irregular, spine-like projections; flower pits usually spirally arranged,
not distantly spaced.
Distribution and habitat:From 14401605N and 61006140W in Dominica, Guadeloupe and
Martinique at medium elevations in lowland or montane rainforest (Fig. 42).
67d. Geonoma undata subsp. edulis (Wendland ex Spruce) Henderson, comb. & stat. nov.
Basionym: Geonoma edulis Wendland ex Spruce (1871: 106). Type: COSTA RICA. Cartago: Turrialba, 1857, H.
Geonoma seleri Burret (1930a: 211). Type: GUATEMALA. Huehuetenango: Yalambohoch, no date, E. Seler 2757
(holotype B, destroyed). Neotype (designated by de Nevers & Grayum 1998): GUATEMALA. Alta Vera Paz:
between Sepacuite and Panzas, 24 June 1904, O. Cook & Doyle 327 (neotype US!).
158
HENDERSON
Geonoma polyneura Burret (1932b: 500). Type: GUATEMALA. Alta Vera Paz: near Finca Sepacuite, 19 March 1902, O.
Cook & R. Griggs 36 (holotype US!, isotype BH!).
Distribution and habitat:From 7181619N and 80069315W in Mexico, Guatemala,
Honduras, Nicaragua, Costa Rica, and Panama at 1523(8502400) m elevation in lowland or montane
There is geographic discontinuity and specimens occur in two areas separated by the lowlands of southern
Nicaragua. There are few significant differences in variables between specimens from these two areas (basal
pinna length, basal pinna width, prophyll length). There is variation in pit arrangement throughout the range
of the subspecies, from spirally arranged to irregularly decussate or tricussate. A few specimens have the pits
loosely spiraled proximally and tricussate or decussate distally, or tricussate proximally and decussate distally.
One specimen (Evans 1459) from Honduras is larger in inflorescence size than other specimens from that
area and has a narrow, elongate prophyll, more like that of subsp. hoffmanniana.
The range of this subspecies overlaps with that of subsp. hoffmanniana in several placesnorthern
Nicaragua, Atlantic and Pacific slopes of the Central Cordillera in Costa Rica, and both slopes on the
Cordillera de Talamanca in Costa Rica and Panama. In this last area specimens are much larger in size than
those from other areas (as are most specimens of subsp. hoffmanniana, which see). Hammel (2003)
considered that specimens of subsp. edulis (as G. edulis) and larger, sympatric specimens of subsp.
hoffmanniana (as G. hoffmanniana) were virtually indistinguishable in this area.
All specimens from the Cordillera de Talamanca and all others from Panama have much thicker peduncles
than other specimens.
67e. Geonoma undata subsp. pulcherrima (Burret) Henderson, comb. & stat. nov.
Basionym: Geonoma pulcherrima Burret (1930a: 195). Type: ECUADOR. Morona-Santiago: confluence of Ro
Bomboiza and Ro Zamora, 700800 m, no date, F. Lehmann 5289 (holotype B, destroyed, isotypes F!, GH!, K!).
Leaves veins raised and rectangular in cross-section adaxially; basal pinna 0.7(0.31.0) cm wide; apical pinna
0.8(0.51.0) cm wide. Inflorescences prophyll margins without irregular, spine-like projections; flower pits
usually spirally arranged, not distantly spaced.
Distribution:From 330418S and 78307841W on eastern Andean slopes in southeastern
Ecuador at 923(7501130) m elevation along river banks in areas subject to flooding (Fig. 43). Plants are
reported to be rheophytes.
67f. Geonoma undata subsp. skovii (Henderson, Borchsenius, & Balslev) Henderson, comb. & stat. nov.
Basionym: Geonoma skovii Henderson, Borchsenius & Balslev (2008: 60). Type: ECUADOR. Morona-Santiago: along
the Limn-Cuenca road 4 km above Plan de Milagro, 304S, 7830W, 1920 m, 19 May 1988, B. Bergmann & H.
Pedersen 62589 (holotype NY!, isotypes AAU!, QCA n.v.).
7.3(2.711.3) cm wide. Inflorescences prophyll margins with irregular, spine-like projections; flower pits
usually spirally arranged, distantly spaced.
Distribution and habitat:From 240338S and 78007830W in Ecuador on eastern Andean
slopes at 1778(14701920) m elevation in montane rainforest (Fig. 43).
Subspecific variation:There is geographic discontinuity, and specimens come from three different
mountain areas. Specimens from the Cordilleras Cutuc and Cndor have more pinnae per side of the rachis
than those from the eastern slopes of the main Cordillera. However, there are too few specimens to test for
differences. Jean-Christophe Pintaud (pers. comm.) reports that this subspecies also occurs in adjacent Peru.
159
FIGURE 43. Distribution maps of Geonoma undata subsp. pulcherrima, G. undata subsp. skovii, G. undata subsp.
stenothyrsa, and G. undata subsp. tacarcunensis.
67g. Geonoma undata subsp. stenothyrsa (Burret) Henderson, comb. & stat. nov.
Basionym: Geonoma stenothyrsa Burret (1930a: 197). Type: COLOMBIA. Antioquia: San Carlos, 16502650 m, 28
January 1880, W. Kalbreyer 1372 (holotype B, destroyed). Neotype (designated by Bernal et al. 1989):
COLOMBIA. Antioquia: carretera Granada-San Luis, 5.5 km adelante de El Choc, 1750 m, 2021 September
1987, R. Bernal & L. Tobn 1387 (neotype COL!, isoneotypes AAU!, BH!, NY!).
Geonoma euterpoidea Burret (1930a: 196). Type: COLOMBIA. Antioquia: Alto Guatap, 1950 m, 26 February 1880, W.
Kalbreyer 1477 (holotype B, destroyed). Neotype (designated by Bernal et al. 1989): COLOMBIA. Antioquia: Mun.
Guatap, vereda Santa Rita, ca. 1900 m, 20 May 1980, R. Bernal & G. Galeano 190 (neotype COL!, isoneotype
NY!).
4.8(0.18.5) cm wide. Inflorescences prophyll margins with irregular, spine-like projections; flower pits
usually spirally arranged, not distantly spaced.
Colombia (Antioquia) at 1675(15001900) m in montane rainforest (Fig. 43).
One specimen (Bernal 190, neotype of G. euterpoidea) has leaves with more pinnae than the others (23
versus 813).
160
HENDERSON
67h. Geonoma undata subsp. tacarcunensis Henderson, subsp. nov. (Appendix IV, Plate 67)
A subspeciebus aliis foveis remote sitis differt.
Type: PANAMA. Darin: Parque Nacional del Darin, Panama/Colombia border, near gold mine at head waters of N
branch of Ro Pucuro, slopes of Cerro Tacarcuna, ca. 6 km N of Cerro Mali, 809N 7715W, 13001500 m, 27
October 1987, G. de Nevers, B. Hammel & H. Herrera 8511 (holotype NY!, isotype MO!).
no data. Inflorescences prophyll margins with irregular, spine-like projections; flower pits usually spirally
arranged, distantly spaced.
Distribution and habitat:At 809N and 7715W on Cerro Tacarcuna on the Panama-Colombia
border at 1612(14001825) m elevation in montane rainforest (Fig. 43).
67i. Geonoma undata subsp. tumucensis Henderson, subsp. nov. (Appendix IV, Plate 68)
A subspeciebus aliis foveis decussatis differt.
Type: SURINAME. Inselberg Talouakem, Massif des Tumuc-Humac, 229N, 5445W, 620 m, 16 August 1993, J.-J. de
Granville, P. Acevedo, A. Boyer & L. Hollenberg 12322 (holotype NY!, isotype US!).
Leaves veins raised and rectangular in cross-section adaxially; basal pinna 3.0 cm wide; apical pinna no data.
Inflorescences prophyll margins with irregular, spine-like projections; flower pits decussately arranged, not
closely spaced.
Distribution and habitat:From 215229N and 54255445W on the Tumuc-Humac mountains
in French Guiana and Suriname at 610(600620) m in lowland rainforest (Fig. 44).
FIGURE 44. Distribution maps of Geonoma undata subsp. tumucensis, G. undata subsp. venezuelana and G. venosa.
67j. Geonoma undata subsp. venezuelana Henderson, subsp. nov. (Appendix IV, Plates 6971)
A subspeciebus aliis pinnis basalibus atque apicalibus angustis differt.
Type: VENEZUELA. Carabobo: Mpio. Bejuma, parte superior de la Fila La Mesa, Valle de Chirgua, finca Monte Sacro,
12501300 m, 28 November1 December 1996, F. Stauffer, A. Fernndez, R. Riina & K. WaltherWeissbeck 262
(holotype VEN n.v., isotype NY!).
161
1.7 cm wide. Inflorescences prophyll margins with irregular, spine-like projections; flower pits usually
spirally arranged, not distantly spaced.
Distribution and habitat:From 10121015N and 68076810W on the Coastal Range in
Venezuela at 1362(12751450) m elevation in montane rainforest (Fig. 44).
68. Geonoma venosa Henderson, sp. nov. (Appendix IV, Plate 72)
A speciebus affinibus venis prominentibus atque rachillis tenuibus differt.
Type: ECUADOR. Imbabura: Cotacachi, Parroquia Garca Moreno, Reserva Biolgica Los Cedros, 019N, 7846W,
1470 m, 25 October 2005, H. Vargas, W. Defas & D. Reyes 6282 (holotype NY!, isotype MO n.v.).
Plants 2.5(2.03.0) m tall; stems no data; internodes no data. Leaves undivided or irregularly pinnate, not
plicate, bases of blades running diagonally into the rachis; sheaths no data; petioles 13.5 cm long, drying
green or yellowish; rachis 27.5 cm long, 2.8(2.03.6) mm in diameter; veins raised and rectangular in crosssection adaxially; pinnae 1? per side of rachis; basal pinna length no data, 14.7 cm wide, forming an angle of
38(3540) with the rachis; apical pinna 24.5 cm long, 12.3 cm wide, forming an angle of 30(2930) with the
rachis. Inflorescences branched at least 3 orders; prophylls and peduncular bracts no data; peduncles no data;
rachillae 8.6(7.010.2) cm long, 0.9(0.80.9) mm in diameter, the surfaces without spiky, fibrous projections
or ridges, drying brown, with faint to pronounced, short, transverse ridges, filiform with extended narrowed
sections between the flower pits; flower pits alternately arranged (sometimes distorted by twisting and
contracting of rachillae), sometimes decussately, then the groups not consistently arranged throughout the
rachillae, glabrous internally; proximal lips without a central notch before anthesis, not recurved after
anthesis, not hood-shaped; proximal and distal lips drying the same color as the rachillae, joined to form a
raised cupule, the margins not overlapping; distal lips well-developed; staminate and pistillate petals not
anthers short and curled over at anthesis; non-fertilized pistillate flowers deciduous after anthesis; staminodial
persistent after anthesis; fruits 8.1 mm long, 7.2 mm in diameter, the bases without a prominent stipe, the
Distribution and habitat:From 018019N and 78467847W on western Andean slopes in
Ecuador (Reserva Biolgica Los Cedros) at 1465(14601470) m elevation in montane rainforest (Fig. 44).
Taxonomic notes:Geonoma venosa is a member of a group of closely related species, the G. lanata
clade. It shares all character states with G. tenuissima, but the only two specimens known are missing
prophylls and peduncular bracts, so that five characters cannot be scored. Given its larger fruits (8.1 mm long
and 7.2 mm in diameter versus 5.3(4.66.0) cm long and 4.8(4.15.5) mm in diameter), veins raised and
rectangular in cross-section adaxially (versus not raised), and higher elevation habitat (1465(14601470) m
versus 597(520700) m elevation) it is kept separate from G. tenuissima, pending more material.
Subspecific variation:No traits vary within species, nor is there any geographic discontinuity.
162
HENDERSON
Acknowledgements
I thank the many people who have helped with this work. At the New York Botanical Garden, the staff of the
herbarium processed the numerous loans I requested, particularly Dr. Tom Zanoni, Lucy Klebieko, and
Wilson Ramos. Dr. Holly Porter Morgan and Hannah Stevens of the GIS Laboratory provided expertise on
map making; Nestor Prez-Molire of the Herbarium Digitization Laboratory provided the type images; YaYi Huang helped with cladistic analysis; and Dr. Scott Mori helped with photography. The curators of the
following herbaria sent these loans, or allowed me to study specimens in their care: AAU, BH, BM, BR, C,
COAH, COL, CR, F, FTG, G, GH, HUA, INB, INPA, K, LE, M, MBML, MEXU, MO, NY, P, PMA, RB, SP,
SPF, U, US, and USM. Drs. Rodrigo Bernal and Gloria Galeano at COL helped in many ways, and Dr. Bernal
wrote the Latin diagnoses; Paula Leitman helped at RB, Hlio de Queiroz Boudet Fernandes at MBML, and
Drs. Jos Pirani and Renata Pardini at SPF. Drs. Anders Barfod, Henrik Balslev, and Finn Borchsenius invited
me to Aarhus University as a Visiting Professor in the summer of 2006. This enabled me to study the rich
collections of Geonoma in the Aarhus Herbarium, and to benefit from their knowledge of Geonoma. Dr.
Borchsenius and Dr. William Baker reviewed the manuscript. A grant from the International Palm Society
allowed me to study specimens in MBML, RB, SP, SPF.
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Appendix I. Qualitative VariablesCharacters and Traits

Characters
Abbreviations in parentheses at the end of each character are the column labels in the Data Matrix
(http://sciweb.nybg.org/Science2/res/Henderson/Geonoma.xls.zip). The states of the characters here are scored as (1)
or (2) etc., and these correspond with the states in the Data Matrix. However, the same character states in the Phylogeny
Data Matrix (Table 1) are scored as one number less, i.e., 0, 1 etc.
0. Internodes yellowish and smooth, or, if short and congested, not scaly (1); internodes covered with reddish or
brownish scales, especially in their distal part (2); internodes covered with dense, brown scales (3). (intercolor)
1. Bases of leaf blades running diagonally into the rachis (1); bases of leaf blades recurved against the rachis (2).
(bladebase)
2. Leaf blades regularly pinnate, the pinnae with 1 main vein and 2 lateral veins on either side of main vein (1); leaf
blades undivided or irregularly pinnate, if regularly pinnate the pinnae with 1 main vein only (rarely with several
lateral veins) (2). (venation)
3. Prophylls and peduncular bracts ribbed with elongate, unbranched fibers, both bracts tubular, narrow, elongate, closely
sheathing the peduncle, more or less persistent (1); prophylls and peduncular bracts not ribbed with elongate,
unbranched fibers, flattened (if tubular, narrow, and elongate then not ribbed), deciduous or persistent (2). (bracts)
4. Prophylls short, asymmetrically apiculate, the margins curved around the stem, the surfaces flat with dense, felty,
brown tomentum, prophyll equal to and early deciduous with the peduncular bract (1); prophylls not short and
asymmetrically apiculate (2). (prophylls)
5. Prophyll surfaces ridged with close, equal, parallel, non-dividing ridges, scarcely tomentose between the ridges (1);
prophyll surfaces not ridged, or if ridged then densely tomentose with widely to closely spaced ridges, these
sometimes dividing (2). This character is most obvious on the abaxial surface of the prophyll. (prosurface)
6. Prophyll surfaces ridged, the ridges unequally wide, often dividing from and rejoining other ridges, the prophyll
between the ridges (1); prophyll surfaces without unequally wide ridges (2). This character is most obvious after the
rachillae have emerged from the bracts, and splitting is observed later in development. (ridged)
7. Peduncular bracts well-developed (1); peduncular bracts vestigial, the prophyll three times or more long, sometimes
the peduncular bract apparently well-developed but then soon disintegrating (2); peduncular bracts absent (3).
(peduncbract)
8. Rachillae surfaces with spiky, fibrous projections or ridges (1); rachillae surfaces without spiky, fibrous projections or
ridges (2). (rachsurface)
9. Rachillae drying brown or yellow-brown, the surfaces without short, transverse ridges (1); rachillae drying brown, the
surfaces with faint to pronounced, short, transverse ridges (2). (rachrugos)
10. Flower pits tricussately or quadricussately arranged throughout the rachillae, the groups of pits closely spaced (1);
flower pits usually spirally arranged, sometimes decussately or tricussately, then the groups not closely spaced nor
consistently arranged throughout the rachillae (2); flower pits decussately arranged throughout the rachillae, the
groups of pits closely spaced (3); flower pits alternately arranged (sometimes distorted by twisting and contracting of
rachillae)(4). (pitarrang)
11. Flower pits densely hairy internally distally only (rarely some hairs on lateral margins of the pit) (1); flower pits
glabrous internally (2); flower pits densely hairy internally proximally and distally (3). (pitsinside)
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12. Proximal lips of flower pits with a central notch before anthesis, often the two sides of the notch overlapping, the lips
more or less heart-shaped (1); proximal lips of flower pits without a central notch before anthesis (but often tearing
in the center after anthesis), not heart-shaped (2); proximal lips of flower pits apiculate and lobed before anthesis,
tearing in the center after anthesis (3). (proxlip)
13. Proximal lips of flower pits not recurved at the apices after anthesis (1); proximal lips of flower pits recurved at the
apices after anthesis (2). (proxrecurv)
14. Proximal lips hood-shaped at anthesis (the margin of the proximal lip straight when viewed from above), sometimes
splitting post-anthesis (1); proximal lips not hood-shaped at anthesis (2). (hood)
15. Proximal and distal lips not joined laterally, with a clear gap between them, not forming a raised cupule, the proximal
lip margins usually overlapping the distal lip margins (1); proximal and distal lips joined laterally with no clear gap
between them, often forming a raised cupule, the margins not overlapping (2). (lipscupul)
16. Distal lips of flower pits absent (1); distal lips of flower pits well-developed (2); distal lips of flower pits a scarcely
raised rim (3). (distalip)
17. Stamens 3 (1); stamens 6 (2); stamens more than 6 (3). (stamenno)
18. Thecae diverging at anthesis, inserted almost directly onto the filament apices, the connectives bifid but scarcely
developed (1); thecae diverging at anthesis, inserted onto bifid and well-developed, non-jointed connectives (2);
thecae diverging or not diverging at anthesis, inserted onto poorly to well-developed, non-split, jointed connectives,
connectives when well-developed alternately long and short (3); thecae diverging at anthesis, inserted directly onto
the apiculate filament apices (4). (connect)
19. Anthers short and curled over at anthesis (1); anthers not short and curled at anthesis, usually elongate, spiraled and
twisted or sometimes remaining straight (2); anthers short at anthesis, remaining straight and parallel (3). (anthers)
20. Gynoecium trilocular (1); gynoecium unilocular (2) (gyno)
21. Staminodial tubes crenulate or shallowly lobed at the apex (1); staminodial tubes lobed at the apex, the lobes
spreading at anthesis, acuminate (2); staminodial tubes lobed at the apex, the lobes not spreading at anthesis, not
acuminate (3). (stamtubes)
22. Staminodial tubes of non-fertilized pistillate flowers projecting and persistent after anthesis (1); staminodial tubes of
non-fertilized pistillate flowers not projecting, deciduous after anthesis (2). (stamtubper)
23. Fruit bases with a prominent, asymmetric stipe (1); fruit bases without a prominent stipe (2). (stipitate)
24. Fruits ovoid, usually with conical apices (1); fruits not ovoid and without conical apices (2). (conical)
25. Fruit surfaces not splitting at maturity (1); fruit surfaces splitting deeply and longitudinally at maturity to reveal
mesocarp with dense layer of radial fibers (2). (splitting)
26. Fruit surfaces with fibers emerging (1); fruit surfaces without fibers emerging (2). (spiny)
27. Fruit surfaces bumpy from the numerous, subepidermal, tangential, short fibers present, these coming to a point at
fruit apices (1); fruit surfaces not bumpy and not apiculate (2); fruit surfaces ridged from the numerous,
subepidermal, meridional, elongate fibers present, these coming to a point at fruit apices (3). (bumpy)
28. Locular epidermis without operculum (1); locular epidermis with operculum (2). (operc).
29. Locular epidermis without pores, or with very few pores (1); locular epidermis with numerous pores (2). (pores)
169
Traits
Abbreviations in parentheses at the end of each variable are the column labels in the Data Matrix.
0. Stems solitary (1); stems clustered (2). (stembranch)
1. Stems not cane-like, the internodes usually wider than long (ratio of stem diameter to internode length greater than
2.2) (1); stems cane-like, the internodes usually longer than wide (ratio of stem diameter to internode length less than
1.8) (2). (internod)
3. Leaves undivided (1); leaves pinnate (2). (leafdivi)
4. Leaves not plicate (1); leaves plicate (2). (plicate)
5. Petioles (and rachis) drying orange-brown or reddish-brown (1); petioles (and rachis) drying green or yellowish (2).
(petcolor)
6. Veins raised and rectangular in cross-section adaxially (1); veins not raised or slightly raised and triangular in crosssection adaxially (2). (veinadax)
7. Inflorescences unbranched (1); inflorescences branched (2). (inflobran)
8. Rachillae filiform with extended narrowed sections between the flower pits (1); rachillae not filiform and not or
scarcely narrowed between the flower pits (2). (rachnarrow)
9. Proximal and distal lips drying the same color as the rachillae (1); proximal and distal lips drying darker brown than
the rachillae (2). (lipcolor)
10. Staminate and pistillate petals emergent, valvate throughout (1); staminate and pistillate petals not emergent, not
valvate throughout (2). (petals)
11. Staminate flowers persistent post-anthesis (1); staminate flowers deciduous post-anthesis (2). (stamfls)
12. Non-fertilized pistillate flowers persistent after anthesis (1); non-fertilized pistillate flowers deciduous after anthesis
(2). (pistfls)
13. Locular epidermis smooth (1); locular epidermis sculpted, usually also with a raised, meridional ridge (2). (sculpt)
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Appendix II. Quantitative variables

Abbreviations in parentheses at the end of each variable are the column labels in the Data Matrix.
0. Plant height (m); data taken from specimen labels. (plheight)
1. Stem height (m); data taken from specimen labels. (stemheight)
2. Stem diameter (cm); data taken from specimens only, not from labels. (stemdiameter)
3. Internode length (cm); data taken from specimens only, not from labels. (internode)
4. Number of leaves per stem; data taken from specimen labels. (leafnumber)
5. Sheath length (cm); measured from base to apex of sheath. The distinction between sheath and petiole is not clear, and
the apex of the sheath was judged to be the place where no more sheath fibers were present. (sheath)
6. Petiole length (cm); measured from apex of leaf sheath to first pinna. Petioles in Geonoma appear to continue
lengthening after the blade had unfolded. (petiole)
7. Rachis length (cm); measured from first pinna to apex of rachis. (rachislen)
8. Rachis diameter (mm); measured at base of leaf blade. (rachiswid)
9. Number of pinnae per side of rachis. (nodivisions)
10. Basal pinna length (cm); measured at the base of the pinna. (baspinlen)
11. Basal pinna width (cm); measured at the base of the pinna. (baspinwid)
12. Basal angle of pinna divergence (); measured at 7.5 cm distance, between the axis of the rachis and margin of basal
pinna. (baspinang)
13. Apical pinnae length (cm); measured from apex of rachis to apex of pinna. (apinlen)
14. Apical pinna width (cm); measured at the base of the pinna; not measured on simple leaves. (apinwid)
15. Apical angle of pinna divergence (); measured at 7.5 cm distance, between the axis of the rachis and margin of
apical pinna. (apinang)
16. Orders of inflorescence branching. (orders)
17. Prophyll length (cm). (prophyll)
18. Peduncular bract length (cm). (pedbract)
19. Distance between peduncular bract and prophyll insertion (cm). (distance)
20. Peduncle length (cm); measured from prophyll scar to first branch. (pedunclelen)
21. Peduncle diameter (mm); measured just below the peduncular bract scar. (pedunclewid)
22. Number of rachillae. (norachillae)
23. Rachilla length (cm). (rachilllen)
24. Rachilla diameter (mm); measured at middle of rachilla. (rachillwid)
25. Fruit length (mm). (fruitlen)
26. Fruit diameter (mm). (fruitdiam)
171
Appendix III. Excluded Names

This list of name comprises published names that have been included in Geonoma but for which no type specimens are
available, or the type specimens are insufficient for identification, or the type host institution is unknown, or the name
has been transferred to another genus. Many names on this list have been typified with illustrations and were accepted by
Wessels Boer (1968) and others. However, it has not been found possible to identify illustrations, and so they are treated
as excluded names.
Geonoma adscendens Dammer ex Burret, 1930a: 175. Type. Peru. Cusco: Santa Ana, 1700 m, 1 July 1905, A.
Weberbauer 5033 (holotype, B, destroyed).
Geonoma allenii Bailey = Calyptrogyne allenii (Bailey) de Nevers
Geonoma altissima Barbosa Rodrigues, 1898: 6. Lectotype, Wessels Boer, 1968: Barbosa Rodrigues, 1903: t. 12B.
Geonoma amabilis Wendland ex Dahlgren = Pholidostachys pulchra Wendland ex Burret
Geonoma amazonica Wendland in Kerchove de Dentergarten, 1878: 245. Type. Not designated.
Geonoma amoena Burret, 1933a: 862. Type. Colombia. Tolima: locality, no date, F. Lehmann 2266 (holotype, B,
destroyed).
Geonoma andicola Dammer ex Burret, 1930a: 218. Type. Peru. Puno: Chunchusmayo, 2500 m, 27 July 1902, A.
Geonoma andina Burret, 1930a: 188. Type. Peru. Cajamarca: Cutervo, Tambillo, 2450 m, 18 March 1878, A. Raimondi
s. n. (holotype, B, destroyed).
Geonoma anomoclada Burret, 1935c: 615. Type. Colombia. El Valle: Bitaco, 1800 m, 11 September 1934, E. Dryander
29 (holotype, B, destroyed).
Geonoma antioquensis Wendland in Kerchove de Dentergarten, 1878: 245. Type. Not designated.
Geonoma aricanga Barbosa Rodrigues, 1879: 40. Lectotype, Wessels Boer, 1968: Barbosa Rodrigues, 1903: t. 25.
Geonoma barbigera Barbosa Rodrigues, 1882: 45. Lectotype, Wessels Boer, 1968: Barbosa Rodrigues, 1903: t. 26.
Geonoma barbosiana Burret, 1938a. 255. Type. Brazil. Rio de Janeiro: no locality, no date, A. Brade & M. Burret 25
(holotype, B, destroyed).
Geonoma bartlettii Dammer ex Burret, 1930a: 183. Type. Guyana. Conawaruk River, September 1905, A. Bartlett 8195
Geonoma beccariana Barbosa Rodrigues, 1888: 33. Lectotype, Wessels Boer, 1968: Barbosa Rodrigues, 1902: t. 17.
Geonoma bijugata Barbosa Rodrigues, 1875: 10. Lectotype, Wessels Boer, 1968: Barbosa Rodrigues, 1903: t. 16.
Geonoma bluntii auct., in Anon, 1881: 766. Type. Not designated.
Geonoma brachyfoliata Barbosa Rodrigues, 1875: 10. Lectotype, Wessels Boer, 1968: Barbosa Rodrigues, 1903: t. 33.
Geonoma brachystachys Burret, 1940a: 23. Type. Ecuador. Pastaza: Mera, 7 September 1938, H. Schultze-Rhonhof 2788
Geomoma brevispatha Barbosa Rodrigues, 1879: 41. Lectotype, Wessels Boer, 1968: Barbosa Rodrigues, 1903: t. 22.
Geonoma caespitosa Wendland ex Drude, 1882: 500. Type. Cultivated plant, no date, H. Wendland s. n. (holotype, HAN
n.v.).
Geonoma calophyta Barbosa Rodrigues, 1882: 48. Lectotype, Wessels Boer, 1968: Barbosa Rodrigues, 1903: t. 32B.
Geonoma campyloclada Burret, 1930a: 189. Type. Colombia. Department unknown: Arizal, 1700 m, 17 May 1881, W.
Kalbreyer 2045 (holotype, B, destroyed).
Geonoma campylostachys Burret, 1940a: 24. Type. Ecuador. Pastaza: Mera, 7 September 1938, H. Schultze-Rhonhof
Geonoma capanemae Barbosa Rodrigues, 1875: 9. Lectotype, Wessels Boer, 1968: Barbosa Rodrigues, 1903: t. 29.
Geonoma carderi Bull = Prestoea carderi (Bull) Hooker.
Geonoma caudescens Wendland ex Drude, 1882: 504. Type not designated.
Geonoma cernua Burret, 1940a: 24. Type. Ecuador. Pastaza: Mera, 10 October 1938, H. Schultze-Rhonhof 2888
Geonoma chapadensis Barbosa Rodrigues, 1898: 4. Lectotype, Wessels Boer, 1968: Barbosa Rodrigues, 1903: t. 12A,
13A.
Geonoma chiriquensis Linden ex Hooker, 1882: 60. Type. Not designated.
Geonoma compacta Linden, 1881: 31. Type not designated.
Geonoma condensata Bailey = Calyptrogyne condensata (Bailey) Wessels Boer.
Geonoma congestissima Burret, 1930a: 224. Type. Peru. Loreto: Moyobamba, 1300 m, 19 August 1904, A. Weberbauer
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Geonoma corallifera Morren = Chamaedorea ernesti-augusti Wendland
Geonoma decora Linden & Rodigas, 1894: 361. Type not designated.
Geonoma decurrens Wendland ex Burret, 1930a: 162. Type. Costa Rica. Heredia: Sarapiqu, August 1857, H. Wendland
s. n. (holotype, B, destroyed).
Geonoma decussata Burret, 1933b: 103. Type. Brazil. Bahia: near Caetete, no date, E. Werdermann 3454 (holotype, B,
destroyed).
Geonoma demarestei Pritzel, 1855: 486 (orthographic variant of Geonoma desmarestii).
Geonoma desmarestii Martius, 1843: 23. Type. Bolivia. Cochabamba: without locality, no date, A. dOrbigny 50
(holotype, P!).
Geonoma dicranospadix Burret, 1930a: 169. Type. Colombia. Antioquia: without locality, no date, W. Kalbreyer s. n.
Geonoma discolor Spruce, 1871: 110. Type. Brazil. Par: Rio Tapajs, no date, R. Spruce 36 (holotype, K n.v.).
Geonoma donnell-smithi Dammer = Calyptrogyne ghiesbreghtiana (Linden & Wendland) Wendland
Geonoma dulcis C. H. Wright ex Griseb. = Calyptronoma plumeriana (Mart.) Lourteig
Geonoma erythrospadice Barbosa Rodrigues, 1879: 41. Lectotype, Wessels Boer, 1968: Barbosa Rodrigues, 1903: t. 24.
Geonoma estevaniana Burret, 1938a: 256. Type. Brazil. Par: Utinga, no date, M. Burret 208 (holotype, B, destroyed).
Geonoma falcata Barbosa Rodrigues, 1875: 10. Lectotype, Wessels Boer, 1968: Barbosa Rodrigues, Sert. Palm. Brasil.
1903: t. 19.
Geonoma fenestrata (Wendland) Wendland = Chamaedorea geonomiformis Wendland
Geonoma ferruginea Linden, 1881. Type not designated.
Geonoma floccosa Dammer ex Burret, 1930a: 203. Type. Peru. Junn: Tarma, Huacapistana, 2700 m, 20 January 1903, A.
Geonoma frigida Linden, 1881: 31. Type not designated.
Geonoma furcifolia Barbosa Rodrigues, 1875: 11. Lectotype, Wessels Boer, 1968: Barbosa Rodrigues, 1903: t. 15.
Geonoma furcifrons Drude, 1882: 502 = Geonoma furcifolia Barbosa Rodrigues
Geonoma gamiova Barbosa Rodrigues, 1907: 13. Type. Brazil. Santa Catarina: Blumenau, no date, Anon s. n. (holotype,
not known).
Geonoma ghiesbreghtiana Linden & Wendland = Calyptrogyne ghiesbreghtiana (Linden & Wendland) Wendland
Geonoma gibbosa Burret, 1936: 342. Type. Ecuador. Pichincha: San Carlos de los Colorados, 150 m, 12 December 1935,
H. Schultze-Rhonhof 1994 (holotype, B, destroyed).
Geonoma glauca Orsted = Calyptrogyne ghiesbreghtiana (Linden & Wendland) Wendland
Geonoma gracilipes Dammer ex Burret, 1930a: 173. Type. Peru. Loreto: Moyobamba, 11001200 m, 12 August 1904,
A. Weberbauer 4557 (holotype, B, destroyed).
Geonoma gracillima Burret, 1930a: 165. Type. Colombia. Antioquia: Murr, 9001000 m, 22 July 1880, W. Kalbreyer
1824 (holotype, B, destroyed). (A neotype was designated for G. gracillima by Bernal et al., 1989Colombia.
Antioquia: Municipio de Frontino, Corregimiento de Murr, La Blanquita, 815 m, 22 March 1982, R. Bernal & G.
Galeano 286 (COL)but this same specimen was also used, in the same publication, as neotpye for Geonoma
cuneatoidea. This specimen, R. Bernal & G. Galeano 286, is retained as the type of G. cuneatoidea, and G.
gracillima is treated as an excluded name.)
Geonoma grandifrons Burret, 1930a: 163. Type. Colombia. Antioquia: Cienegatas, 1400 m, 23 July 1880, W. Kalbreyer
Geonoma grandisecta Burret, 1930a: 258. Type. Brazil. Amazonas: Manaus, August 1928, G. Huebner 106 (holotype, B,
destroyed).
Geonoma granditrijuga Burret, 1930a: 171. Type. Peru. Hunuco: Monzon, Huallaga, 700800 m, September 1903, A.
Geonoma heinrichsiae Burret, 1934b: 43. Type. Ecuador. Tungurahua: Ambato, near Baos, 2100 m, 18 January 1933,
E. Heinrichs 225 (holotype, B, destroyed; isotype, F!).
Geonoma helminthoclada Burret, 1930a: 222. Type. Peru. Amazonas: Chachapoyas, no date, A. Raimondi 509 (holotype,
B, destroyed).
Geonoma herbstii auct., in Anon, 1889: 463. Type not designaged.
Geonoma herthae Burret, 1939: 325. Type. Ecuador. Pacayacu, 200 m, 11 June 1937, H. Schultze-Rhonhof 2394
Geonoma hoppii Burret, 1931b: 235. Type. Ecuador. Napo: Archidona, no date, W. Hopp 1039 (holotype, B, destroyed).
173
Geonoma huebneri Burret, 1930a: 254. Type. Colombia. Amazonas: Sierra de Yupat near La Pedrera, no date, G.
Huebner 43 (holotype, B, destroyed).
Geonoma humilis auct. = Chamaedorea geonomiformis Wendland
Geonoma imperialis Linden, 1881: 31. Type not designated.
Geonoma insignis Burret, 1940a: 28. Type. Ecuador. Pastaza: Mera, no date, H. Schultze-Rhonhof 2892 (holotype, B,
destroyed).
Geonoma intermedia (Wendland) Williams = Calyptronoma plumeriana (Martius) Lourteig
Geonoma iodoneura Burret, 1930a: 210. Type. Colombia. Santander: Teorama, 13001700 m, 15 January 1881, W.
Geonoma iraze Linden, 1881: 31. Type not designated.
Geonoma kalbreyeri Burret, 1930a: 168. Type. Colombia. Antioquia: Pulperia, 2000 m, 7 May 1880, W. Kalbreyer 1642
Geonoma lacerata auct., in Dombrain, 1869: plate 446. Type not designated.
Geonoma lakoi Burret, 1930a: 253. Type. Brazil. Amazonas: Rio Manacaparu, 10 May 1929, C. Lak/G. Huebner 116
Geonoma lanceolata Burret, 1930c: 7. Type. Brazil. Amazonas: Rio Ia, May 1930, C. Lak 18 (holotype, B, destroyed).
Geonoma latifolia Burret, 1933b: 102. Type. Brazil. Sergipe: Serra do Itabaiana, no date, E. Werdermann 3065 (holotype,
B, destroyed).
Geonoma latifrons Burret = Chamaedorea ernesti-augusti Wendland
Geonoma latisecta Burret, 1930a: 255. Type. Brazil. Amazonas: near Manaus, no date, G. Huebner 30 (holotype, B,
destroyed).
Geonoma leptoclada Burret, 1933a: 863. Type. Guatemala. Quezaltenango: Volcan de Santa Maria, 2000 m, 22 June
1882, F. Lehmann 1613 (holotype, B, destroyed).
Geonoma leucotricha Burret, 1930a: 204. Type. Colombia. Department unknown: Tibajes, no date, W. Kalbreyer 2035
Geonoma longipes hort. ex Wendland in Kerchove de Dentergarten, 1878, 245. Type not designated.
Geonoma macroclada Burret, 1930a: 220. Type. Colombia. Antioquia: Titiribi, 23002650 m, 22 March 1880, W.
Geonoma macrophylla Burret, 1940a: 27. Type. Ecuador. Pastaza: Mera, 1000 m, 7 September 1938, H. SchultzeRhonhof 2794 (holotype, B, destroyed).
Geonoma macrosiphon Burret, 1930a: 214. Type. Ecuador. Province unknown: Nauegol, August 1874, L. Sodiro s. n.
Geonoma macroura Burret, 1930a: 202. Type. Colombia. Antioquia: Tambu, 23002550 m, 16 July 1880, W. Kalbreyer
Geonoma martiana Wendland = Asterogyne martiana (Wendland) Wendland ex Drude
Geonoma megaloptila Burret, 1930a: 247. Type. Colombia. Santander: Catatumbo, 800900 m, 16 February 1881, W.
Geonoma microstachys Wendland ex Burret, 1930a: 228. Type. Costa Rica. Heredia: Sarapiqu valley, no date, H.
Wendland s. n. (holotype, K n.v.).
Geonoma molinillo Burret, 1937b: 491. Type. Colombia. El Valle: Ro Nima, 2700 m, 4 January 1937, J. Duque 537
Geonoma olfersiana Klotzsch ex Drude, 1882: 506. Type. Brazil. Rio de Janeiro: no locality, no date, F. Sellow s. n.
(holotype, not known).
Geonoma oligoclada Burret, 1930c: 9. Type. Brazil. Amazonas: Rio I, no date, C. Lak 7 (Huebner 138) (holotype, B,
destroyed).
Geonoma pachyclada Burret, 1930a: 214. Type. Colombia. Santander: San Pedro, 20002830 m, December 1877, W.
Geonoma palustris Barbosa Rodrigues, 1875: 11. Lectotype, Wessels Boer, 1968: Barbosa Rodrigues, 1903: t. 27.
Geonoma paraguanensis Karsten, 1856: 410. Type. Venezuela. Falcn: Peninsula Paraguan, Monte Santa Ana, no date,
H. Karsten s. n. (holotype, LE, destroyed).
Geonoma pilosa Barbosa Rodrigues, 1882: 43. Lectotype, Wessels Boer, 1968: Barbosa Rodrigues, 1903: t. 23.
Geonoma pleeana Martius, 1843: 33. Type. Venezuela. Zulia: Maracaibo, no date, A. Ple s. n. (holotype, P n.v.).
Geonoma pleioneura Burret, 1931b: 234. Type. Ecuador. Napo: Archidona, no date, W. Hopp 1040 (holotype, B;
holotype image!).
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Geonoma plumeriana Martius = Calyptrogyne plumeriana (Martius) Lourteig

Geonoma polyclada Burret, 1940a: 26. Type. Ecuador. Pastaza: Mera, 1000 m, 7 September 1938, H. Schultze-Rhonhof
Geonoma porteana Wendland, 1856: 340. Type. Cultivated plant, no date, H. Wendland s. n. (holotype, K n.v.).
Geonoma preussii Burret, 1930a: 242. Type. Mexico. State unknown: Isthmus of Tehuantepec, 18 April 1900, H. Preuss
Geonoma princeps Linden, 1881: 31. Type not designated.
Geonoma pulchella Linden, 1881: 31. Type not designated.
Geonoma pynaertiana Sander = Iguanura wallichiana (Martius) Beccari
Geonoma raimondii Burret, 1930a: 182. Type. Peru. Amazonas: without locality, no date, A. Raimondi 978 (holotype, B,
destroyed).
Geonoma rectifolia Wallace, 1853: 67. Lectotype, Wessels Boer, 1968: Wallace, 1853: t. 25.
Geonoma rhytidocarpa Burret, 1930a: 189. Type. Colombia. Santander: Sisabita, no date, W. Kalbreyer 1115 (holotype,
B, destroyed).
Geonoma riedeliana Wendland in Kerchove de Dentergarten, 1878: 245. Type not designated.
Geonoma robusta Burret, 1930a: 259. Type. Guyana. Canawaruk River, September 1905, A. Bartlett 6/8189 (holotype,
B, destroyed).
Geonoma rodeiensis Barbosa Rodrigues, 1882: 42. Lectotype, Wessels Boer, 1968: Barbosa Rodrigues, 1903: t. 11.
Geonoma rupestris Barbosa Rodrigues, 1882: 47. Lectotype, Wessels Boer, 1968: Barbosa Rodrigues, 1903: t. 31A.
Geonoma seemanni auct., in Dombrain, 1869: plate 428. Type not designated.
Geonoma solitaria (Engel) Jahn ex Hill, 1929: 102. Roebelia solitaria Engel, 1865: 680. Lectotype, Glassman, 1972:
Engel 1865: t. 3, fig. 5.
Geonoma speciosa Barbosa Rodrigues, 1875: 9. Lectotype, Wessels Boer, 1968: Barbosa Rodrigues, 1903: t. 18.
Geonoma spicigera Koch = Calyptrogyne ghiesbreghtiana (Linden & Wendland) Wendland
Geonoma spruceana subsp. spruceana var. micra Trail, 1876: 329. Type. Brazil. Par: Lago Juruty, no date, J. Trail 29
(holotype, K n.v.).
Geonoma stuebelii Burret, 1930a: 220. Type. Colombia. Cauca: Popayan, Cerro Munchique, June 1869, A. Stuebel 321e
Geonoma swartzii Grisebach & Wendland = Calyptrogyne occidentalis (Swartz.) Gmez
Geonoma synanthera Martius = Pholidostachys synanthera (Martius) Moore
Geonoma tenuifolia auct., in de Bosschere, 1895, 186. Type not designated.
Geonoma tenuifolia Burret, 1940a: 25. Type. Ecuador. Pastaza: Mera, 1200 m, 15 November 1938, H. Schultze-Rhonhof
Geonoma tessmannii Burret, 1930a: 181. Type. Peru. Amazonas: Ro Maraon, 7 October 1924, G. Tessmann 4225
Geonoma tomentosa Barbosa Rodrigues, 1882: 44. Lectotype, Wessels Boer, 1968: Barbosa Rodrigues, 1903: t. 26A.
Geonoma trichostachys Burret, 1933a: 862. Type. Colombia. Cauca: Guanacas, 28003200 m, 27 February 1883, F.
Lehmann 2640 (holotype, B, destroyed).
Geonoma trigonostyla Barbosa Rodrigues, 1882: 46. Lectotype, Wessels Boer, 1968: Barbosa Rodrigues, 1903: t. 20, 21.
Geonoma trifurcata Orsted = Asterogyne martiana (Wendland) Wendland ex Drude
Geonoma trijugata Barbosa Rodrigues, 1875: 12. Lectotype, Wessels Boer, 1968: Barbosa Rodrigues, 1903: t. 14.
Geonoma uncibracteata Burret, 1930a: 215. Type. Colombia. Nario: Cuchilla de Patascoy, near Pasco, 3200 m, August
1869, A. Stuebel 367a (holotype, B, destroyed).
Geonoma uliginosa Barbosa Rodrigues, 1875: 11. Lectotype, Wessels Boer, 1968: Barbosa Rodrigues, 1903: t. 28.
Geonoma ventricosa Engel, 1865: 688. Type. Venezuela. Trujillo, no date, F. Engel s. n. (holotype, unknown).
Geonoma verdugo Linden, 1881: 31. Type not designated.
Geonoma verschaffelti hort. ex Wendland in Kerchove de Dentergarten, 1878: 246. Type not designated.
Geonoma wallisii Linden ex Wendland in Kerchove de Dentergarten, 1878: 246. Type not designated.
Geonoma wendlandiana Burret, 1930a: 192. Type. Colombia. Antioquia: Concordia, 21502350 m, 27 March 1880, W.
Geonoma wendlandii auct., in Anon, 1878: 440. Type. Not designated.
Geonoma werdermannii Burret, 1930a: 173. Type. Bolivia. Beni: Mission Todos Santos, 300 m, 2 August 1926, E.
Werdermann 2183 (holotype, B, destroyed).
Geonoma woronowii Burret, 1930c: 6. Type. Colombia. Caquet: Getuch, Ro Orteguaza, 21 July 1926, G. Woronow &
175
S. Juzepczuk 6119 (holotype, B, destroyed).

Geonoma yauaperyensis Barbosa Rodrigues, 1902: 88. Lectotype, Wessels Boer, 1968: Barbosa Rodrigues, 1903: t. 30.
Geonoma zamorensis Linden ex Wendland in Kerchove de Dentergarten, 1878: 246. 1878. Type not designated.
Taenianthera gracilis Burret, 1930c: 14. Type. Brazil. Amazonas: Rio Ia, April 1930, C. Lak 10 (holotype, B,
destroyed).
Taenianthera lakoi Burret, 1930c: 11. Type. Brazil. Roraima: Rio Catrimany, Aamaro, Cachoeira do Mirity, November
1929, C. Lak (G. Huebner 128) (holotype, B, destroyed).
Taenianthera minor Burret, 1939: 324. Type. Ecuador. Pastaza: Canelos, 350 m, 12 January 1937, H. Schultze-Rhonhof
Taenianthera weberbaueri Burret, 1930a: 269. Type. Peru. Hunuco: Huamelies, Monzon, 9001000 m, 27 July 1903, A.
Vouay Aublet 1775: 99. Type not designated.
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Appendix IV. Plates of Type Images
Appendix IV, Plate 1. Isotype of Geonoma bernalii (R. Bernal 1399, NY).
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178
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Appendix IV, Plates 2 & 3. Isotype of Geonoma brongniartii subsp. pascoensis (A. Henderson, E. Ferreira & M. Arakaki
3012, NY).
179
180
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Appendix IV, Plates 4 & 5. Holotype of Geonoma concinna subsp. simplex (A. Gentry & M. Monsalve 48188, NY).
181
182
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Appendix IV, Plates 6 & 7. Holotype of Geonoma concinnoidea (G. de Nevers & H. Herrera 6942, NY).
183
Appendix IV, Plate 8. Isotype of Geonoma concinnoidea subsp. coclensis (A. Henderson & E. Ferreira 3028, NY).
184
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Appendix IV, Plate 9. Holotype of Geonoma concinnoidea subsp. jefensis (S. Mori & L. Joly 7933, MO).
185
186
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187
188
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189
190
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Appendix IV, Plates1015. Holotype of Geonoma congesta subsp. osensis (H. Moore 6534, NY).
191
Appendix IV, Plate 16. Holotype of Geonoma cuneata subsp. guanacastensis (L. Gmez et al. 19053, NY).
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Appendix IV, Plate 17. Holotype of. Geonoma cuneata subsp. indivisa (G. de Nevers, A. Henderson, H. Herrera, G.
McPherson & L. Brako 6293, NY).
193
Appendix IV, Plate 18. Holotype of Geonoma cuneata subsp. minor (G. de Nevers, A. Henderson, H. Herrera, G
McPherson & L. Brako 6355, NY).
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195
Appendix IV, Plates 19 & 20. Holotype of Geonoma cuneata subsp. rubra (R. Bernal, W. Devia, E. Linhares & J. Angulo
1770, COL).
196
HENDERSON
197
198
HENDERSON
199
Appendix IV, Plates 2124. Holotype of Geonoma deneversii (G. de Nevers, F. Almeda & G. McPherson 8823, NY).
200
HENDERSON
Appendix IV, Plate 25. Holotype of Geonoma deversa subsp. belizensis (M. Balick, R. Arvigo, P. Cocom, R. Cocom, H.
Robinson & G. Shropshire 2698, NY).
201
Appendix IV, Plate 26. Isotype of Geonoma deversa subsp. peninsularis (A. Henderson, G. Galeano & B. Hammel 1817,
NY).
202
HENDERSON
203
204
HENDERSON
205
Appendix IV, Plates 2730. Isotype of Geonoma deversa subsp. quadriflora (G. Galeano, R. Bernal, A. Henderson & S.
Churchill 2112, NY).
206
HENDERSON
207
Appendix IV, Plates 31 & 32. Holotype of Geonoma dindoensis (A. Gentry & M. Monsalve 48419, NY).
208
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Appendix IV, Plate 33. Holotype of Geonoma ferruginea subsp. nicaraguensis (J. Pipoly 5112, NY).
209
Appendix IV, Plate 34. Isotype of Geonoma fosteri (R. Aguinda, N. Pitman & R. Foster 1315, F).
210
HENDERSON
211
212
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Appendix IV, Plates 3537. Isotype of Geonoma galeanoae (R. Callejas, J. Betancur, A. Arbelaez & H. Correa 4208,
NY).
213
Appendix IV, Plate 38. Holotype of Geonoma gentryi (A. Gentry & M. Fallen 17623, MO).
214
HENDERSON
Appendix IV, Plate 39. Holotype of Geonoma lehmannii subsp. corrugata (G. de Nevers & S. Charnley 6684, NY).
215
216
HENDERSON
Appendix IV, Plates 40 & 41. Isotype of Geonoma longivaginata subsp. copensis (G. de Nevers, A. Henderson, H.
Herrera, G. McPherson & L. Brako 6392, NY).
217
218
HENDERSON
219
Appendix IV, Plates 4244. Holotype of Geonoma longivaginata subsp. sanblasensis (G. de Nevers & H. Herrera 7957,
NY).
220
HENDERSON
Appendix IV, Plate 45. Isotype of Geonoma longivaginata subsp. vallensis (A. Henderson & R. Bernal 2039, NY).
221
222
HENDERSON
223
Appendix IV, Plates 4648. Holotype of Geonoma maxima subsp. dispersa (R. Bernal, G. Galeano & D. Restrepo 1148,
COL).
224
HENDERSON
225
Appendix IV, Plates 49 & 50. Holotype of Geonoma maxima subsp. multiramosa (H. Balslev, A. Barfod, A. Henderson,
F. Skov & A. Argello 60731, NY).
226
HENDERSON
Appendix IV, Plate 51. Isotype of Geonoma maxima subsp. sigmoidea (G. Galeano & J. Huitoto 1279, NY!).
227
Appendix IV, Plate 52 Isotype of Geonoma operculata (W. Meier 3401, NY).
228
HENDERSON
Appendix IV, Plate 53. Holotype of Geonoma peruviana (J. Schunke 9416, MO).
229
Appendix IV, Plate 54. Isotype of Geonoma pohliana subsp. linharensis (G. Farias 258, NY).
230
HENDERSON
Appendix IV, Plate 55. Holotype of Geonoma pohliana subsp. rodriguesii (G. Gottsberger 14-191166, NY).
231
Appendix IV, Plate 56. Isotype of Geonoma pohliana subsp. unaensis (A. Amorim, S. SantAna, J. Jardim, E. Santos & J.
Hage 1119, NY).
232
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Appendix IV, Plate 57. Isotype of Geonoma sanmartinensis (I. Snchez Vega & M. Dillon 8658, NY).
233
234
HENDERSON
Appendix IV Plates 58 & 59. Holotype of Geonoma schizocarpa (R. Kyap 1212, MO).
235
Appendix IV, Plate 60. Holotype of Geonoma stricta subsp. antioquiensis (A. Cogollo & R. Torres 2397, COL).
236
HENDERSON
Appendix IV, Plate 61. Holotype of Geonoma stricta subsp. bracteata (H. Rainer P22-10988, NY).
237
Appendix IV, Plate 62. Holotype of Geonoma stricta subsp. divaricata (R. Rojas, A. Pea & E. Chvez 101, NY).
238
HENDERSON
Appendix IV, Plate 63. Holotype of Geonoma stricta subsp. pendula (R. Rojas, A. Pea & E. Chvez 256, NY).
239
Appendix IV, Plate 64. Isotype of Geonoma stricta subsp. pliniana (J.-J. de Granville & G. Cremers 13148, NY).
240
HENDERSON
Appendix IV, Plate 65. Holotype of Geonoma stricta subsp. quibdoensis (M. Grayum, B. Hammel, J. Kress & G. Brown
7645, MO).
241
Appendix IV, Plate 66. Holotype of Geonoma stricta subsp. submontana (D. Smith 2019, NY).
242
HENDERSON
Appendix IV, Plate 67. Holotype of Geonoma undata subsp. tacarcunensis (G. de Nevers, B. Hammel & H. Herrera
8511, NY).
243
Appendix IV, Plate 68. Holotype of Geonoma undata subsp. tumucensis (J.-J. de Granville, P. Acevedo, A. Boyer & L.
Hollenberg 12322, NY).
244
HENDERSON
245
246
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Appendix IV, Plates 6971. Isotype of Geonoma undata subsp. venezuelana (F. Stauffer, A. Fernndez, R. Riina & K.
Walther-Weissbeck 262, NY).
247
Appendix IV, Plate 72. Holotype of Geonoma venosa (H. Vargas, W. Defas & D. Reyes 6282, NY).
248
HENDERSON
Appendix V. Numerical List of Taxa and Specimens Examined

Numerical List of Taxa
1. Geonoma aspidiifolia
1a. Geonoma aspidiifolia subsp. aspidiifolia
1b. Geonoma aspidiifolia subsp. fusca
2. Geonoma baculifera
3. Geonoma bernalii
4. Geonoma braunii
5. Geonoma brenesii
6. Geonoma brongniartii
6a. Geonoma brongniartii subsp. brongniartii
6b. Geonoma brongniartii subsp. pascoensis
7. Geonoma calyptrogynoidea
8. Geonoma camana
9. Geonoma chlamydostachys
10. Geonoma chococola
10a. Geonoma chococola subsp. chococola
10b. Geonoma chococola subsp. awaensis
11. Geonoma concinna
11a. Geonoma concinna subsp. concinna
11b. Geonoma concinna subsp. simplex
12. Geonoma concinnoidea
12a. Geonoma concinnoidea subsp. concinnoidea
12b. Geonoma concinnoidea subsp. coclensis
12c. Geonoma concinnoidea subsp. jefensis
13. Geonoma congesta
13a. Geonoma congesta subsp. congesta
13b. Geonoma congesta subsp. osensis
14. Geonoma cuneata
14a. Geonoma cuneata subsp. cuneata
14b. Geonoma cuneata subsp. guanacastensis
14c. Geonoma cuneata subsp. indivisa
14d. Geonoma cuneata subsp. irena
14e. Geonoma cuneata subsp. linearis
14f. Geonoma cuneata subsp. minor
14g. Geonoma cuneata subsp. procumbens
14h. Geonoma cuneata subsp. rubra
14i. Geonoma cuneata subsp. sodiroi
15. Geonoma deneversii
16. Geonoma deversa
16a. Geonoma deversa subsp. deversa
16b. Geonoma deversa subsp. belizenesis
16c. Geonoma deversa subsp. peninsularis
16d. Geonoma deversa subsp. quadriflora
17. Geonoma dindoensis
18. Geonoma divisa
19. Geonoma elegans
20. Geonoma epetiolata
21. Geonoma euspatha
22. Geonoma ferruginea
22a. Geonoma ferruginea subsp. ferruginea
22b. Geonoma ferruginea subsp. microspadix
249
22c. Geonoma ferruginea subsp. nicaraguensis

23. Geonoma fosteri
24. Geonoma frontinensis
25. Geonoma galeanoae
26. Geonoma gentryi
27. Geonoma hollinensis
28. Geonoma hugonis
29. Geonoma interrupta
29a. Geonoma interrupta subsp. interrupta
29b. Geonoma interrupta subsp. magnifica
29c. Geonoma interrupta subsp. purdieana
29d. Geonoma interrupta subsp. rivalis
30. Geonoma lanata
31. Geonoma laxiflora
32. Geonoma lehmannii
32a. Geonoma lehmannii subsp. lehmannii
32b. Geonoma lehmannii subsp. corrugata
33. Geonoma leptospadix
34. Geonoma longipedunculata
35. Geonoma longivaginata
35a. Geonoma longivaginata subsp. longivaginata
35b. Geonoma longivaginata subsp. copensis
35c. Geonoma longivaginata subsp. sanblasensis
35d. Geonoma longivaginata subsp. vallensis
36. Geonoma macrostachys
37. Geonoma maxima
37a. Geonoma maxima subsp. maxima
37b. Geonoma maxima subsp. ambigua
37c. Geonoma maxima subsp. camptoneura
37d. Geonoma maxima subsp. chelidonura
37e. Geonoma maxima subsp. compta
37f. Geonoma maxima subsp. dispersa
37g. Geonoma maxima subsp. hexasticha
37h. Geonoma maxima subsp. multiramosa
37i. Geonoma maxima subsp. sigmoidea
37j. Geonoma maxima subsp. spixiana
38. Geonoma monospatha
39. Geonoma mooreana
40. Geonoma multisecta
41. Geonoma occidentalis
42. Geonoma oldemanii
43. Geonoma oligoclona
44. Geonoma operculata
45. Geonoma orbignyana
45a. Geonoma orbignyana subsp. orbignyana
45b. Geonoma orbignyana subsp. hoffmanniana
46. Geonoma paradoxa
47. Geonoma pauciflora
48. Geonoma peruviana
49. Geonoma pinnatifrons
49a. Geonoma pinnatifrons subsp. pinnatifrons
49b. Geonoma pinnatifrons subsp. binervia
49c. Geonoma pinnatifrons subsp. martinicensis
49d. Geonoma pinnatifrons subsp. membranacea
250
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49e. Geonoma pinnatifrons subsp. mexicana

49f. Geonoma pinnatifrons subsp. oxycarpa
49g. Geonoma pinnatifrons subsp. platybothros
49h. Geonoma pinnatifrons subsp. ramosissima
49i. Geonoma pinnatifrons subsp. vaga
50. Geonoma poeppigiana
51. Geonoma pohliana
51a. Geonoma pohliana subsp. pohliana
51b. Geonoma pohliana subsp. fiscellaria
51c. Geonoma pohliana subsp. gastoniana
51d. Geonoma pohliana subsp. kuhlmannii
51e. Geonoma pohliana subsp. linharensis
51f. Geonoma pohliana subsp. rodriguesii
51g. Geonoma pohliana subsp. rubescens
51h. Geonoma pohliana subsp. trinervis
51i. Geonoma pohliana subsp. unaensis
51j. Geonoma pohliana subsp. weddelliana
51k. Geonoma pohliana subsp. wittigiana
52. Geonoma poiteauana
53. Geonoma sanmartinensis
54. Geonoma santanderensis
55. Geonoma schizocarpa
56. Geonoma schottiana
57. Geonoma scoparia
58. Geonoma simplicifrons
59. Geonoma spinescens
60. Geonoma stricta
60a. Geonoma stricta subsp. stricta
60b. Geonoma stricta subsp. antioquiensis
60c. Geonoma stricta subsp. arundinacea
60d. Geonoma stricta subsp. bracteata
60e. Geonoma stricta subsp. divaricata
60f. Geonoma stricta subsp. pendula
60g. Geonoma stricta subsp. pliniana
60h. Geonoma stricta subsp. quibdoensis
60i. Geonoma stricta subsp. submontana
61. Geonoma talamancana
62. Geonoma tenuissima
63. Geonoma triandra
64. Geonoma triglochin
65. Geonoma trigona
66. Geonoma umbraculiformis
67. Geonoma undata
67a. Geonoma undata subsp. undata
67b. Geonoma undata subsp. appuniana
67c. Geonoma undata subsp. dussiana
67d. Geonoma undata subsp. edulis
67e. Geonoma undata subsp. pulcherrima
67f. Geonoma undata subsp. skovii
67g. Geonoma undata subsp. stenothyrsa
67h. Geonoma undata subsp. tacarcunensis
67i. Geonoma undata subsp. tumucensis
67j. Geonoma undata subsp. venezuelana
68. Geonoma venosa
251
Specimens Examined
Specimens are arranged by collector (with first initial, when known) in alphabetical order, followed by collectors
number in increasing order (s. n. = without number), followed by species number in parentheses.
Abraho, G. 5815 (56)
Acevedo, P. 1595 (37d); 1719 (14a); 4880 (42); 6752 (16a); 6862 (63); 8140 (16a); 8145 (60c); 8290 (16a); 9775 (60c);
9960 (6a); 9991 (29a)
Acosta, L. 527 (45b); 1301 (13b); 1408 (16c); 1534 (35a); 1643 (35a); 2153 (35a)
Adams, C. 92 (49i)
Affonso, P. 307 (56)
Agostini, G. 89 (58); 689 (58)
Aguilar, G. 889 (57); 2169 (13b); 4983 (29b)
Aguilar, M. 411 (37c)
Aguilar, R. 1824 (29b); 3248 (16c); 4025 (45b); 4575 (14a);
Aguinda, R. 696 (60c); 1046 (64); 1069 (67a); 1086 (64); 1240 (23); 1315 (23); 1700 (60c)
Alexiades, M. 233 (60c); 923 (37c); 1010 (16a)
Alfaro, E. 3903 (45b)
Alston, A. 1838 (14c)
Altamirano, S. 3445 (37d)
Alvarado, A. 104 (36)
Alvarez, A. 1611 (67a)
Alverson, W. 92 (9)
Amaral, I. 1015 (51j); 1311 (36); 1657 (67b)
Amorim, A. 640 (51a); 1119 (51i); 4100 (51a); 4208 (51a); 4457 (47); 6124 (47)
Ancuash, E. 14 (50); 267 (60c); 1418 (55)
Anderson, A. 20 (7); 22 (14h); 26 (10a); 274 (1a)
Anderson, W. 6289 (51j); 6613 (51j); 8138 (51j); 10592 (36); 11127 (37d); 35977 (51j)
Andrade, P. 1430 (56)
Andrade-Lima, D. 552331 (51a)
Anon s. n. (60c); 42 (37c); 54 (16a); 16920 (13a)
Antezana, A. 146 (45a)
Antonio, T. 1214 (35b); 1864 (38); 2599 (32b); 2784 (28); 3046 (35b); 3956 (14a); 4092 (28); 4331 (14g)
Araquistain, M. 2523 (22c); 2524 (13a); 2588 (22c); 2698 (22c)
Araujo, A. 1107 (29a)
Araujo, D. 1059 (19); 1836 (19); 2069 (51k)
Arbo, M. 4767 (51j)
Archer, W. 74 (51j); 1370 (32a); 1980 (7); 2162 (16a)
Argello, A. 393 (14a); 636 (67e)
Arias, J. 1186 (43); 1372 (33)
Aristeguieta, L. 1876 (49a); 7415 (2)
Aronson, J. 729 (31); 923 (60c)
Arroyo, L. 608 (51j)
Asplund, E. 9354 (36)
Atwood, J. 50 (29b)
Aulestia, C. 322 (10b); 418 (7); 1042 (30); 1146 (30); 1200 (30); 1244 (30)
Aulestia, M. 275 (60c); 1005 (10b); 1645 (60c); 1835 (60c); 2077 (64); 2081 (37h); 2085 (50)
Avril, C. 2 (33)
Ayala, F. 2366 (60c); 2776 (37d); 2778 (60c); 3425 (37d)
Aymard, G. 3091 (29c); 3967 (33); 4003 (37b); 5344 (37b); 6117 (16a); 7290 (2); 7322 (33); 7825 (2); 7848 (16a); 7857
(16a); 8161 (52); 8503 (2); 8631 (16a); 8923 (37g); 9719 (2); 9721 (37d)
Bacon, C. 62 (14a); 63 (13a)
Bailey, L. 697 (19)
Baker, M. 5638 (67a); 5911 (36); 5960 (36); 6368 (36); 6671 (29a); 6788 (36); 6905 (37h); 6912 (60c); 7018 (8)
252
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Bale, W. 1549 (2); 1591 (37a); 2675 (2); 2965 (33); 3347 (2)
Balick, M. 923 (36); 926 (16a); 928 (36); 932 (37d); 936 (52); 938 (16a); 939 (36); 942 (37d); 944 (36); 947 (60c); 1043
(6a); 1045 (50); 1046 (16a); 1131 (8); 1137 (8); 1138 (60c); 1139 (36); 1140 (36); 1141 (6a); 1154 (29a); 1202 (16a);
1303 (2); 1364 (37e); 1587 (51j); 1726 (67d); 2698 (16b); 2704 (29b)
Balslev, H. 1594 (36); 1595 (60c); 2160 (67a); 2335 (6a); 2391 (36); 4270 (49a); 4271 (67a); 4273 (67a); 4276 (60c);
4280 (67a); 4284 (67a); 4292 (67a); 4297 (14a); 4300 (16a); 4302 (60c); 4303 (34); 4306 (37h); 4312 (16a); 4319
(36); 4320 (60c); 4325 (36); 4372 (36); 4415 (21); 4420 (60c); 4424 (60c); 4437 (67a); 4456 (67a); 4542 (36); 4569
(36); 4624 (8); 4626 (60c); 4668 (14a); 4723 (14a); 4777 (40); 4781 (64); 4782 (60c); 4783 (34); 4791 (16a); 4801
(40); 4811 (60c); 4814 (6a); 4816 (36); 4825 (36); 4828 (6a); 4838 (60c); 4861 (36); 4863 (36); 4872 (36); 6080
(60c); 6148 (60c); 6151 (60c); 6156 (60c); 6201 (60c); 6216 (60c); 6255 (60c); 6296 (60c); 6297 (60c); 6326 (60c);
6384 (64); 6385 (37h); 6387 (60c); 6396 (60c); 6400 (40); 6402 (16a); 6405 (50); 6418 (27); 6419 (60c); 6420 (36);
6422 (36); 6430 (36); 6434 (60c); 6442 (21); 6454 (60c); 6472 (67a); 6475 (34); 6481 (60c); 6482 (60c); 6501 (45a);
6521 (67a); 6553 (8); 6566 (6a); 6569 (36); 6570 (36); 6583 (6a); 10552 (67a); 10629 (36); 60506 (36); 60517 (36);
60520 (37h); 60536 (40); 60571 (6a); 60579 (36); 60584 (60c); 60609 (36); 60640 (45a); 60643 (67f); 60644 (67a);
60649 (29a); 60661 (67a); 60662 (60c); 60663 (67a); 60691 (6a); 60692 (29a); 60731 (37h); 60734 (60c); 60739
(40); 62009 (67a); 62018 (14d); 62019 (14i); 62025 (14d); 62030 (14a); 62035 (60c); 62043 (49a); 62053 (60c);
62054 (16a); 62055 (34); 62060 (60c); 62061 (36); 62063 (36); 62064 (64); 62066 (40); 62071 (64); 62075 (36);
62081 (60c); 62082 (45a); 62086 (45a); 62087 (67a); 62099 (30); 62101 (14a); 62112 (14a); 62201 (60c); 62206
(64); 62207 (36); 62209 (60c); 62210 (36); 62211 (34); 62216 (8); 62406 (34); 62407 (60c); 62409 (36); 62410
(60c); 62411 (40); 62412 (60c); 62430 (60c); 62434 (36); 62448 (67a); 62468 (64); 62471 (60c); 62475 (29a);
62489 (67a); 62492 (45a); 62498 (36); 62501 (36); 62502 (29a); 67201 (40); 69024 (60c); 69026 (16a); 69027 (34);
69028 (40); 69033 (60c); 69040 (40); 69041 (60c); 69042 (40); 69043 (64)
Bang, M. 877 (45a)
Barbosa, C. 6279 (14a); 6480 (7); 6533 (63); 6616 (14a)
Barbour, P. 446 (60e); 2742 (45a); 3718 (45a); 4126 (67a); 4778 (60c); 4932 (60c); 5184 (37c)
Barfod, A. 60042 (67a); 60044 (67a); 60075 (49a); 60115 (14e); 60160 (60c); 60163 (21); 60180 (67a)
Barringer, K. 2541 (5); 3614 (35a); 4001 (5)
Barros, F. 1789 (51a)
Bartlett, H. 16685 (14g); 16743 (14g); 16748 (49b)
Bausen, E. 129 (19)
Beck, H. 163 (42); 238 (16a); 1757 (46); 1758 (14a); 1769 (10b); 2176 (10b); 2218 (30); 2251 (46); 2263 (30); 2271
(10b); 2311 (14a); 3018 (67a)
Beck, S. 13707 (6a); 16466 (6a); 16513 (16a); 16550 (41); 16746 (6a); 18258 (6a)
Belem, R. 3884 (19)
Bello, E. 2661 (49e); 3009 (67d); 5018 (49e)
Beltrn, H. 708 (37h)
Bennett, B. 3571 (60c); 4118 (36)
Berg, C. 699 (2)
Bergmann, B. 62121 (60c); 62126 (60c); 62127 (34); 62128 (37h); 62133 (40); 62163 (8); 62166 (60c); 62169 (21);
62587 (67a); 62588 (60c); 62589 (67f); 62590 (67f); 62591 (67f); 62593 (8); 62596 (60c); 62597 (36); 62598 (60c);
62599 (60c); 67202 (60c); 67206 (60c); 67207 (60c); 67209 (60c); 67212 (8); 67222 (60c); 67225 (36); 67226 (6a);
67227 (60c); 67231 (36); 67236 (30); 97824 (21); 97827 (60c); 97830 (60c); 97847 (60c); 97859 (60c); 97860
(60c); 97874 (67e)
Berlin, B. 214 (55); 476 (55); 576 (55); 671 (50); 964 (55); 1845 (55); 1938 (60e); 2038 (50); 3567 (36)
Bernal, R. 12 (32a); 190 (67g); 217 (67a); 226 (45a); 235 (45a); 285 (49a); 286 (14a); 306 (49h); 365 (67a); 380 (67a);
382 (32a); 390 (32a); 445 (14a); 446 (29b); 448 (7); 468 (29b); 493 (14a); 536 (45a); 569 (32a); 571 (32a); 638
(45a); 655 (14a); 657 (14a); 659 (7); 676 (18); 746 (18); 841 (45a); 843 (45a); 845 (45a); 861 (14a); 864 (16a); 905
(67a); 925 (24); 928 (49a); 979 (67a); 1047 (45a); 1074 (10a); 1077 (14e); 1080 (10a); 1084 (18); 1101 (14a); 1113
(49a); 1114 (14a); 1133 (49a); 1148 (37f); 1161 (14a); 1188 (14a); 1320 (14e); 1342 (45a); 1351 (67a); 1358 (67a);
1362 (45a); 1369 (67a); 1377 (45a); 1383 (24); 1385 (11a); 1387 (67g); 1388 (24); 1394 (67g); 1395 (67a); 1398
(29d); 1399 (3); 1402 (45a); 1408 (36); 1409 (40); 1410 (6a); 1415 (34); 1419 (40); 1421 (64); 1423 (67a); 1427
(36); 1440 (67a); 1443 (67a); 1456 (10a); 1762 (46); 1769 (46); 1770 (14h); 1777 (10a); 2020 (37d); 2021 (60c);
2042 (40); 2043 (8); 2056 (60c); 2072 (6a); 2077 (60c); 2085 (8); 2098 (37g); 2114 (37g); 2140 (14a); 2142 (7);
2153 (14a); 2162 (37f); 2174 (14a); 2519 (6a); 2520 (36); 2521 (36); 2522 (6a); 2537 (36); 2539 (36); 2540 (16d);
253
2542 (6a); 2547 (8); 2558 (50); 2563 (36); 2572 (37i); 2578 (60c); 2620 (60c); 2623 (37d); 2833 (45a); 2870 (67a);
2889 (64); 2899 (45a); 2900 (45a); 2901 (45a); 2930 (16d); 2933 (6a); 2961 (14a); 3483 (54); 3490 (45a); 3512
(45a); 3513 (45a); 3534 (54); 3597 (60c); 3626 (37e); 3627 (37d); 3676 (37d)
Bernal, H. 348 (67a)
Bernardi, A. 831 (21); 2825 (16a); 5699 (58); 5913 (58); 6870 (45a)
Berry, P. 1474 (37g); 1514 (37g)
Betancur, J. 1119 (45a); 1273 (21); 2262 (6a); 2262 (21); 2811 (67a); 2818 (49h); 3818 (67a); 4264 (67a); 4858 (67a);
5429 (32a); 5477 (49a); 5705 (45a); 5714 (45a); 6220 (45a); 6241 (45a); 8007 (63)
Blanc, P. 9391 (52)
Blanchet, M. 25 (51g); 194 (51a)
Blicher-Mathiesen, U. 62601 (34); 62603 (60c)
Boeke, J. 1282 (36); 2114 (67a)
Bonifaz, C. 3799 (63); 3891 (14e)
Boom, B. 4436 (37d); 7634 (67b); 8967 (1b); 9010 (1b); 9222 (67b); 10309 (16a); 10408 (2); 10429 (16a); 10785 (66)
Borchsenius, F. 42 (30); 94 (30); 172 (36); 219 (14a); 220 (14d); 258 (14a); 280 (62); 286 (14d); 298 (45a); 303 (67a);
304 (60c); 305 (36); 310 (67a); 318 (67a); 327 (67a); 332 (45a); 334 (30); 339 (67a); 340 (14a); 343 (7); 345 (14a);
348 (7); 627 (45a); 91426 (45a); 91430 (60c); 91431 (36); 91434 (16a); 91437 (60c); 91438 (8)
Bordenave, B. 8481 (33)
Bosque, C. 10 (49a); 19 (67a); 20 (67a); 27 (67a); 29 (67a); 31 (67a)
Boutin, F. 5145 (29b)
Bovini, M. s. n. (51k)
Boyle, B. 838 (45b); 931 (67d); 1026 (67d); 1081 (67d); 1104 (67d); 2216 (67a); 3440 (67a); 4499 (45a); 6069 (13a)
Brade, A. 15878 (51h); 16345 (51k); 16477 (51d); 18126 (51f); 18319 (51a); 18454 (51f); 19951 (51f)
Braga, J. 1898 (19); 4058 (19)
Brand, J. 419 (14a); 520 (7); 924 (29b)
Brandbyge, J. 30003 (36); 31074 (60c); 31233 (60c); 31410 (60c); 31831 (60c); 31923 (60c); 31940 (34); 32004 (36);
33970 (60c); 36011 (60c)
Braun, A. s. n. (58)
Breedlove, D. 33936 (29b); 34415 (67d); 35114 (67d)
Brenes, A. 4473 (5); 13588 (5); 21848 (49e)
Bristan, N. 556 (63)
Britton, N. 1805 (49i); 1933 (49i); 2278 (49i)
Broadway, W. s. n. (49i); 2701 (49i); 4073 (49i); 5676 (49i); 5928 (49i); 9851 (49i)
Bryon 299 (31)
Buchtien, O. 3670 (45a); 5338 (45a)
Bunting, G. 8512 (49a); 9526 (49a); 10255 (29c); 10836 (16a); 11167 (49a); 11443 (16a)
Burger, W. 3973 (22a); 4263 (35a); 4305 (29b); 4442 (14a); 4569 (67d); 4580 (67d); 4914 (14a); 5196 (14g); 5424 (14a);
6413 (45b); 8373 (45b); 8563 (5); 8703 (67d); 8739 (45b); 8910 (16c); 9785 (45b); 10744 (22a)
Busey, P. 430 (29b)
Byg, A. 61 (60c)
Cabrera, I. 2002 (16a); 2406 (16a); 3328 (36)
Cadena, J. 25 (45a)
Caldern, C. 2427 (51j)
Callejas, R. 2138 (32a); 2386 (14a); 2386 (49a); 2441 (14a); 2628 (14a); 3410 (3); 4098 (29d); 4208 (25); 4824 (14a);
5399 (49a); 8040 (14a); 8560 (9); 8992 (67g); 9205 (60b); 9257 (14a)
Calonje, M. 626 (20)
Calzada, J. 813 (29b)
Camp, W. 1391 (67f)
Campos, J. 1042 (6a); 2828 (32a); 3568 (32a); 4206 (67a); 5495 (45a); 5895 (32a); 5941 (45a); 6286 (32a); 6506 (32a)
Campos, M. 415 (36); 519 (60c); 923 (60c); 925 (6a); 932 (6a); 933 (60c); 940 (60c); 954 (6a)
Campos, M. T. 13624 (56)
Carauta, J. 1527 (56); 2208 (56); 2505 (51a); 2539 (56)
Carbon, E. 824 (14a)
Crdenas, D. 370 (16a); 469 (7); 588 (16a); 668 (7); 670 (14a); 771 (7); 1012 (16a); 1114 (14a); 1149 (37f); 1602 (16a);
254
HENDERSON
1627 (29b); 1920 (16a); 2013 (37f); 2631 (60b); 2666 (29b)
Crdenas, M. 1981 (41); 6291 (67a)
Cardiel, J. 112 (41); 1051 (37d)
Cardona, F. 922 (2)
Cardoso, S. 200 (51d)
Carvalho, A. 3465 (47); 3466 (51g); 6116 (51i); 6775 (51g)
Casari, M. 234 (51a)
Casas, F. 8103 (16a); 8136 (36); 8169 (36); 8254 (37c); 8309 (36)
Cascante, A. 151 (20); 608 (61); 1131 (14b); 1219 (38); 1463 (22a); 1957 (45b)
Castro, R. 737 (51f)
Cazalet, P. 7775 (60c)
Cern, C. 427 (21); 567 (60c); 720 (36); 1093 (60c); 1690 (36); 2264 (36); 2284 (36); 2307 (60c); 2897 (36); 2961 (67a);
2968 (27); 2981 (36); 3008 (60c); 3191 (60c); 3213 (64); 3244 (50); 3271 (37h); 3280 (8); 3379 (8); 4022 (50); 4229
(36); 4918 (60c); 4935 (31); 4998 (60c); 5052 (37h); 5133 (8); 6050 (36); 6436 (36); 6552 (60c); 6589 (36); 6630
(36); 7454 (60c); 7472 (36); 9946 (36)
Chacn, I. 59 (20); 974 (16c); 1608 (14a)
Chamas, C. 356 (56)
Chambers, K. 2561 (49c)
Chaparro, M. 40 (21)
Chavarra, M. 104 (38)
Chvez, C. 239 (22a)
Chvez, F. 643 (16a); 652 (6a); 685 (29a); 688 (36); 689 (36); 692 (6a); 693 (16a); 695 (36); 696 (36); 697 (36); 698
(36); 699 (36); 702 (29a); 705 (6a); 710 (36); 718 (6a); 745 (6a); 761 (36)
Chazdon, R. 17 (20); 18 (20); 19 (20); 20 (20); 21 (20); 25 (16a); 106 (22b); 107 (16a); 110 (22b); 119 (20); 120 (20);
132 (22b)
Choo, J. 158 (36); 213 (50); 301 (16d); 323 (60c); 351 (37d); 378 (16d); 403 (16d); 436 (16d); 437 (60c); 438 (16d); 439
(16d)
Churchill, H. 3989 (14c); 4163 (35b); 4224 (49b); 4249 (49b); 4343 (16a); 4800 (28); 5100 (67d); 5109 (14a); 5117
(14a); 5335 (14a); 5553 (14c); 5752 (28); 5800 (28); 5801 (14a); 5802 (14a); 5848 (32b); 5886 (67d); 6118 (14a);
6119 (14a); 6120 (14a); 6121 (14a); 6169 (14a); 6185 (28); 6186 (14a); 6187 (14a); 6188 (14a); 6189 (32b); 6192
(14a)
Cid, C. 7 (33); 530 (16a); 545 (60c); 842 (60c); 1479 (42); 1634 (16a); 1803 (16a); 7089 (16a); 8629 (60c)
Clark, J. 2759 (29a); 4065 (14a); 5253 (67a); 5302 (36); 5306 (67a); 5355 (60c); 5405 (67a)
Clarke, D. 339 (33); 403 (37b); 486 (2); 648 (37b); 1155 (21); 1298 (16a); 1316 (2); 1490 (2); 1899 (37b); 2205 (52);
2242 (37b); 2243 (33); 2256 (52); 2532 (16a); 2830 (16a); 3177 (37a); 3427 (16a); 3776 (2); 4167 (37b); 4336
(16a); 4390 (52); 4558 (33); 4584 (2); 4895 (2); 4896 (33); 5213 (37b); 5978 (37b); 6238 (16a); 6294 (37b); 7597
(52); 7921 (2); 7951 (16a); 8464 (52); 8723 (2); 8793 (16a); 8975 (67b); 9016 (1b); 9134 (67b); 9213 (1b); 9307
(67b); 9505 (67b); 10129 (37b); 10290 (1b); 10357 (67b); 11184 (2); 11911 (67b)
Clewell, A. 4141 (13a)
Coelho 313 (60c)
Coello, F. 43 (60c); 87 (60c)
Cogollo, A. 711 (29d); 1028 (29d); 2087 (63); 2397 (60b); 2822 (14a); 3747 (60b); 3763 (9); 3807 (25); 4915 (16a)
Colella, M. 1595 (16a); 1814 (2); 2027 (37g)
Contreras, E. s. n. (29b); 2372 (29b)
Cook, O. s. n. (13a); s. n. (49e); 11 (29b); 35 (49e); 36 (67d); 128 (49b); 135 (22a); 143 (14h); 153 (22a); 327 (67d); 727
(29b); 728 (49e); 822 (49e); 1092 (67a); 1108 (67a); 1187 (67a); 5808 (49b)
Cooper, G. 493 (35a)
Cordeiro, I. 1691 (51k)
Cordero, Z. 196 (45a); 398 (36); 517 (43)
Cornejo, F. 2801 (45a)
Cornejo, X. 4524 (30); 6359 (10b); 6537 (30)
Correa, M. 9826 (67d)
Costa, J. 670034 (47); 680082 (51a); 680089 (47)
Costa, M. 740 (1a)
255
Couvreur, T. 121 (27)

Cremers, G. 11958 (16a); 13009 (60a); 14764 (37a); 15318 (2); 15379 (66)
Croat, T. 2259 (12c); 5314 (29b); 5760 (14c); 5786 (29b); 6491 (29b); 6513 (14g); 6515 (14c); 7432 (29b); 8029 (14g);
8728 (29b); 8790 (14c); 8791 (49b); 9303 (29b); 11237 (14c); 12148 (14c); 13390 (14c); 13683 (45b); 14429 (12c);
14530 (29b); 15200 (12c); 15928 (67d); 16162 (45b); 16275 (49b); 16430 (49b); 16828 (49b); 16844 (14a); 17232
(14c); 17621 (37d); 18114 (37c); 18458 (50); 18474 (50); 18493 (50); 18524 (50); 19032 (16d); 20164 (16d); 20312
(60c); 20340 (16a); 20846 (60c); 20853 (6a); 20929 (60d); 21004 (6a); 21429 (58); 22880 (49b); 23120 (39); 24468
(29b); 24535 (29b); 25216 (14c); 25931 (39); 26056 (16a); 26462 (67d); 26469 (45b); 26688 (67d); 27121 (14a);
27369 (20); 27589 (13a); 27707 (14c); 27718 (13a); 27792 (20); 33610 (49b); 34013 (14a); 37150 (32b); 37221
(32b); 37283 (28); 37350 (35d); 37478 (14c); 37691 (29b); 37809 (14a); 40855 (49d); 41334 (67d); 43574 (49e);
43599 (22a); 43616 (22a); 44645 (35b); 48515 (32b); 48593 (67d); 48653 (28); 48869 (32b); 48961 (38); 49192
(35b); 49946 (28); 51638 (6a); 51943 (32a); 53933 (58); 56755 (14a); 58194 (53); 59311 (16a); 59452 (67b); 59717
(16a); 59756 (16c); 62795 (10a); 66475 (32b); 66643 (28); 66892 (20); 66912 (39); 67108 (14a); 67233 (67d);
67680 (13b); 67686 (14a); 67809 (28); 67831 (14a); 67969 (32b); 68733 (20); 68743 (12b); 69293 (10a); 70171
(10a); 71699 (14a); 72564 (60c); 74394 (58); 74669 (45a); 76685 (14a); 76769 (14a); 76772 (12b); 76826 (39);
78176 (32a); 79346 (14a); 79904 (24); 85555 (16a); 85561 (16); 92021 (32a); 97242 (67e); 98964 (32a)
Croizat, L. 369 (16a)
Cruxent, J. 110 (16a)
Cuadros, H. 2947 (49g); 3589 (14a); 3608 (49a)
Cuatrecasas, J. 8673 (45a); 8875 (37h); 11612 (67a); 12339 (67a); 12357 (67a); 12733 (67a); 12790 (45a); 14018 (46);
15497 (67a); 15731 (46); 15772 (7); 16042 (14e); 16981 (49h); 17160 (14e); 17428 (46); 21749 (67a); 23614 (32a)
Cuello, N. 1611 (32a)
Curran, H. 694 (67a)
Custodo Filho, A. 141 (51f); 4518 (51a)
DArcy, W. 18069 (67d)
Daly, D. 1213 (2); 1844 (21); 5823 (67g); 5901 (67g); 6057 (24); 7852 (36); 7913 (16a); 8242 (6a); 8319 (6a); 8580 (36);
8701 (60c); 8727 (60c); 8769 (6a); 8780 (60c); 8792 (8); 8803 (60c); 9119 (36); 9166 (6a); 9295 (37e); 9421 (36);
9818 (36); 9852 (37c); 9854 (16a); 10248 (36); 10423 (6a); 10466 (60c); 10550 (6a); 10582 (6a); 10716 (6a); 10947
(6a); 11194 (36); 11198 (36); 11402 (36); 11706 (60c); 11724 (6a); 11815 (36)
Daniel, Bro. 569 (32a); 4167 (32a)
da Sivla, A. 1937 (51j)
Davidse, G. 2333 (13a); 5498 (29b); 5666 (60c); 13582 (58); 16528 (16a); 16703 (58); 17316 (16a); 18521 (29c); 18605
(16a); 21617 (16a); 21746 (16a); 22157 (67a); 23349 (22a); 23485 (20); 23519 (20); 23547 (13a); 24419 (14a);
26197 (45b); 26208 (45b); 26231 (14a); 26599 (37g); 26695 (37g); 26711 (37d); 26715 (37d); 27069 (16a); 27163
(16a); 27237 (37g); 28010 (67a); 28584 (61); 28640 (67d); 28955 (45b); 29105 (61); 31106 (35a); 31112 (14a);
31287 (13a); 31333 (14a); 32026 (29b); 32290 (29b); 32441 (49e); 34255 (67d); 35682 (29b)
Davis, E. 850 (60c); 853 (50); 870 (36); 872 (36); 960 (60c); 961 (36); 1015 (37h)
Deago, J. 238 (67d)
de Benavides, O. 744 (67a); 8886 (67a); 8998 (67a)
de Granville, J.-J. 46 (2); 291 (16a); 757 (37a); 977 (33); 1348 (67i); 1892 (2); 1932 (42); 1993 (52); 1993 (60a); 2141
(2); 2210 (16); 2390 (42); 2592 (16); 3519 (66); 3774 (16); 4803 (60a); 6382 (66); 7067 (2); 7542 (21); 7543 (66);
7665 (60a); 7982 (21); 8422 (60a); 8584 (66); 8617 (37a); 8787 (60a); 8801 (21); 9096 (42); 9200 (60a); 9239 (2);
9240 (37a); 9510 (37a); 9692 (16); 10158 (2); 10212 (66); 10216 (60g); 10303 (16a); 10410 (16a); 10484 (66);
10514 (16); 10621 (21); 10883 (21); 11297 (16a); 11496 (2); 11881 (2); 11899 (60a); 11989 (21); 11994 (2); 11996
(33); 12226 (37a); 12322 (67i); 12489 (2); 12544 (60a); 12610 (33); 12611 (52); 12612 (21); 12683 (42); 12797
(37a); 12999 (16a); 13145 (16a); 13148 (60g); 13234 (60a); 13389 (37a); 13886 (66); 16838 (37a); 16897 (60a)
de Jong, W. 159 (50); 161 (36)
De La Cruz, J. 4234 (37b)
Delascio, F. 9164 (67a)
Del Carpio, C. 1779 (31)
Delgado, L. 774 (16a)
Delgado, R. 12 (22a)
Delinks, T. 311 (36)
de Lima, H. 3496 (51k)
256
HENDERSON
Delnatte, C. 1310 (60a)

Demuner, V. 2502 (19); 2632 (19); 2824 (19); 3911 (51a); 4032 (19); 4346 (56); 4449 (19); 4573 (51a)
de Nevers, G. 3785 (49b); 4058 (14c); 4457 (49b); 4462 (16a); 4482 (35c); 4711 (14c); 4721 (35c); 4792 (49b); 4864
(14c); 4959 (35c); 4960 (49b); 4961 (35c); 4974 (12a); 5037 (14g); 5080 (16a); 5120 (14a); 5152 (14c); 5367 (12a);
5381 (12c); 5427 (14a); 5433 (14a); 5434 (20); 5492 (14a); 5554 (14c); 5557 (14a); 5717 (14g); 5724 (16a); 5965
(14c); 5991 (14a); 6036 (32b); 6038 (45b); 6108 (14c); 6109 (14a); 6147 (12a); 6166 (16a); 6207 (14c); 6245 (20);
6286 (12c); 6293 (14c); 6295 (14a); 6303 (14a); 6316 (14a); 6331 (49e); 6353 (67d); 6355 (14f); 6372 (12b); 6379
(20); 6385 (14f); 6392 (35b); 6393 (14f); 6414 (16a); 6439 (49b); 6465 (16a); 6477 (16a); 6616 (12a); 6617 (35c);
6618 (14a); 6684 (32b); 6708 (16a); 6717 (20); 6718 (14f); 6728 (12b); 6730 (16a); 6732 (13a); 6745.1 (14a); 6755
(14c); 6766 (29b); 6786 (14a); 6811 (32b); 6856 (28); 6862 (13a); 6864 (35a); 6867 (13a); 6879 (13a); 6911 (14a);
6939 (16a); 6942 (12a); 6963 (35c); 7007 (14a); 7214 (14c); 7251 (35c); 7751 (16c); 7752 (14a); 7757 (57); 7783
(22a); 7789 (5); 7796 (29b); 7805 (49e); 7841 (16a); 7957 (35c); 8032 (35c); 8305 (7); 8312 (14a); 8319 (7); 8511
(67h); 8538 (16a); 8743 (14a); 8745 (67d); 8823 (15); 8838 (49e); 8854 (28); 8858 (28); 8903 (12b); 8923 (32b);
8924 (28); 8983 (14a); 8989 (14a); 10556 (38); 10601 (39); 10603 (14c); 10608 (32b); 10611 (32b); 10649 (35a);
10650 (13a); 10676 (16a)
de Paula, C. 164 (51d)
de Queiroz, L. 2830 (51j)
Devia, W. 2718 (10a)
Daz, C. 28 (36); 688 (6a); 1186 (50); 2531 (45a); 2981 (45a); 3364 (45a); 3731 (32a); 3986 (45a); 7009 (60e); 7048
(37c); 7058 (36); 7118 (36); 7120 (36); 7131 (37e); 7327 (60c); 7405 (36); 7721 (60e); 7769 (36); 7798 (45a); 7817
(32a); 7819 (45a); 7822 (45a); 8747 (32a); 8774 (45a); 8778 (32a); 9233 (16a); 9352 (16a); 9519 (60c); 10001
(45a); 10008 (45a); 10196 (32a); 10256 (45a); 10274 (45a); 10311 (32a); 10533 (45a); 10535 (32a); 10783 (32a)
Daz, S. 1530 (45a); 3150 (45a); 3231 (45a)
Daz, W. 106 (58); 450 (16a); 3195 (16a); 3828 (67b); 5414 (37b)
Dick, C. 154 (52)
Dillon, M. 4394 (67a)
Dodson, C. 7510 (62); 10073 (14d); 13804 (62); 14542 (62); 14820 (62); 14822 (14i); 14887 (67a); 15213 (67a); 15241
(67a)
Donzo, L. 8 (67d); 10 (67d)
Dorr, L. 6533 (45a); 7181 (49a); 7194 (49a); 7219 (45a); 7315 (45a); 7412 (32a); 7454 (32a); 7861 (49a); 8015 (32a);
8063 (45a); 8212 (45a); 9044 (67a)
dos Santos, R. s. n. (51a)
dos Santos, T. 1487 (19)
Doyle, C. 6 (49b); 10 (49e); 15 (45a); 21 (45a); 27 (67a)
Dransfield, J. 4860 (14a); 4863 (7)
Dressler, R. 4777 (20); 5182 (38)
Dryander, E. 3000 (29b)
Duarte, A. 3615 (19); 4037 (51e); 7316 (16)
Dudley, T. 13064 (45a); 13343 (45a)
Dueas, H. 3146 (67a)
Duivenvoorde, J. 951 (60c)
Duke, J. 242 (29b); 246 (29b); 5278 (49b); 5443 (29b); 5795 (29b); 9948 (37f); 9949 (37f); 11333 (49h); 13282 (37f);
13598 (7); 13864 (16a); 14901 (16a); 15043 (29b); 15113 (67d)
Dumont, K. 7551 (49i)
Duno, R. 1488 (67a)
Duque, A. 6035 (37d)
Dusn, P. 12136 (56)
Duss, A. 22 (67c); 3313 (67c); 4198 (67c)
Dwyer, J. 4499 (35d); 4547 (14f); 8016 (14c); 8963 (14c); 9404 (16a); 10921 (29b); 11312 (49e); 12096 (16b)
Eggers, H. 5825 (49i)
Egler, W. 46691 (16)
Eiten, G. 2056 (51a); 2122 (51j); 5196 (37d); 6397 (56); 7815 (51f); 7826 (56); 9200 (51j)
Ek, R. 599 (37b)
Ekman, E. 3781 (49f)
257
Engel, F. s. n. (45a). s. n. (45a)

Ernst, W. 1824 (49c)
Escobar, L. 4006 (14h)
Espinoza, S. 130 (33); 147 (34); 241 (37h); 290 (36)
Evans, R. 1459 (67d); 1741 (67d); 2747 (49e)
Ewan, J. 16871 (46)
Ezechias 17115 (51j)
Faber-Langendoen, D. 474 (10a); 1563 (49h)
Fanshawe, D. 573 (37b)
Farias, G. 258 (51e)
Farinaccio, M. 355 (51j)
Farias, M. 321 (16a); 491 (16a)
Farney, C. 251 (56); 265 (51f); 1752 (37d)
Fendler, A. 728 (49i); 2461 (67a); 2467 (58)
Fennell, T. 138964 (49f)
Fernandes, H. 763 (51a); 1098 (56); 1135 (51a); 1603 (56); 1833 (19); 1837 (19); 1957 (51a); 2002 (51a); 2022 (51e);
2028 (19); 2036 (19); 2037 (51f); 2046 (19); 2055 (51e); 2056 (51f); 2084 (19); 2187 (51f); 2193 (51f); 2197 (51e);
2199 (51f); 2609 (56); 2617 (51f); 2757 (51b); 2763 (51b); 2976 (19); 3073 (56); 3083 (56); 3095 (19); 3103 (19);
3107 (51a); 3204 (51j); 3218 (51a); 3219 (19); 3222 (56); 3225 (19); 3226 (51a); 3230 (56); 3253 (51a); 3254 (51a);
3290 (19); 3307 (56); 3355 (19); 3376 (56); 25848 (51a)
Fernndez, A. 1831 (2); 2673 (2); 4124 (2); 7084 (2); 7106 (16a); 7502 (16a);
Fernndez, J. 8982 (14a); 11036 (64); 16430 (45a)
Fernndez, Y. 418 (16a)
Ferreira, E. 71 (16a); 87 (36); 104 (60c); 110 (16d); 111 (8); 112 (6a); 113 (6a); 117 (36); 125 (8); 131 (16); 136 (8); 148
(37c); 161 (60c); 163 (8); 165 (8); 173 (6a); 181 (16a); 182 (60c); 185 (60c); 186 (33); 193 (36); 256 (16a); 307 (6a);
369 (36); 431 (36); 7093 (60c)
Feuillet, C. 9933 (37a); 10062 (16a); 10310 (37a)
Fiaschi, P. 8 (56); 26 (56); 149 (56); 151 (19); 163 (51a); 253 (51e); 287 (51j); 422 (56); 464 (51a); 552 (51a); 604 (51f);
613 (56); 654 (51a); 655 (56); 720 (56); 726 (51a); 913 (56); 1051 (51g); 1273 (47); 2033 (51g); 2517 (47); 2596
(47); 2834 (47); 3154 (19); 3155 (19)
Figueiredo, C. 761 (31); 802 (36)
Fleck, D. 369 (36); 373 (16d); 382 (60c); 776 (8)
Fletes, E. 1 (45b); 411 (16c); 626 (16a)
Fogg, J. 22180 (12c)
Foldats, E. 9189 (16a)
Folli, D. 1660 (19)
Folsom, J. 2329 (20); 3339 (38); 5301 (28); 6603 (12a)
Fonnegra, R. 1749 (14a); 2002 (63); 2180 (63); 2204 (49a); 2695 (29d); 2902 (16a); 3027 (25); 4057 (9); 4756 (9); 5332
(32a); 5634 (32a)
Fontoura, T. s. n. (51b)
Forero, E. 1340 (18); 1364 (60h); 1790 (49a); 3027 (67a); 3213 (14a); 4442 (14a); 7370 (49h)
Forest Department 1148 (1b); 3038 (60a); 6899 (37b); 6335 (66)
Forzza, R. 2242 (51f); 2867 (56)
Foster, M. 1421 (45a); 1673 (16a); 2315 (14h)
Foster, P. 471 (51j)
Foster, R. 4353 (16a); 4457 (36); 6787 (60c); 7305 (60c); 7326 (67a); 7327 (45a); 7632 (45a); 7640 (65); 7770 (32a);
7846 (6a); 7904 (33); 7924 (60d); 7949 (36); 7983 (6b); 8863 (16a); 8984 (45a); 9116 (32a); 9310 (6b); 9475 (64);
9566 (50); 9576 (6a); 9721 (6a); 10001 (6b); 10047 (64); 10050 (6b); 10759 (60c); 10760 (64); 10761 (64); 10768
(60c); 10791 (37c); 11904 (60c); 11961 (36); 13393 (6a)
Fothergill, M. 27 (51j)
FPR 4046 (37d)
Fraga, C. 2117 (51a)
Franco, P. 1702 (45a); 2353 (32a); 5244 (24); 5818 (37g)
Freire, E. 86 (36); 495 (60c); 2566 (34); 3381 (21); 4355 (32a)
258
HENDERSON
Fres, R. 19920 (51a); 19999 (47); 24994 (1a); 25501 (37e); 28707 (37g); 28712 (37d); 32444 (37a); 33784 (37e)
Fuchs, H. 21987 (10a); 21994 (7); 21994 (10a); 22114 (14a); 22159 (49h); 22274 (14a)
Fuentes, A. 3911 (6a); 3959 (16a); 5361 (6a); 6091 (45a); 7422 (45a); 7838 (41); 8777 (45a); 8846 (45a); 8970 (67a)
Fuentes, Z. 218 (5)
Funk, V. 8106 (41); 8272 (60c)
Furlan, A. 1495 (51a)
Galdames, C. 3383 (61); 3410 (61)
Galeano, G. 32 (32a); 67 (32a); 127 (14a); 132 (67a); 247 (45a); 248 (45a); 316 (32a); 323 (32a); 427 (10a); 447 (60h);
463 (67a); 476 (32a); 479 (32a); 485 (45a); 486 (45a); 797 (67a); 1249 (37f); 1279 (37i); 1284 (33); 1285 (37d);
1293 (37i); 1303 (36); 1304 (16a); 1305 (8); 1310 (60c); 1323 (50); 1331 (37i); 1336 (10a); 1339 (18); 1340 (37f);
1344 (36); 1353 (37e); 1354 (37i); 1355 (37d); 1466 (50); 1487 (60c); 1492 (37j); 1564 (8); 1619 (60c); 1718 (6a);
1735 (36); 1762 (36); 1770 (37j); 1811 (37j); 1821 (31); 1869 (43); 1887 (36); 1890 (36); 1954 (67a); 1956 (24);
1959 (24); 1965 (11a); 1973 (37e); 1995 (60c); 1998 (43); 2025 (37j); 2030 (36); 2034 (43); 2046 (29b); 2074 (60c);
2075 (37d); 2079 (8); 2080 (37i); 2096 (36); 2097 (6a); 2098 (60c); 2100 (60c); 2101 (60c); 2109 (37d); 2111 (16d);
2112 (16d); 2386 (32a); 2751 (60c); 3680 (63); 6122 (54); 6811 (54); 6884 (54); 7613 (49g); 7614 (49g)
Gamboa, B. 366 (45b); 708 (45b); 1055 (61)
Garca, M. 364 (16a); 419 (21)
Garca-Barriga, H. 10400 (67a); 13652 (16a); 14324 (43); 14628 (16a); 14696 (37d); 15005 (37e); 18221 (29c)
Gaviria, O. 18752 (63)
Gentle, P. 4883 (16b); 5103 (29b); 6362 (29b); 6363 (29b); 7177 (29b); 7558 (16b); 8201 (29b); 8587 (16b); 9157 (29b);
9161 (16b); 9289 (16b)
Gentry, A. 465 (13a); 3771 (29b); 6359 (49b); 7344 (18); 7449 (14c); 7883 (16b); 7987 (49e); 8036 (29b); 9090 (60c);
13856 (63); 13857 (49a); 13998 (67h); 15273 (49b); 15632 (36); 15877 (36); 16276 (41); 17438 (18); 17574 (14a);
17623 (26); 17719 (14a); 18047 (62); 18991 (60c); 20429 (36); 20474 (60c); 20921 (16a); 22019 (40); 22880 (45a);
23964 (14a); 24048 (14a); 24049 (14a); 24431 (14e); 24432 (14e); 25631 (60c); 25638 (50); 26127 (36); 26230
(60c); 26244 (37d); 26484 (67a); 27603 (60c); 27652 (16a); 27724 (36); 28657 (67a); 28666 (14a); 29080 (50);
29081 (6a); 29155 (50); 29340 (60c); 29498 (60c); 29741 (50); 29749 (60c); 30030 (16a); 31388 (60c); 31450
(16a); 31518 (16a); 31621 (60c); 31629 (37d); 31779 (41); 31861 (60c); 31924 (60c); 34989 67a); 35847 (45a);
35852 (32a); 35877 (32a); 36023 (37c); 36450 (37d); 37124 (16a); 37936 (21); 38798 (36); 39164 (60c); 39542
(50); 39548 (37d); 39570 (50); 39571 (6a); 39674 (50); 39724 (50); 40486 (14h); 40519 (14h); 40520 (14a); 40630
(46); 40633 (46); 40654 (14h); 40660 (14a); 40665 (46); 40738 (11b); 41172 (29c); 41266 (67a); 41322 (6a); 41368
(6a); 41877 (6b); 42003 (6b); 42004 (33); 42125 (6b); 42131 (37c); 42154 (8); 42609 (37d); 42649 (50); 42651 (50);
42656 (36); 42722 (50); 43160 (31); 43418 (36); 43607 (16a); 43898 (22c); 44006 (22c); 44009 (49e); 44046 (45b);
44279 (6a); 44447 (45a); 44560 (45a); 44716 (67a); 45134 (53); 45154 (45a); 45312 (45a); 45395 (53); 45403
(45a); 45512 (45a); 45513 (45a); 45538 (45a); 45902 (60c); 45997 (60c); 46133 (60c); 46547 (67b); 47660 (45a);
47797 (67a); 47806 (10a); 48188 (11b); 48419 (17); 48592 (14a); 49466 (56); 49784 (19); 49920 (47); 53252 (67a);
53356 (14e); 53400 (10a); 53400 (49h); 53684 (49h); 53744 (10a); 53925 (45a); 54033 (67a); 54536 (36); 54605
(36); 54606 (6a); 54901 (67a); 55014 (67a); 55033 (67a); 55125 (67a); 55412 (49g); 55449 (29c); 55534 (67a);
55702 (60c); 55739 (8); 55814 (36); 56144 (50); 56265 (36); 56357 (50); 56390 (37d); 56657 (10a); 56681 (14h);
57334 (7); 57498 (6a); 57500 (6a); 57521 (41); 57658 (60c); 58442 (6a); 58890 (51a); 58992 (19); 58993 (51a);
60249 (60c); 60832 (8); 60899 (60c); 60907 (36); 61490 (45a); 61835 (36); 61842 (60c); 61844 (6a); 62866 (46);
63278 (6b); 63342 (6b); 64322 (37h); 64670 (67a); 64745 (49g); 65334 (67a); 65800 (36); 65845 (60c); 66024 (36);
68666 (36); 69545 (60c); 69546 (41); 69557 (16a); 71006 (45a); 71565 (67d); 71686 (67a); 72321 (36); 72534
(67a); 74652 (45a); 75826 (14a); 76020 (67a); 76249 (49g); 76654 (60c); 76755 (41); 76827 (8); 76872 (41); 76928
(60c); 76937 (41); 77127 (6a); 78695 (16c); 78759 (16c); 78847 (67a); 78944 (14a); 79753 (49g); 80158 (67a);
80171 (67f); 80360 (65); 80753 (67a); 80978 (67e); 400003 (65)
Gerlach, G. 213 (21); 214 (6a); 216 (41); 220 (67a)
Gillespie, L. 1588 (37b); 2087 (2); 2091 (2); 2164 (2); 2377 (2)
Giraldo, J. 40 (67a)
Girn, A. 7848 (67d)
Giulietti, A.-M. 12588 (51j)
Glassman, S. 2112 (67d)
Glaziou, A. 1178 (56); 2754 (51h)
Gmez, A. 362 (14a)
259
Gmez, L. 9053 (14b); 19171 (22a); 19245 (45b); 19507 (13b); 19537 (29b); 19732 (45b); 19820 (49e); 19824 (22a);
20448 (13a); 20450 (35a); 20849 (14a); 22429 (45b); 23112 (14a)
Gmez-Pompa, A. 4466 (29b)
Gonzales, G. 19 (16a)
Gonzlez, F. 3311 (16a)
Gordillo, E. 408 (37g)
Gordon, B. 43 (13a)
Gottsberger, G. 1123671 (56); 1128671 (51a); 1521671 (56); 1522671 (56); 11291200 (14c); 12020183 (2); 14191166
(51f)
Graham, J. 2516 (37c); 2517 (36); 2519 (60c)
Grndez, C. 1692 (31)
Grant, M. 9177 (45a)
Grayum, M. 1817 (29b); 3400 (16c); 3540 (22b); 3723 (45b); 3854 (45b); 3986 (13b); 4049 (14a); 4103 (16c); 4845
(49e); 4899 (29b); 4986 (22a); 4998 (22a); 5021 (20); 5960 (14a); 5982 (14a); 6350 (22a); 6650 (20); 6698 (22b);
6721 (13a); 6895 (35a); 6927 (14a); 7025 (67d); 7124 (45b); 7539 (29b); 7540 (14a); 7624 (29d); 7645 (60h); 7689
(14a); 7703 (13a); 7704 (35a); 8005 (35a); 8108 (22a); 8359 (29b); 8732 (35a); 8734 (14a); 8991 (35a); 9185 (35a);
9190 (14a); 9383 (62); 9396 (14a); 9467 (49e); 10377 (67d); 10862 (22a); 11033 (61)
Griggs, R. 40 (67d)
Grijalva, A. 52 (22c); 434 (22c); 501 (22c); 501 (49e); 3401 (45b); 3644 (22c); 3664 (49e)
Gruezmacher, M. 43 (8); 45 (37d)
Gudio, E. 5 (60c); 552 (60c); 863 (60c)
Guilln, R. 2393 (51j)
Gunn, B. 37 (49c); 519 (49c); 534 (49c)
Gutierrez, G. 894 (16a)
Guzmn, L. 27 (14a)
Haber, W. 5329 (67d)
Hage, J. 1081 (51a)
Hahn, W. 293 (7); 1309 (49c); 3467 (67a); 3625 (37a); 3773 (42); 4220 (37b); 4304 (1b); 4394 (67b); 4395 (67b); 4397
(67b); 5000 (67a); 5454 (67b); 5799 (37b)
Hall, F. 592 (2)
Hamilton, C. 680 (14c); 1084 (14g); 1451 (29b); 2652 (38); 2982 (38); 3045 (14a); 3093 (32b); 3732 (32b); 3749 (32b);
4069 (14c)
Hammel, B. 796 (20); 1111 (49b); 1685 (13a); 1789 (14a); 2116 (28); 2120 (32b); 2398 (35b); 2615 (12b); 2655 (49b);
3074 (32b); 3107 (20); 3184 (12c); 3186 (14a); 3459 (39); 3496 (14c); 4112 (14a); 4139 (20); 4166 (35b); 4614
(14a); 4616 (38); 5602 (14c); 6134 (28); 6225 (28); 7204 (20); 7523 (32b); 7524 (32b); 11622 (29b); 11765 (13a);
12082 (35a); 12480 (35a); 12804 (49e); 13851 (67d); 13862 (45b); 13864 (67d); 13875 (5); 14044 (14g); 14234
(22a); 14498 (35a); 14769 (14c); 14803 (49b); 14829 (49b); 14832 (14g); 15594 (49e); 15954 (36); 16761 (16c);
17801 (49e); 17810 (22a); 19829 (45b); 20151 (14a); 20326 (14g); 20450 (14a); 20503 (49b); 21447 (60a); 21636
(2); 21674 (37a); 21757 (60a)
Hammer, N. 83 (35a)
Hampshire, R. 892 (28); 922 (28)
Hansen, B. 7786 (62)
Harley, R. 17852 (47); 17853 (51a); 19613 (51j); 20187 (51j); 24894 (51j); 26143 (51j)
Harling, G. 3260 (60c); 3569 (36); 3577 (6a); 3600 (60c); 3762 (60c); 3892 (67a); 24557 (36)
Harmon, W. 2068 (29b)
Hartman, R. 12113 (7); 12489 (14a)
Hassler, E. 4715 (51j)
Hatschbach, G. 2975 (56); 7794 (19); 8066 (56); 16014 (19); 16035 (56); 16319 (19); 18642 (19); 35344 (51j); 35450
(56); 35907 (51j); 46160 (51j); 47799 (19); 52233 (51a); 52769 (56); 65025 (51j); 67206 (51j)
Haught, O. 1850 (7)
Hawkins, T. 385 (67d); 492 (67d); 1195 (49e)
Hayes, S. 98 (14c); 530 (14g); 896 (13a); 897 (49b)
Heinrichs, E. 375 (36); 501 (16a)
Henao, G. 348 (67a)
260
HENDERSON
Henao, J. 288 (24); 289 (24); 292 (32a)

Henderson, A. 9 (37g); 16 (16a); 30 (67b); 31 (67b); 39 (37g); 48 (14a); 55 (13a); 56 (29b); 61 (16a); 64 (29b); 69 (14a);
75 (14a); 90 (14g); 95 (16a); 102 (49b); 110 (49a); 117 (6a); 120 (67a); 152 (24); 157 (24); 158 (24); 187 (1a); 189
(37j); 191 (16a); 195 (36); 200 (33); 201 (16a); 203 (37j); 230 (37d); 250 (16a); 301 (60c); 302 (60c); 303 (60c); 482
(1a); 524 (45a); 528 (6a); 529 (45a); 530 (67a); 532 (45a); 538 (65); 633 (37d); 639 (1a); 643 (37j); 645 (1a); 649
(37a); 655 (1a); 659 (37j); 662 (60c); 664 (37a); 670 (37d); 678 (37d); 710 (16a); 717 (49b); 725 (14g); 816 (36);
818 (16a); 819 (60c); 828 (6a); 829 (36); 835 (37d); 836 (16a); 843 (8); 845 (50); 850 (60c); 851 (60c); 853 (6a);
860 (16a); 862 (36); 863 (8); 866 (60c); 877 (33); 878 (60c); 879 (16a); 880 (16a); 885 (60c); 886 (8); 889 (60c);
890 (60c); 891 (16a); 893 (2); 976 (37g); 996 (2); 1009 (67b); 1043 (60c); 1048 (37d); 1050 (16a); 1053 (37j); 1055
(37a); 1056 (37d); 1066 (60c); 1075 (37a); 1079 (1a); 1100 (31); 1102 (31); 1105 (60c); 1106 (8); 1108 (60c); 1110
(36); 1112 (60c); 1115 (36); 1121 (36); 1122 (37d); 1128 (8); 1132 (60c); 1143 (16); 1149 (37d); 1152 (1a); 1161
(37d); 1163 (37a); 1181 (49f); 1512 (37d); 1517 (37d); 1518 (37d); 1520 (60c); 1521 (33); 1523 (36); 1528 (60c);
1529 (37d); 1531 (60c); 1533 (16a); 1536 (36); 1538 (33); 1550 (16a); 1553 (33); 1554 (37e); 1557 (37j); 1558
(37j); 1578 (60c); 1579 (60c); 1580 (60c); 1585 (43); 1586 (16a); 1595 (60c); 1597 (60c); 1625 (49i); 1632 (16a);
1633 (41); 1634 (37c); 1636 (50); 1637 (6a); 1638 (60c); 1649 (6a); 1651 (36); 1652 (6a); 1661 (8); 1668 (60c);
1673 (16a); 1679 (16); 1680 (36); 1682 (60c); 1684 (16); 1687 (37d); 1690 (8); 1700 (60c); 1705 (16a); 1707 (36);
1712 (37d); 1803 (61); 1808 (35a); 1810 (14a); 1813 (57); 1816 (16c); 1817 (16c); 2039 (35d); 2042 (14a); 2043
(32b); 2044 (28); 2052 (12b); 3004 (29a); 3005 (29a); 3012 (6b); 3020 (49e); 3021 (29b); 3024 (16a); 3028 (12b);
3029 (14a); 3043 (14a); 3044 (14a); 3045 (14c); 3047 (13a); 3051 (16a); 3053 (49b)
Henkel, T. 500 (2); 1074 (21); 1359 (1b); 1530 (67b); 3302 (52); 4301 (1b); 4485 (67b); 4701 (2)
Heringer, E. 15 (51j); 16951 (51j); 18328 (51j); 8589783 (51j)
Hernndez, H. 1171 (49e); 1376 (29b)
Hernndez, J. 594 (60b)
Herrera, G. 1048 (13a); 1242 (14b); 1652 (14g); 2487 (22a); 2785 (61); 2789 (61); 3427 (45b); 4371 (16c); 4372 (16c);
4820 (57); 8535 (61)
Herrera, H. 752 (14g); 938 (29b); 973 (14g); 1327 (14a); 1333 (14g); 1401 (14a); 1483 (49b)
Heyde, E. 4656 (49d)
Hill, S. 24065 (49c)
Hind, D. 50061 (51j)
Hodel, D. 1240 (38); 1242 (28)
Hodge, W. 307 (49c); 1430 (67c); 1987 (49c); 6079 (6a); 6999 (14a); 7002 (63); 7014 (63); 7074 (63)
Hodges, V. 242 (36)
Hoehne, F. 17386 (51h); 17391 (56)
Hoff, M. 6686 (42); 6700 (42); 6773 (16a); 7062 (16a)
Hoffman, B. 824 (37b); 1403 (37b); 1577 (37b); 3309 (21); 3320 (1b); 3686 (2); 3905 (37b); 3993 (37b)
Holdridge, L. 2149 (49f); 5112 (35a)
Holm, R. 74 (49e); 907 (14g); 919 (13a); 962 (13a)
Holm-Nielsen, L. 19808 (36);19852 (60c);19928 (31);20069 (60c);21308 (64);21316 (6a);21601 (6a);21664 (8);21809
(60c);21896 (36);22073 (60c);22124 (34);22128 (60c);22157 (34);22224 (60c);22264 (36);22267 (60c);22461
(60c);22479 (36);25437 (14e);25841 (16a);26565 (45a)
Holst, B. 2368 (16a); 5293 (49e)
Homeier, J. 1977 (27)
Hoover, W. 2489 (30); 3764 (30)
Huaman, E. 24 (8)
Huashikat, V. 674 (36); 1023 (50); 1277 (36); 1868 (34); 2009 (36)
Huber, W.149 (57)
Huber, O. 173 (59); 6256 (49a); 8921 (67b); 9038 (67b); 12602 (67b); 12796 (67b)
Huertas, G. 586 (67a)
Huft, M. 1617 (16a); 1670 (49b); 1688 (14g)
Hurtado, F. 1033 (36); 1547 (60c); 2083 (36); 2291 (36); 2659 (6a)
Hutchinson, P. 8954 (51a); 8955 (19); 8991 (51a)
Idrobo, J. 197 (24); 709 (29b); 1618 (67a); 2498 (21); 9658 (45a); 10519 (24); 11308 (16a)
INBIO 6 (67d)
Induni, G. 237 (16c)
261
Irwin, H. 5422 (51j); 6276 (51j); 8851 (51j); 12479 (51j); 15633 (51j); 19233 (51j); 21810 (51j); 24838 (51j); 28195
(56); 47282 (60g); 47351 (16); 47455 (37a); 47588 (16a); 47627 (52); 47631 (16); 47707 (52); 47714 (2); 47795
(16); 48082 (16); 55587 (2)
Itaip Binacional 940 (51j)
Ivey, C. 68 (5); 261 (45b)
Jangoux, J.1615 (2); 10189 (29c); 85001 (60c); 85029 (36); 85030 (60c); 85050 (60c); 85070 (16); 85075 (36); 85076
(8)
Janovec, J. 1281 (14a); 1369 (60c); 1443 (60c); 1486 (60c); 1499 (67e); 1502 (67e); 1552 (67a); 1882 (60c); 2163 (41);
2370 (60c)
Jansen-Jacobs, M. 368 (37b); 823 (21); 824 (33); 1508 (2); 1521 (33); 1730 (16a); 2384 (37b); 2386 (2); 2944 (37b);
2945 (37b); 2946 (2); 5438 (37b); 5727 (16a); 5784 (16a); 6343 (60a)
Janzen, D. 11089 (14a)
Jaramillo, J. 4242 (29a); 6832 (60c); 6874 (60c); 12622 (64); 13953 (34); 14690 (16a); 14722 (60c); 30701 (60c)
Jaramillo, N. 275 (36); 499 (36); 1099 (36)
Jaramillo, R. 438 (34); 7141 (24); 7332 (45a)
Jardim, J. 845 (51a); 1098 (51a); 2961 (47)
Jtiva, C. 231 (14a)
Jimnez, Q. 891 (14a)
Jorgensen, P. 61362 (45a)
Jouvin, P. 145 (51f)
Juchum, F. 78 (51j)
Juncosa, A. 601 (16a); 649 (49b); 660 (49b); 739 (60b); 1041 (32a); 1228 (14a); 1251 (7); 1444 (14e); 1547 (10a); 1641
(14a); 1694 (14a); 1723 (7); 1726 (18); 1770 (18); 1869 (63); 1870 (14a); 1908 (14a); 2162 (67a); 2168 (67a)
Kahn, F. 578 (1a); 1681 (29a); 1697 (60c); 1715 (37d); 1757 (60c); 1886 (60c); 1892 (8); 1893 (8); 1894 (50); 1899 (33);
1900 (33); 2020 (50); 2064 (60c); 2117 (37c); 2119 (16a); 2133 (41); 2152 (60c); 2154 (60c); 2157 (37c); 2182
(60c)
Kajekai, C. 300 (60c)
Kallunki, J. 535 (47); 581 (47); 669 (51a)
Karsten, H. s. n. (45a); s. n. (58); s. n. (58)
Katan, T. 76 (36)
Kawasaki, L. 974 (56)
Kayap, R. 210 (60e); 361 (55); 552 (34); 797 (50); 1212 (55)
Kellerman, W. 7130 (29b)
Kennedy, H. 660 (49b); 1763 (14c); 1803 (49b); 2791 (14g); 2850 (29b)
Kernan, C. 521 (16c); 969 (16c)
Kiem, S. 603 (16b)
Killeen, T. 4120 (67a)
Killip, E. 8787 (24); 11137 (67a); 14866 (16a); 15336 (49a); 16044 (45a); 23006 (6a); 23860 (36); 24700 (67a); 24702
(36); 25821 (32a); 26170 (33); 26190 (60c); 26247 (6b); 26414 (33); 26430 (16a); 26431 (64); 26455 (6b); 26532
(16a); 26594 (16a); 26722 (6b); 26987 (37d); 27295 (36); 27360 (36); 27654 (31); 27977 (37c); 28026 (50); 28142
(36); 28145 (36); 28541 (50); 28895 (36); 29373 (36); 29433 (36); 29621 (8); 29673 (36); 29951 (37d); 33623 (7);
33942 (67a); 34274 (21); 37398 (33); 37437 (2); 37802 (58)
Kinloch, J. 55 (16b)
Kirizawa, M. 935 (19)
Kirkbride, J. 649 (39); 1172 (63); 1349 (7); 1884 (45a); 2375 (67a); 2410 (45a); 4194 (45a)
Klug, G. 448 (36); 448 (36)
Knab-Vispo, C. 531 (2)
Knapp, S. 893 (12c); 1203 (12c); 1509 (61); 1712 (16a); 1985 (12b); 2577 (38); 2864 (37b); 2998 (49b); 3103 (7); 3161
(49b); 3460 (14a); 3464 (20); 3800 (35b); 3804 (20); 3810 (35b); 3812 (38); 4209 (32b); 4342 (38); 4696 (14a);
4837 (32b); 4987 (14a); 5078 (28); 5118 (28); 5242 (14c); 5308 (35d); 5384.1 (14c); 5387 (39); 5618 (28); 6041
(29b); 6057 (67d); 7113 (36); 7616 (60f); 7673 (60c); 7692 (60e); 7750 (45a); 7925 (37c); 8133 (36)
Knudsen, J. 519 (60c); 525 (60c); 528 (60c); 583 (34); 585 (60c); 614 (49b); 615 (35c); 626 (14a); 633 (32b); 635 (32b);
641 (32b); 644 (28); 646 (28); 647 (28); 648 (32b); 651 (35a); 660 (35a); 664 (45b); 1014 (35a); 9578 (62); 9618
(40); 9625 (40); 9626 (60c); 9627 (60c); 9628 (60c); 9653 (60c); 9654 (60c); 9655 (60c); 9710 (36); 9713 (6a); 9722
262
HENDERSON
(60c); 9723 (60c); 9733 (50); 9734 (50)

Koczicki, C. 205 (51a)
Kollmann, L. 4170 (19); 5401 (56); 6214 (51k); 6324 (19); 8729 (51f); 9321 (19); 9579 (56); 9590 (51k); 10630 (19)
Koortjvar, B. s. n. (56)
Koptur, S. 331 (67d); 334 (67d)
Koyama, T. 7326 (21)
Krapovickas, A. 23181 (56); 23246 (56); 36983 (51a); 38462 (56); 46029 (51j)
Kress, J. 913251 (2)
Krukoff, B. 4543 (60c); 5361 (36); 6082 (2); 6772 (37e); 7157 (37j); 7158 (43); 8144 (36); 10808 (6a); 10836 (6a);
10982 (21); 10983 (16a); 11154 (45a)
Kuhlmann, J. 141 (51d); 1231 (33); 1236 (43); 1237 (37d); 3790 (51j)
Kujikat, A. 98 (60e); 159 (50); 332 (55); 434 (55)
Kunkumas, P. 215 (60c)
Kuntze, O. s. n. (6a)
Laegaard, S. 55213 (45a)
Langlois, A. s. n. (22b)
Lanna, J. 476 (37g); 573 (56); 575 (51a); 576 (19); 584 (51f); 590 (56); 884 (56); 888 (56); 1129 (19); 1252 (51b); 2174
(37g)
La Rotta, C. 594 (7); 619 (18)
Lau, P. 3 (49a)
Lawesson, J. 43328 (31); 43336 (31); 43496 (50); 44367 (16a); 44487 (36)
Le Fiell, C. 4 (37d)
LeDoux, D. 2580 (14c)
Lehmann, F. 5289 (67e); 7233 (32a); 8957 (46)
Leito Filho, H. 33102 (19); 34584 (19)
Leitman, P. 79 (51k)
Lellinger, D. 1007 (45b)
Lent, R. 34 (16a); 2137 (13a); 3335 (22a)
Len, H. 1293 (14a)
Leoni, L. 1496 (56)
Le Prieur, F. s. n. (60a); s. n. (60a); s. n. (60a)
Leveau, J. 83 (37c); 190 (37c)
Lewis, W. 211 (29b); 2162 (35a); 10114 (37d); 10263 (36); 10332 (60c)
Licata, A. 249 (32a); 658 (32a)
Liebman, F. 10804 (49e)
Liesner, R. 284 (29b); 791 (14f); 815 (14c); 2861 (29b); 3216 (13b); 5978 (37b); 8224 (49a); 8369 (49a); 9341 (16a);
9779 (49a); 9858 (45a); 10017 (45a); 10570 (16a); 10943 (49a); 10991 (16a); 11368 (16a); 11799 (45a); 12349
(49a); 13734 (58); 13995 (52); 13999 (2); 14828 (22a); 15028 (49b); 15049 (13a); 15262 (49e); 15309 (49e); 17331
(16a); 17554 (16a); 17569 (37g); 17684 (67b); 17843 (37g); 17939 (67b); 18846 (16a); 19184 (33); 19594 (2);
21630 (21); 22428 (67b); 24278 (16a); 24531 (37g); 24765 (67b); 24767 (67b); 25096 (67b); 25180 (67b); 25587
(37g); 25833 (37g)
Lindeman, J. 389 (16a); 4184 (2)
Ling, F. 12 (35a)
Listabarth, C. 112190 (36); 116589 (36); 1121289 (29a); 11141188 (6a); 11120594 (37g); 12120594 (16a); 12240594 (2)
Lleras, E. 16642 (37e); 16916 (36); 17168 (36); 17313 (60c); 17401 (60c)
Lloyd, F. 930 (49c)
Loiselle, B. 101 (20)
Ljtnant, B. 13437 (36); 12261 (67a); 13534 (60c)
Lombardi, J. 348 (51a); 1550 (51a); 1846 (51j); 5156 (19); 5491 (47)
Loomis, H. 3 (13a); 10 (35a)
Lpez, R. 1666 (21); 2146 (21); 3603 (29b); 3606 (21); 4264 (43); 7020 (32a)
Lourteig, A. 2374 (19)
Lowrie, S. 690 (16a)
Lozano, G. 2444 (45a); 2458 (67a); 2639 (67a); 4902 (67a); 5923 (14e); 7273 (9); 7696 (24)
263
Lozano, P. 606 (45a); 882 (67e)

Luederwaldt, H. 12230 (19); 12235 (56); 12238 (51a)
Luetzelburg, P. s. n. (19); s. n. (19); 20045 (56); 22278 (37d)
Lundell, C. 2691 (49e); 6823 (29b)
Luteyn, J. 6609 (45a); 8213 (58); 8321 (67a); 8518 (6a); 8571 (6a); 8679 (36); 8719 (36); 13746 (45a)
Lyra-Lemos, R. 1871 (47)
Maas, P. 3285 (19); 3290 (56); 3526 (37b); 6668 (16a); 6846 (37g); 11065 (16a); 12704 (60c)
Mabberley, D. 1851 (49b)
Macbride, J. 4775 (45a); 4867 (67a)
Macedo, A. 3346 (51j)
Macia, M. 3986 (6a); 4631 (21)
Madison, M. 5691 (1a)
Madrigal, M. 81 (14a)
Magnago, L. 1542 (51a)
Maguire, B. 22939 (37b); 24094 (2); 24095 (33); 24117 (52); 24154 (2); 24166 (60a); 28800 (43); 29347 (16a); 29361
(37g); 37156 (67b); 39058 (60a); 40822 (52); 46740 (2); 60530 (67b); 65662 (67b)
Malagn, W. 8 (6a); 11 (37h); 26 (45a); 48 (64); 58 (37h); 70 (40); 74 (34)
Marcato, A. 1 (51a); 4 (19); 38 (56); 78 (56); 112 (51j); 123 (56); 130 (51j); 170 (51j); 201 (56); 211 (56); 339 (51a); 345
(19)
Marqueta, R. 2182 (51f)
Martn, S. 818 (35a); 991 (22b)
Martinelli, G. 2226 (51e); 2253 (19); 4032 (51d); 5014 (51e); 6727 (51a); 7474 (51j); 11792 (51k); 12042 (51k); 12246
(51k); 13314 (19)
Martnez, E. 6981 (29b); 8270 (29b); 8800 (49e); 12243 (29b)
Martins, H. 333 (56)
Mathias, M. 5113 (6a)
Matuda, E. 3477 (49e); 16384 (29b); 16820 (29b); 18568 (49d)
Maxon, W. 6825 (49b); 6903 (13a)
Mayo, S. 1013 (51a)
McDaniel, S. 10950 (36); 11032 (60c); 16892 (37d); 27585 (50); 30253 (31)
McDonagh, J. 508 (49b); 510 (49b)
McDowell, T. 425 (13a); 2696 (37b); 2708 (21); 3367 (37b); 3441 (1b); 3487 (37b); 4268 (37b); 4398 (52); 4479 (37b);
4914 (1b)
McPherson, G. 7195 (13a); 9388 (32b); 11448 (14a); 11485 (29b); 13201 (67a)
Meave J. 1051 (49e)
Meerow, A. 956 (51j); 1088 (5)
Meier, W. 377 (67a); 3401 (44); 3405 (58); 3409 (58); 4469 (49a); 4543 (49a); 4605 (67a); 4664 (49a); 4782 (49a); 5183
(4); 5654 (49a); 5716 (49a); 5848 (45a); 6466 (67a); 6565 (49a); 7176 (49a); 7235 (67j); 7907 (59); 8582 (67a);
9228 (49i); 9718 (4)
Mejia, K. 642 (16a)
Mejia, R. 130 (29b); 3542 (24)
Mello-Silva, R. 829 (51a); 844 (19); 851 (51h); 917 (51a); 961 (19); 1569 (51a); 1570 (51a); 1622 (56); 1667 (51j); 3015
(51j); 10329 (51j)
Mendoza, H. 2747 (21)
Menezes, N. 4767 (51j)
Merino, B. 4777 (45a)
Mexia, I. 4664 (56); 6135 (50); 7027 (67a); 7763 (32a); 7764 (32a)
Milanowski, D. 176 (45a)
Miller, J. 744 (14c); 1111 (14g); 1742 (16a)
Milliken, W. 460 (2); 706 (37b); 711 (16a)
Molina, A. 14979 (16a); 17666 (14g); 17683 (13a); 20379 (67d)
Mondragn, M. 55 (67a); 63 (67a)
Monsalve, M. 314 (10a); 337 (10a); 350 (7); 400 (14h); 421 (14h); 473 (10a); 866 (49h); 987 (49h); 1040 (16a); 1066
(10a); 1272 (14e); 1385 (46); 1417 (46); 1447 (14a); 3093 (14h); 1080 (49h)
264
HENDERSON
Monteagudo, A. 3845 (67a); 4085 (6b); 4185 (36)

Montenegro, W. 54 (67f)
Moore, H. 6534 (13b); 6536 (13b); 6538 (16a); 6566 (22b); 6586 (22b); 6592 (29b); 6607 (22b); 6612 (22b); 6675 (45b);
6695 (22a); 6722 (22a); 6727 (22a); 8219 (16b); 8350 (67a); 8355 (6a); 8428 (37d); 8434 (60c); 8450 (50); 8474
(60c); 8497 (31); 8503 (50); 8520 (37c); 8530 (50); 8577 (6a); 8592 (6a); 9455 (46); 9469 (16a); 9472 (7); 9530
(1a); 9534 (37a); 9543 (16a); 9545 (37j); 9553 (37a); 9716 (67b); 9839 (58); 9840 (59); 9874 (16a); 9875 (7); 10126
(35a); 10188 (16a); 10207 (14e); 10222 (63); 10532 (35b)
Mora, E. 1681 (14a)
Moraes, M. 295 (16a); 676 (6a); 833 (67a); 834 (67a); 839 (45a); 848 (45a); 849 (67a); 1864 (6a); 1865 (16a); 1866 (41);
1872 (45a)
Moraga, C. 331 (67d); 544 (13a); 761 (13a); 831 (14b); 951 (16a)
Morales, C. 1433 (60c); 1686 (67a); 1693 (67a)
Morales, J. 5707 (67d); 10297 (35a)
Moreno, L. D. 19 (45a)
Moreno, L. R. 17 (51j); 34 (41); 35 (6a); 43 (16a); 51 (41); 58 (37c); 64 (6a); 78 (67a); 96 (45a); 99 (67a); 103 (41); 105
(60c); 108 (6a); 109 (36); 114 (67a); 124 (6a); 125 (21); 126 (50); 141 (51j); 155 (16a); 161 (37e); 166 (16a); 180
(33); 196 (60c); 197 (37c); 198 (41); 200 (37e); 204 (37d); 211 (37c); 223 (21); 224 (6a); 227 (6a); 237 (37d); 253
(60c); 254 (60c); 257 (37c); 305 (50); 312 (37d); 346 (21)
Moreno, P. 7480 (67d); 7515 (22c); 7990 (67d); 12994 (14g); 15157 (14g); 18142 (22c); 20203 (22c); 23083 (14g);
23704 (29b); 23881 (13a); 13033 (49e); 14819 (13a); 15013 (13a); 15126 (13a); 17296 (29b); 20467 (49e); 20523
(49e); 20604 (13a); 21021 (49e); 25573 (13a)
Mori, S. 1943 (14c); 2120 (13a); 2235 (16a); 2496 (14c); 2620 (14a); 2811 (49b); 2895 (16a); 3173 (14a); 3307 (49b);
3909 (20); 4007 (14a); 4119 (12a); 4804 (14a); 5134 (49b); 6433 (14a); 6561 (14c); 6577 (35d); 7500 (32b); 7933
(12c); 8154 (37b); 8155 (37b); 8156 (37b); 8628 (2); 8937 (60a); 9012 (37e); 9188 (60c); 9910 (51a); 10181 (47);
10517 (51a); 10661 (47); 11020 (47); 11022 (51i); 11746 (47); 11759 (51g); 12106 (47); 12538 (51j); 12814 (47);
12822 (47); 15552 (37a); 17204 (16); 17265 (37a); 17607 (16); 17611 (52); 19271 (1a); 21591 (37a); 21835 (37d)
Mullins, J. 1 (51j)
Murphy, H. 88 (31)
Murray, N. 3 (45b)
Mutchnick, P. 1514 (37b); 820 (16a); 844 (2); 1064 (2); 1515 (33)
Nadruz, M. 1829 (51d)
Nakajima, J. 266 (51j)
Narvez, E. 925 (62); 954 (62)
Navas, A. 514 (7)
Nee, M.11252 (39); 17842 (58); 28387 (13a); 28412 (22c); 28420 (14a); 30640 (67b); 30885 (16a); 31617 (41); 34335
(16a); 34372 (16a); 34436 (41); 34903 (33); 34994 (16a); 42341 (60c); 42897 (60c); 49528 (36); 51564 (36)
Neill, D. 1602 (49e); 1861 (22c); 1915 (13a); 3539 (13a); 3800 (29b); 3828 (22c); 6677 (60c); 6770 (34); 6784 (36);
6862 (36); 7187 (60c); 8073 (36); 8183 (64); 8195 (60c); 8251 (64); 8622 (36); 8832 (60c); 8833 (60c); 8876 (60c);
8984 (60c); 9356 (6a); 9561 (64); 9635 (67e); 10560 (60c); 10563 (67a); 10606 (36); 11229 (60c); 13910 (14a);
13961 (30); 14058 (36); 14058 (60c); 14982 (32a); 15024 (32a); 15026 (67f)
Nelson, C. 9145 (13a); 9259 (16a)
Neruyi, E. 386 (51f)
Nicolson, D. 1920 (49c)
Nielsen, V. 43 (16c)
Noblick, L. 3209 (51j); 3334 (51j); 4404 (47); 4692 (51a); 4715 (51a); 4726 (51g); 4747 (51a); 4748 (51a); 4749 (47);
4755 (51a); 4756 (51a); 4761 (47); 4769 (47); 4778 (51a); 4789 (51a); 4790 (19); 4799 (56); 4875 (19); 4926 (51j);
4940 (49a); 4949 (67a); 4952 (51a); 5001 (51g); 5008 (51j); 5011 (37a)
Nez, P. 8787 (67a); 9826 (60c); 9839 (16a); 10590 (6a); 11939 (45a); 11993 (45a); 12242 (6a); 14187 (67a); 23326
(45a); 23383 (45a); 23937 (29a)
Oersted, A. 6564 (49b)
Ohba, T. 127 (67a); 133 (45a); 588 (21)
Oldeman, R. 161 (16a); 3164 (52)
Oliveira, E. 3861 (37a); 4676 (37a)
llgaard, B. 9035 (60c); 9236 (60c); 34750 (60c); 34754 (34); 35332 (60c); 39183 (60c); 57518 (30); 57626 (30); 74507
265
(67a); 90560 (45a); 98478 (60c)

Orjuela, M. s. n. (45a)
Ortega, F. 2567 (67a)
Ortiz, F. 779 (13a)
Oviedo, E. 3 (21)
Pabn, M. 62 (52)
Paixo, J. 470 (47)
Palacios, W. 692 (34); 702 (37h); 728 (36); 1556 (36); 1557 (60c); 2208 (36); 2361 (60c); 2440 (60c); 4080 (36); 4114
(36); 4143 (67a); 4223 (60c); 5054 (45a); 5110 (60c); 5505 (64); 5547 (64); 5582 (36); 6067 (67a); 6501 (34); 7612
(34); 7674 (60c); 7683 (8); 8223 (64); 8595 (60c); 8850 (37h); 8960 (64); 9136 (34); 9137 (60c); 9270 (37h); 9414
(60c); 9424 (6a); 9602 (45a); 11191 (14a); 11209 (14e); 12553 (67a); 15664 (34); 15724 (60c); 15762 (21); 15777
(36)
Paniagua, N. 5870 (45a)
Pardini, R. 1 (51a); 2 (19); 10 (37d); 11 (8); 13 (16d); 14 (60c); 15 (6a); 17 (60c); 19 (60c); 23 (8); 25 (36); 28 (60c); 29
(16); 32 (37j); 35 (31); 44 (37d); 45 (8); 48 (16d); 49 (36); 50 (16a); 51 (37e); 52 (60c); 53 (37j); 56 (60c); 60 (31);
63 (33); 75 (19); 76 (51a)
Paula-Souza, J. 5818 (51d)
Pavn, J. s. n. (29a)
Pedersen, H. 97601 (14d); 97607 (60c); 97611 (34); 97616 (60c); 97619 (33); 97639 (67a); 97649 (60c); 104007 (8);
104019 (16a); 104080 (67a)
Peixoto, A. 3140 (19); 3499 (51a); 3518 (56)
Pea, F. 29 (36); 43 (36)
Pea, M. 1061 (49e)
Pennell, F. 3413 (24); 4235 (14a); 7642 (45a); 8787 (24); 9057 (24); 9314 (45a)
Pennington, T. 30 (14i)
Pereira, B.1555 (51j)
Prez, A. 20 (33); 790 (42)
Persaud, R. 93 (67b)
Peyton, B. 1454 (67a); 1496 (67a)
Philcox, D. 4606 (51j)
Philipson, W. 1464 (21); 1962 (21); 1979 (49a); 2327 (21)
Phillips, J. 469 (37i)
Picado, A. 133 (14a); 358 (45b)
Pinard, M. 807 (36); 808 (16a); 837 (37c); 859 (37c)
Pinheiro, R. 141 (47)
Pinilla 667 (16d)
Pinkus, A. 140 (67b)
Pipoly, J. 3770 (13a); 3816 (16a); 4706 (49e); 4892 (29b); 4917 (49e); 5112 (22c); 5246 (22c); 5307 (29b); 5308 (13a);
6111 (22c); 6144 (22c); 11049 (21); 12325 (31); 12444 (8); 12675 (60c); 12873 (6a); 12892 (36); 13001 (60c);
13040 (60c); 13059 (36); 13292 (37d); 13434 (37d); 13454 (60c); 13739 (36); 13908 (37d); 13922 (60c); 13946
(60c); 14144 (8); 14478 (16d); 14496 (37d); 14930 (36); 15628 (60c); 15638 (37d); 16576 (67a)
Pirani, J. 1258 (51j); 2305 (51j); 2926 (47); 3116 (19); 3118 (51a); 11712 (51j)
Pires, J. 50395 (16); 50396 (52); 50759 (16); 51397 (16); 52221 (37a)
Pitman, N. 358 (60c)
Pittier, H. 4411 (16a); 4423 (32b); 8030 (58); 9109 (58); 10012 (67a); 11371 (58); 13915 (49a); 13840 (59)
Plowman, T. 4388 (34); 5062 (6a); 5937 (50); 9586 (2); 11461 (50); 11539 (36); 11677 (50); 12197 (60c); 12235 (2);
12236 (33); 12247 (36); 12419 (37d); 13366 (67a); 13653 (67b)
Polak, M. 136 (52)
Poliquesi, C. 630 (56)
Poll, G. 228 (49e)
Poole, J. 1652 (52)
Porter, D. 4389 (14c); 4741 (35a); 4808 (16a)
Pounds, W. 339 (45b)
Prance, G. 1598 (2); 1675 (2); 1831 (66); 1992 (52); 2239 (37a); 2383 (31); 2766 (31); 2801 (36); 2838 (16); 4559 (16a);
266
HENDERSON
4989 (16a); 5529 (37d); 5803 (16a); 5980 (37e); 7319 (6a); 7596 (37d); 8533 (60c); 8536 (16a); 9362 (2); 10036
(21); 10036 (33); 10266 (16a); 11835 (36); 12093 (60c); 13488 (31); 14667 (60c); 15311 (60c); 16363 (31); 16491
(33); 16492 (16a); 16493 (36); 17781 (1a); 21691 (37e); 23838 (37d); 23847 (60c); 23909 (36); 24055 (60c); 24572
(37d); 25489 (16a); 25503 (60c); 26519 (37a); 29195 (67b); 29451 (16a); 30213 (16a); 59110 (51j)
Prvost, M. 2417 (2)
Priess, J. 138 (16a); 140 (16a)
Proctor, G. 26927 (13a); 26968 (16a); 32326 (14g)
Quelal, C. 175 (30); 425 (67a)
Quesada, J. 900 (16a); 1563 (67d)
Quevedo, R. 2621 (51j); 2545 (51j)
Quinet, A. 392 (19); 1043 (51a)
Quipuscoa, V. 340 (50); 405 (45a)
Quizhpe, W. 868 (32a); 1338 (60c); 2222 (32a); 2471 (32a)
Rabelo, B. 1861 (2); 2310 (16); 2393 (16)
Rainer, H. 11.8888 (6a); 4988 (29a); 7988 (60c); 133288 (45a); 217988 (36); 1321988 (36); 1330188 (45a); 1418188
(37c); 1625188 (36); 2131888 (33); 2210988 (60d); 2214988 (45a); 2223888 (37c); 2314988 (45a); 2513988 (45a)
Rambo, B. 42217 (56); 47141 (51a)
Ramrez, C. 1 (14a)
Ramrez, J. 85 (29d); 1196 (9); 1361 (9); 4919 (64); 5034 (64); 5146 (45a); 5401 (64)
Ramos, J. 160 (51j); 21825 (1a)
Rangel, O. 2021 (45a); 2173 (45a); 2462 (32a); 5334 (45a)
Read, R. 2003 (49c); 2021 (49c); 3615 (56); 8115 (20); 8138 (20); 74212 (49c)
Redden, K. 3997 (37b); 4105 (1b)
Regnell, A. 448 (56)
Reina, A. 729 (10a)
Reitz, P. 1530 (19); 2008 (19)
Renteria, E. 1773 (37f); 2650 (29d); 2746 (9)
Restrepo, D. 916 (45a)
Revilla, J. 1433 (37d); 2325 (36); 2329 (16a); 3728 (37d); 8376 (2); 8438 (16a); 8600 (60c)
Ribas, O. 2702 (51a)
Ribeiro, B. 1660 (66)
Riedel, L. 732 (19); 734 (51h)
Rimachi, M. 3150 (31); 3935 (36); 10372 (36); 10600 (36); 10965 (37d); 11275 (37d); 11396 (37d); 12210 (36)
Rivera, D. 1024 (45a); 1035 (67a)
Rivera, G. 184 (45b); 720 (14a); 1680 (22a); 1839 (22a); 2337 (57)
Robles, R. 1252 (13a); 1519 (29b); 1903 (35a)
Robleto, W. 628 (49e); 1977 (49e)
Rodrguez, A. 3939 (14g); 4691 (16a)
Rodrguez, E. 261 (60c); 513 (36); 568 (60c); 581 (36); 584 (50); 693 (32a); 981 (60e); 1065 (21); 1136 (8); 1228 (32a)
Rodrguez, J. 139 (43)
Rojas, E. 178 (16a)
Rojas, R. 99 (36); 101 (60e); 256 (60f); 467 (36); 502 (45a); 591 (60c); 592 (60c)
Rojas, W. 443 (67a)
Romero, R. 1273 (51j); 2811 (51j)
Romero-Castaeda, R. 7838 (67a); 7737 (67a); 7795 (45a)
Romoleroux, K. 2080 (60c)
Roncal, J. 3 (36); 5 (6a); 7 (36); 8 (29b); 14 (37c); 19 (16a); 103 (36); 106 (36); 115 (36); 117 (36); 122 (36); 126 (8); 130
(36); 131 (36); 137 (60c); 141 (60i); 142 (60i); 150 (6b); 172 (60d); 177 (60i); 182 (6a); 186 (36); 190 (36); 304
(49e); 308 (14a); 312 (14a); 313 (14a); 317 (20); 320 (14a); 321 (14a); 327 (14a); 329 (14a); 332 (14a); 336 (35b);
340 (14a); 341 (13a); 351 (67d); 353 (14a); 354 (32b); 355 (67d); 356 (14a); 358 (28); 359 (28); 360 (14a); 361
(14a); 364 (32b); 365 (67d); 366 (32b); 367 (14a); 368 (14a); 369 (32b); 371 (14a); 373 (32b); 375 (14a); 382 (16a);
383 (16a); 391 (14c); 394 (12a); 395 (12a); 396 (42); 397 (42); 400 (2); 402 (60g); 495 (5)
Rosa, N. 1600 (37g)
Rossato, M. 9 (56)
267
Rubio, D. 395 (62); 1090 (30); 1146 (7); 1184 (30); 1203 (30); 1258 (10b); 2184 (30)
Rudas, A. 2916 (60c); 3101 (37d); 3531 (8); 5798 (8)
Rueda, R. 2669 (49b); 6823 (49e); 8307 (14g); 9153 (35a); 9521 (14g); 9622 (49e); 10062 (14g); 10373 (49e); 10674
(22c); 14981 (14g)
Ruz, J. 220 (36); 1248 (36); 1300 (60c)
Rusby, H. 411 (37b)
Russell, G. 683 (22a); 893 (22a)
Saboro, J. 105 (14a)
Salaroli, S. 11 (2)
Saldanha, J. 7090 (51h); 8607 (51f)
Saldias, M. 3565 (51j)
Salick, J. 8102 (13a)
Snchez Vega, I. 6281 (45a); 6696 (45a); 8469 (53); 8658 (53); 11235 (21)
Sanders, R. 1781 (14h)
Sandino, J. 219 (22c); 500 (29b); 2372 (29b); 2825 (22c); 4802 (13a)
Sanoja, E. 3391 (16a)
Sant'Ana, S. 311 (51j)
Santos, J. 872 (37d)
Sastre, C. 115 (2); 1567 (60a); 3445 (37i)
Scariot, A. 419 (56); 553 (36); 562 (37c); 563 (41); 572 (16a)
Schatz, G. 1106 (49e)
Schipp, W. 94 (16b); 397 (29b)
Schmalzel, R. 1099 (14c); 1168 (49b); 1595 (32b)
Schultes, R. 3467 (37h); 3884 (37d); 6656 (31); 8290 (31); 8992 (37d); 12733 (16a); 12781 (16a); 13667 (16a); 13968
(33); 14455 (37d); 15838 (36); 16011 (37d); 16258 (16a); 17353 (33); 17854 (16a); 17948 (37g); 18038 (37g);
46315 (31)
Schulz, J. 466 (67a)
Schunke, J. 1406 (37c); 1557 (6a); 5416 (60c); 5505 (37c); 5507 (33); 5603 (60c); 6111 (36); 6613 (29a); 6615 (50);
7194 (50); 7382 (16a); 7612 (50); 7716 (29a); 8080 (6a); 8421 (36); 9274 (36); 9413 (45a); 9416 (48); 9451 (6a);
9912 (6a); 9961 (36); 9962 (50); 10192 (48); 10407 (36); 10804 (50); 10944 (50); 14756 (6a); 15199 (31); 15228
(36); 15262 (60c); 15351 (31); 15360 (6a); 15380 (36); 15598 (6a); 15622 (60c); 16036 (6a); 16061 (60c); 16280
(6a); 16286 (36); 16590 (60c); 514989 (6a)
Schwalke 10970 (51f)
Secco, R. 265 (21); 304 (37a)
Seele, C. 299 (51k)
Seemann, B. s. n. (63)
Seibert, R. 1597 (13a)
Seidel, R. 2930 (16a)
Seymour, F. 5612 (13a)
Shank, P. 4897 (14g)
Shattuck, O. 396 (49b)
Shillingford, C. 428 (49c)
Silva, J. 299 (16a); 1759 (19)
Silveira, M. 1253 (8); 1255 (36)
Silverstone-Sopkin, P. 5182 (29b); 5667 (29b); 5732 (29b); 5919 (29b); 468 (16a); 1381 (67a); 1780 (67a); 4400 (67a);
9424 (14a)
Simonelli, M. 1350 (56)
Skov, F. 60120 (60c); 60123 (60c); 60125 (40); 60126 (40); 64710 (60c); 64716 (36); 64719 (29a); 64722 (67a); 64723
(21); 64724 (21); 64728 (45a); 64729 (67a); 64731 (67a); 64732 (67a); 64740 (14a); 64741 (30); 64743 (14a);
64744 (14a); 64745 (30); 64752 (67a); 64756 (45a); 64760 (45a); 64762 (60c); 64763 (21); 64765 (60c); 64766
(60c); 64767 (60c); 64768 (36); 64775 (67a); 64779 (36); 64781 (67a); 64782 (6a); 64785 (37h); 64788 (29a);
64790 (6a); 64792 (40); 64800 (31); 64804 (36); 64805 (60c); 64806 (60c); 64807 (16a); 64808 (34); 64809 (31);
64810 (60c); 64815 (6a); 64834 (30); 64837 (14d)
Smith, A. 1220 (22a); 2556 (37b); 2770 (2); 10010 (49i)
268
HENDERSON
Smith, D. 240 (41); 1519 (45a); 1935 (60i); 1961 (36); 2019 (60i); 2445 (6b); 2782 (65); 2855 (60i); 2865 (6b); 3676
(6b); 3810 (37c); 3849 (6a); 3964 (60c); 4149 (67a); 4156 (45a); 4195 (67a); 4381 (67a); 4417 (53); 4475 (32a);
4590 (45a); 4842 (45a); 4870 (67a); 4926 (45a); 5259 (6a); 5535 (6a); 5576 (67a); 5791 (45a); 6720 (21); 8201
(67a); 8420 (60i); 8658 (67a); 12898 (21); 12982 (6a); 13044 (16a); 13045 (60c); 14242 (6a)
Smith, H. 2340 (49g); 2341 (45a)
Smith, J. 3 (16b); 530 (5); 1346 (67d); 6845 (22b); 14774 (5)
Smith, L. 5712 (19)
Sneidern, K. 5362 (14a)
Sobel, G. 2000 (67a)
Soejarto, D. 3433 (7); 4448 (29d)
Soerensen, M. 82 (32a)
Solomon, J. 6216 (37e); 6292 (33); 7977 (37e); 8497 (67a); 9260 (45a); 9605 (45a); 9674 (67a); 10875 (31); 14801 (6a);
15640 (67a); 16893 (16a); 17060 (31); 17152 (31)
Souza, D. 89 (56)
Sperling, C. 6210 (37a); 6412 (16a); 6428 (60c); 6611 (36)
Spichiger, R. 1106 (36)
Sprague, T. 349 (32a)
Spruce, R. 29 (37g); 44 (37d); 75 (1a)
Sthl, B. 1899 (60c); 1900 (21); 1920 (34); 1922 (16a); 1925 (36); 1979 (60c); 1980 (64); 2200 (45a); 2359 (45a); 2873
(36); 2874 (67f); 2881 (60c); 2975 (60c); 3259 (60c); 3468 (60c); 3489 (60c); 3490 (60c); 3559 (34); 4375 (60c);
4441 (60c)
Standley, P. 27954 (13a); 37328 (22a); 42046 (45b); 42159 (45b); 42172 (45b); 42692 (45b); 43988 (45b); 44784 (22a);
48954 (13a); 48974 (35a); 48976 (13a); 49004 (13a); 51430 (45b); 72791 (16b); 86906 (49d); 86970 (49d)
Stannard, B. 2414 (51j)
Stauffer, F. 262 (67j); 266 (4); 543 (59)
Stehl, H. 308 (49c); 1528 (49c); 4536 (49c)
Stein, B. 2740 (45a)
Stergios, B. 6567 (32a); 7554 (37d); 9860 (16a); 10221 (16a); 13001 (16a); 13227 (2); 15005 (16a); 15357 (60c); 20104
(45a)
Stern, W. 1841 (29b)
Stevens, W. 4779 (14g); 4979 (14g); 6351 (13a); 6688 (22c); 6796 (13a); 7069 (13a); 8432 (49e); 8434 (16a); 8967
(14a); 11341 (67d); 12092 (49e); 12110 (16a); 13098 (13a); 13104 (29b); 13734 (14a); 13979 (45b); 14770 (49e);
15105 (67d); 16546 (16a); 16634 (29b); 18738 (29b); 19546 (16a); 20455 (45b); 20990 (22c); 21143 (22c); 21217
(22c); 22131 (67d); 22486 (67d); 23438 (35a); 23485 (16a); 23982 (49e); 24281 (13a); 24544 (13a); 24788 (35a);
25578 (22); 25868 (49e)
Stevenson, D. 803 (37d); 817 (43); 867 (37g); 1041 (16a); 1042 (16a)
Stevenson, P. 414 (36)
Steward, W. 277 (37d); 13024 (36)
Steyermark, J. 906 (67b); 9066 (33); 17220 (29b); 33339 (49d); 37091 (49d); 38757 (49b); 41603 (29b); 41873 (22);
44387 (49e); 45442 (49b); 46079 (29b); 47919 (49d); 48742 (67d); 53632 (32a); 55305 (67a); 56391 (45a); 60789
(21); 74634 (16a); 74750 (33); 75212 (67b); 87274 (37b); 88126 (37b); 89119 (37b); 90576 (2); 90627 (52); 91607
(67a); 91859 (58); 94660 (67a); 95204 (58); 95308 (49a); 95360 (59); 97668 (4); 98180 (67b); 98971 (67a); 99046
(67a); 99584 (16a); 99868 (29c); 100313 (4); 101326 (16a); 102463 (37d); 103029 (37g); 103333 (67a); 103879
(67b); 103998 (67b); 106031 (67b); 106230 (4); 106805 (4); 116170 (67b); 116266 (58); 119381 (37f); 120417
(49a); 120582 (16a); 120593 (49a); 120627 (49i); 121520 (49i); 122226 (16a); 122698 (29c); 123796 (4); 124073
(67b); 124821 (67a); 125511 (58); 125565 (49a); 125642 (37g); 125699 (16a); 125997 (67b); 126615 (67a); 127374
(37b); 128223 (67b); 128808 (67b); 129705 (16a); 130435 (67b); 130864 (67b)
Stijfhoorn, E. 798 (49c)
Strudwick, J. 4671 (37a); 5001 (16a); 5007 (2); 5009 (37a); 5010 (16a)
Suarez, E. 3481 (57)
Suclli, E. 2817 (32a)
Sucre, D. 5494 (51e); 8649 (51e); 8819 (19)
Sullivan, G. 365 (28); 366 (32b); 1181 (67a)
Svenning, J.-C. 105 (60c); 140 (60c); 141 (36); 147 (60c); 148 (36); 150 (49a); 151 (40); 168 (60c); 179 (60c)
269
Sylvestre, L. 1490 (51c)

Sytsma, K. 2419 (14f); 2508 (14c); 2771 (12a); 3366 (14a); 3879 (14c); 4082 (14a); 4384 (20); 4943 (32b)
Tamayo, F. 871 (67a); 1130 (67a); 1239 (67a); 1558 (67a); 2529 (32a)
Tannebaum, B. 43 (67a)
Tate, G. 166 (2); 355 (37g); 394 (52)
Taylor, J. 11587 (29b); 11643 (14a); 18020 (14a); 18172 (22a)
Teixeira, L. 1033 (16a)
Tessmann, G. 3317 (6a); 5087 (37d)
Thomas, W. 3023 (16a); 5147 (16a); 5626 (51j); 6088 (19); 9251 (51j); 9438 (51g); 10521 (47); 10688 (47); 10929 (47);
12020 (47); 13151 (47); 13266 (51a); 14115 (51a); 120999 (47)
Tillett, S. 70 (67a); 44921 (37b); 44931 (1b); 45047 (1b); 45458 (37b); 45509 (37b); 45614 (37b); 45662 (21); 45713
(52); 45726 (33); 45855 (33); 673299 (32a); 752303 (67b)
Timan, M. 1588 (36); 1612 (6a)
Tipaz, G. 1322 (30)
Tiwari, S. 905 (37b)
Tomlin, S. 79 (67d)
Tonduz, A.12979 (22a)
Torres, J. 2662 (45a); 2994 (60c); 3020 (60c); 3132 (37i); 3152 (8); 3157 (60c); 3187 (37e); 3245 (37e)
Torres, R. 856 (49e)
Tovares, A. 270 (16a)
Trail, J. 963 (33); 972 (37j); 974 (37j); 976 (37j); 983 (37d); 984 (37d); 985 (37d); 987 (37e); 988 (37e); 989 (37d); 990
(37e); 991 (37d); 993 (37e); 995 (37e); 998 (37d); 999 (37d); 1002 (37d); 1004 (37d); 1005 (37d); 1006 (37d); 1007
(37d); 1008 (37d); 1019 (43)
Triana, J. 723 (45a)
Tsugaru, S. 1833 (16a)
Tunqui, S. 24 (36); 65 (36); 270 (60c); 360 (37e); 484 (37c); 489 (37c); 555 (37c); 578 (34)
Tyson, E. 3235 (12c)
Ule, E. s. n. (51a); 865 (19); 8805 (67b)
Utley, J. 2389 (45b); 3215 (14b); 3993 (49e)
Valenzuela, J. 407 (10b); 635 (30)
Valera, A. 63 (60c)
Valverde, . 736 (13b); 1145 (35a)
Van Andel, T. 245 (60c); 329 (60c)
van der Werf, H. 4684 (6a); 4703 (29c); 5487 (45a); 5647 (49a); 6077 (32a); 13197 (60c)
Vargas, C. 15405 (6a); 15769 (6a)
Vargas, H. 1581 (64); 4001 (67a); 4947 (67a); 6282 (68); 6469 (68)
Vargas, I. 2162 (45a); 2495 (36)
Vargas, L. 863 (60c)
Vargas, O. 143 (22b); 238 (20)
Vargas, W. 1748 (67a)
Vsquez, M. 1557 (49e)
Vsquez, R. 42 (31); 238 (37d); 354 (36); 461 (36); 472 (36); 476 (60c); 477 (37d); 636 (16a); 955 (36); 1052 (37d);
1426 (37d); 1427 (16a); 1818 (36); 2137 (50); 2224 (41); 2228 (60c); 2275 (36); 2448 (37c); 2450 (36); 3153 (36);
3724 (36); 3840 (60c); 4189 (60c); 4555 (8); 4719 (8); 4737 (50); 5061 (16a); 5245 (37e); 5687 (36); 6105 (60c);
6106 (36); 6142 (8); 6271 (36); 6626 (37e); 6850 (36); 6925 (36); 7083 (37e); 7314 (60c); 7413 (60c); 7591 (37e);
7811 (8); 8240 (50); 8531 (60c); 8555 (50); 8795 (36); 8806 (50); 9004 (33); 9005 (60c); 9056 (60c); 9744 (6a);
9745 (60c); 10006 (33); 10103 (36); 10723 (60c); 10724 (50); 10836 (60c); 10885 (60c); 11080 (8); 11143 (8);
11535 (16a); 11773 (60c); 11775 (36); 11779 (36); 12057 (36); 12059 (36); 12164 (36); 12167 (36); 12191 (50);
12209 (50); 12220 (36); 12237 (60c); 12304 (16a); 12362 (36); 13019 (36); 13110 (36); 13112 (60c); 13235 (36);
13261 (16a); 13470 (36); 13851 (60c); 14195 (60c); 14327 (60c); 14413 (50); 14439 (60c); 14444 (16a); 14446
(37d); 14455 (60c); 14456 (50); 16011 (8); 17303 (8); 18420 (50); 18428 (8); 18434 (36); 18508 (60e); 18522 (60e);
18555 (60f); 18741 (60c); 18758 (8); 18850 (60c); 18927 (36); 19403 (36); 19645 (50); 20034 (60e); 20215 (32a);
20217 (45a); 20223 (45a); 20286 (60c); 20435 (45a); 21111 (60e); 21851 (45a); 22263 (36); 22273 (50); 22357
(60f); 22507 (36); 22511 (64); 22526 (60e); 22591 (60e); 23063 (60c); 23068 (8); 23238 (37d); 23925 (50); 24035
270
HENDERSON
(60f); 24166 (60e); 24195 (60c); 24213 (36); 24318 (50); 24320 (60e); 24322 (60c); 24385 (36); 24830 (36); 26268
(45a); 26597 (67a); 26645 (45a); 26686 (32a)
Vaz, A. 532 (51b); 660 (51k)
Velsquez, M. 186 (29d); 7866 (24)
Vlez, N. 1694 (67a)
Vervloet, R. 3306 (56)
Viana, P. s. n. (51j)
Vieira, C. 17 (51b); 576 (16a)
Vieira, M. 816 (21)
Villalobos, R. 158 (13a)
Villarreal, D. 1006 (49e)
Vimercat, J. 290 (51a)
von Hagen, C. 1023 (67d); 1321 (16a); 2016 (32b)
von Tuerckheim, H. 8332 (49e)
Vormisto, J. 40 (50); 44 (60c); 76 (50); 239 (50); 247 (33); 298 (60c); 308 (16d); 363 (6a); 506 (36); 510 (50); 520 (16d);
527 (8); 534 (60c); 547 (60c); 551 (6a); 554 (16d); 557 (8); 558 (50); 600 (6a); 636 (60c); 692 (6a); 693 (36); 698
(36); 720 (8); 760 (6a)
Wachter, T. 135 (60c)
Waechter, J. 2289 (51a)
Wallace, B. 4188 (20)
Wallnfer, B. 13514 (66); 1717688 (67a); 12311287 (60d)
Warner, R. 275 (63)
Warush, A. 97 (34)
Wasshausen, D. 402 (49c); 892 (60e)
Weberbauer, A. 6800 (67a)
Webster, G. 13563 (49c)
Wendt, T. 4748 (29b)
Wessels Boer, J. 176 (2); 181 (16); 194 (37a); 199 (37a); 278 (2); 281 (37a); 320 (16a); 325 (2); 335 (2); 336 (16a); 349
(2); 388 (2); 460 (16); 912 (33); 942 (2); 1064 (60a); 1513 (60a); 1558 (60a); 1598 (60a); 1635 (49i); 1776 (67a);
1869 (29c); 2013 (49a); 2024 (67a); 2026 (67a); 2038 (4); 2048 (45a); 2108 (14a); 2111 (16a); 2255 (37g); 2375
(37d); 2399 (36); 2456 (29c); 2459 (49a); 6508 (60a); 54392 (21); 54437 (21)
White, O. 1094 (41)
Whitefoord, C. 59 (49b); 172 (49a); 178 (49a); 200 (29b); 381 (49b); 417 (7); 3663 (49c)
Williams, L. 737 (36); 740 (36); 939 (6a); 941 (6a); 1089 (41); 1090 (16a); 7836 (37c); 8954 (29b); 13496 (58); 15198
(2); 18756 (22); 23507 (45b); 27791 (45b); 29271 (45b); 29291 (36)
Wilson, P. 405 (29b); 407 (29b)
Wolfe, F. 12335 (45a)
Woodson, R. 1946 (14a)
Woronow, G. 5862 (40); 5885 (21)
Wurdack, J. 43188 (16a)
Young, K. 42 (41); 59 (60c); 69 (16a); 366 (45a); 373 (67a); 985 (36); 1005 (6a)
Young, S. 17 (67a)
Yuncker, T. 5004 (29b); 8473 (49e)
Zak, V. 4638 (60c); 4664 (36)
Zamora, N. 1533 (14a); 1761 (57)
Zappi, D. 9963 (51j)
Zardini, E. 48731 (51j); 48769.1 (51j); 54807 (51j)
Zarucchi, J. 1712 (37e); 1747 (16a); 1748 (60c); 4755 (32a); 5959 (32a); 7010 (45a)
Zent, S. 108516 (16a)
Zona, S. 944 (49c)
Zumbado, M. 59 (67d)
271

Revision Geonoma Phytotaxa 2011

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Phytotaxa 17: 1271

ISSN 1179-3155 (print edition)

A revision of Geonoma (Arecaceae)

Accepted by W. Baker: 2 Nov. 2010; published: 18 Feb. 2011

FIRST PUBLISHED IN 2011 BY

2011 Magnolia Press

Phytotaxa 17 2011 Magnolia Press

REVISION OF GEONOMA (ARECACEAE)

Phytotaxa 17 2011 Magnolia Press

Materials and Methods

Phytotaxa 17 2011 Magnolia Press

REVISION OF GEONOMA (ARECACEAE)

Phytotaxa 17 2011 Magnolia Press

Phytotaxa 17 2011 Magnolia Press

REVISION OF GEONOMA (ARECACEAE)

Phytotaxa 17 2011 Magnolia Press

TABLE 1. Phylogeny Matrix ( = missing data)

Phytotaxa 17 2011 Magnolia Press

Phytotaxa 17 2011 Magnolia Press

. . . . . . Figure 1 continued on the next page

Phytotaxa 17 2011 Magnolia Press

Figure 1 continued from the last page

Phytotaxa 17 2011 Magnolia Press

. . . . . . Figure 2 continued on the next page

Phytotaxa 17 2011 Magnolia Press

Figure 2 continued from the last page

REVISION OF GEONOMA (ARECACEAE)

Phytotaxa 17 2011 Magnolia Press

Phytotaxa 17 2011 Magnolia Press

FIGURE 3. Distribution of all Geonoma specimens examined.

Phytotaxa 17 2011 Magnolia Press

Variability, abundance and range

Phytotaxa 17 2011 Magnolia Press

REVISION OF GEONOMA (ARECACEAE)

Phytotaxa 17 2011 Magnolia Press

Phytotaxa 17 2011 Magnolia Press

Phytotaxa 17 2011 Magnolia Press

Phytotaxa 17 2011 Magnolia Press

terra firme, Loreto

terra firme, Madre de

terra firme, Pasco

REVISION OF GEONOMA (ARECACEAE)

Phytotaxa 17 2011 Magnolia Press

Phytotaxa 17 2011 Magnolia Press

REVISION OF GEONOMA (ARECACEAE)

Phytotaxa 17 2011 Magnolia Press

Phytotaxa 17 2011 Magnolia Press

REVISION OF GEONOMA (ARECACEAE)

Phytotaxa 17 2011 Magnolia Press

Phytotaxa 17 2011 Magnolia Press

REVISION OF GEONOMA (ARECACEAE)

Phytotaxa 17 2011 Magnolia Press

Phytotaxa 17 2011 Magnolia Press

Phytotaxa 17 2011 Magnolia Press

Key to the species of Geonoma

Hispaniola, Lesser Antilles, Trinidad and Tobago ....................................................................................................... 2

Phytotaxa 17 2011 Magnolia Press

Flower pits spirally, alternately, or rarely decussately arranged................................................................................... 7

REVISION OF GEONOMA (ARECACEAE)

Phytotaxa 17 2011 Magnolia Press

Phytotaxa 17 2011 Magnolia Press

REVISION OF GEONOMA (ARECACEAE)

Phytotaxa 17 2011 Magnolia Press