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Phytotaxa 35: 1–88 (2011) 

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PHYTOTAXA 

35 

A revision of Desmoncus (Arecaceae) 

ANDREW HENDERSON 

ISSN 1179-3155 (print edition) 

PHYTOTAXA 

ISSN 1179-3163 (online edition) 

The New York Botanical Garden, Bronx, NY 10458–5126, U.S.A. E-mail: ahenderson@nybg.org 

Magnolia Press 

Auckland, New Zealand 

Accepted by Maarten Christenhusz: 27 Oct. 2010; published: 14 Dec. 2011

ANDREW HENDERSON 

A revision of Desmoncus (Arecaceae) 

(Phytotaxa 35) 

88 pp.; 30 cm. 

14 Dec. 2011 

ISBN 978-1-86977-839-2 (paperback) 

ISBN 978-1-86977-840-8 (Online edition) 

FIRST PUBLISHED IN 2011 BY 

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ISSN 1179-3155 (Print edition) 

ISSN 1179-3163 (Online edition) 

2 Phytotaxa 35 © 2011 Magnolia Press 

HENDERSON

Table of contents 

Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 

Materials and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 

Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 

Taxonomic Treatment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 

Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47 

References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47 

Appendix I. Qualitative Variables–Characters and Traits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50 

Appendix II. Quantitative Variables . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52 

Appendix III. Excluded Names . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53 

Appendix IV. Plates of Type Images. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56 

Appendix V. Numerical List of Taxa and Specimens Examined. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77 

Appendix VI. Index of Names. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86 

Abstract 

A taxonomic revision of the Neotropical palm genus Desmoncus based on morphological data and morphometric 

methods was carried out. Eight hundred and fifty-one herbarium specimens were scored for 16 qualitative variables and 

16 quantitative variables. Qualitative variables were divided into 15 characters and one trait. Using the Phylogenetic 

Species Concept, characters were applied to recognize 24 species. These are widely distributed in Central and South 

America from southern Mexico to Bolivia and Paraguay, and to Trinidad, Tobago, and the Lesser Antilles. Analysis of 

each species for geographic distribution and quantitative variables led to recognition of 9 subspecies in two of the 

species, giving a total of 31 taxa. Seven new species (D. kunarius, D. interjectus, D. loretanus, D. madrensis, D. moorei, 

D. obovoideus, D. osensis) and two new subspecies (D. horridus subsp. occidentalis, D. mitis subsp. ecirratus) are 

described. Five new combinations are made. One of the most variable species is considered to be a species complex and 

is divided into morphotypes: groups of similar specimens without formal taxonomic status. Nomenclature, descriptions, 

and distribution maps are provided for each species and subspecies. Images of type specimens of all new taxa are also 

given. 

Key words: climbing palm, morphometrics, Neotropics, Palmae 

Introduction 

Desmoncus is distinctive among Neotropical palms in its climbing habit. The name is derived from the Greek 

words desmos, meaning a band, and ogkos, meaning a hook, in reference to the climbing hooks, or 

acanthophylls on the leaves. However, not all Desmoncus species are climbers, and not all Neotropical 

climbers are Desmoncus. One other genus, Chamaedorea Willdenow (1806: 638) contains a single species 

with climbing stems, C. elatior Mart. In fact, so similar are the acanthophylls of both genera that specimens of 

C. elatior are commonly misidentified as Desmoncus. The climbing organs of Desmoncus are also remarkably 

similar to those of some Old World rattans, although the two are not closely related. 

Desmoncus was established by Martius (1824). This original brief description was expanded by the same 

author (Martius 1823–1837), who also increased the number of species to seven in the description of his 

Amazon collections. Drude (1881), in the treatment of the palms for Flora Brasiliensis, recognized 17 

species, and Barbosa Rodrigues (1903), also working only on Brazilian palms, recognized 28 species. 

The first revision of the entire genus was that of Burret (1934), a German botanist who worked in the 

Berlin herbarium. In this revision, 41 species were recognized, nine of them new. In a series of papers, Bailey 

(1943, 1947, 1948, 1949) described 14 new species, only two of which are recognized here. Bailey, as well as 

Barbosa Rodrigues and Burret, have been criticized for employing an extremely narrow species concept 

(Wessels Boer 1965, Henderson 1999). Wessels Boer (1965) gave a detailed discussion of variation in 

Desmoncus. In particular he considered that plants of the same species growing in different but adjacent 

A REVISION OF DESMONCUS (ARECACEAE) 

Phytotaxa 35 © 2011 Magnolia Press 3

habitats, for example open areas or forest understory, could exhibit very different leaf morphology, and that 

leaves of young and old plants could also appear quite different. 

Apart from this environmental and ontogenetic variation, there are other sources of taxonomic problems 

in Desmoncus. Because of their spiny nature, plants tend to be shunned by collectors and so there are 

relatively few specimens in herbaria. Apparently Desmoncus plants seldom flower (Henderson 1995) and this 

means that there are few fertile (and many sterile) specimens in herbaria. Fertile specimens that do exist are 

often poorly collected and pressed, making it difficult to measure and score them. There are also many areas 

from which there are few if any specimens, particularly in the Amazon region, making taxonomic decisions 

difficult. Conversely, some small areas are particularly well collected and specimens from these areas have a 

tendency to be over-emphasized. Another problem in Desmoncus is the apparent presence of numerous 

hybrids, discussed below in the Morphology section. 

Because of these issues, Desmoncus has been regarded as a taxonomically difficult genus, and this has led 

to great variation in the number of species recognized over time. Burret (1934) recognized 41 species, 

whereas Henderson et al. (1995), following Wessels Boer (1965, 1988), recognized only seven. Desmoncus 

thus presents a common problem in taxonomy—how to reconcile the splitting of one botanist with the 

lumping of others. 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, 2005b, 2011). 

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). 

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 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 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 geography 

and quantitative variables. 


HENDERSON

Species delimited under the PSC function as hypotheses and these can be tested (Wheeler & Platnick 

2000). This test depends on distinguishing characters from traits, i.e., the test is, that characters are not traits, 

and traits are not characters. In the former 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 

Eight hundred and fifty-one specimens from the following herbaria were examined and scored: A, AAU, BH, 

BM, CEN, COAH, COL, CR, F, GH, HEPH, IBGE, INB, INPA, K, M, MG, MICH, MO, NY, P, PMA, R, S, 

SPF, UB, and US (herbarium abbreviations from Holmgren et al. 1990). Fragmentary type specimens from 

some herbaria (e.g., M, P) were examined but not necessarily scored. Sometimes, more than one duplicate of 

a collection was used in scoring. All sterile specimens, including types, were excluded from this study 

because of the difficulty of identifying them. There are two exceptions. Specimens of Desmoncus cirrhifer 

and D. stans can be unequivocally identified even when sterile, and all specimens of these species were 

included. 

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 a survey of specimens. A dissecting microscope was used to survey 

floral variables. Sixteen qualitative variables were found and scored (Appendix I). 

A search was made for quantitative variables that could be taken from specimen labels (where, in case of 

ranges, median values were used) or measured from specimens. Variables were counted or measured with a 

ruler, digital calipers, or protractor. Sixteen quantitative variables were found and scored (Appendix II). Two 

are from stems, six from leaves, and eight from reproductive structures. Thirteen 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/Desmoncus.xls.zip). Additional columns recorded a specimen identification number, 

collector, collector’s number, herbarium, country, latitude, longitude, and elevation. 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 851 specimens in the matrix, three spatial variables and 32 morphological variables were 

recorded, giving a potential total of 29,785 data points. However, approximately 46% of these potential data is 

missing in the matrix. Specimens are often fragmentary or incomplete, and various organs are often missing 

(especially staminate flowers). Data on plant height and stem 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 

log 10 -transformed before analysis. 

Species delimitation 

All specimens were assigned a preliminary species identification, either based on a previous determination or 



y using keys in local floras (e.g., Henderson 1995, Hammel et al. 2003, Galeano & Bernal 2010). 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. 

Three variables with many missing data (stem branching, rachillae tomentum, stamen number) were 

excluded from this analysis. Subsequently, stem branching was treated as a trait because its states varied 

within some species, and rachillae tomentum and stamen number were treated as characters. 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 geographic distributions and quantitative 

variables. The purpose of these analyses was to look for evidence of presence of discrete subgroups (i.e., 

subspecies). Traits were not used in subspecies delimitation because only one trait, stem branching, was found 

and this had many missing data. 

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. 

Quantitative variation was analyzed. A t-test (two-sample, separate variance test on log 10 -transformed 

variables) or, with more than two variables, a one-way ANOVA (on log 10 -transformed variables) was used to 

test for geographic subgroup differences for each quantitative variable. The Bonferroni pair wise procedure 

was used to see which pairs of means differed significantly (P

Environmental variation 

Linear regression was used to analyze relationships within species and subspecies between log 10 -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

the morphology of several attributes of Desmoncus not used in delimiting species is discussed. Sources of 

morphological variation are also discussed. 

Stems are mostly clustered. Only one species, Desmoncus giganteus has solitary stems, although this is 

recorded from only two specimens. Stems in all but one species, D. stans, are not free-standing, and climb by 

means of acanthophylls. However, one subspecies of D. mitis, subsp. ecirratus is recorded on several 

specimen labels as being non-climbing. Despite the climbing habit of most species, stems of Desmoncus 

seldom reach great heights. The mean plant height of all climbers is only 6.2 m. Isnard et al. (2005) have 

discussed the biomechanics and development of the climbing habit of two species of Desmoncus, D. horridus 

subsp. horridus (as D. orthacanthos) and D. polyacanthos. 

Leaves of Desmoncus are spiny. These spines occur on sheaths, petioles, rachises, cirri, and pinnae and 

provide several useful taxonomic characters. However, there is considerable variation in spininess of leaf 

parts both amongst and within species. Spininess also appears to vary according to age of the stem and habitat 

(Wessels Boer 1965). 

Leaves are arranged all along the stems, and are recorded on labels as being distichously arranged. The 

leaf consists of sheath, petiole, rachis, pinnae, and cirrus. The elongate, basal sheaths tightly enclose the stem 

and persist even after the leaf has died. Sheaths have well-developed ocreas, extending for several centimeters 

above the insertion of the petiole. Petioles are usually rather short, and only in D. cirrhifer (7.0–16.5 cm long), 

D. costaricensis (8–15 cm long), and D. giganteus (15–30 cm long) are they well-developed. Rachises are 

well-developed in all species, and range from 8–190 cm long. Rachis spines are useful taxonomically and 

species can be divided into two groups based on these spines. In one group, rachis spines are straight with 

briefly swollen bases, usually more than 1 cm long, and mostly adaxially or laterally placed on the rachis (Fig. 

1A). In the second group, rachis spines are recurved with markedly swollen bases, usually less than 1 cm long, 

and mostly abaxially placed on the rachis (Fig. 1B). 

The rachis is extended into a cirrus. This climbing organ can be defined as a distal section of the rachis 

which does not bear pinnae but rather acanthophylls. These are pinnae which are modified into reflexed 

hooks. Cirri can take various forms. They may be well-developed with acanthophylls (Fig. 1C); poorlydeveloped 

with the rachis terminating in a short cirrus with or without acanthophylls; virtually absent with the 

rachis terminating beyond the distal most pair of pinnae in a short ‘stub’ (Fig. 1D); or poorly-developed with 

the rachis terminating in a short cirrus with some small, acanthophyll-like pinnae present (Fig. 1E). 

Arrangement of spines on the cirri differs somewhat, and is useful in identification. In one group of species, 

the cirri have few spines abaxially, mostly on proximal part of the cirri only (Fig. 1C); in a second group the 

cirri have spines abaxially throughout (Fig. 1F); in a third group cirri have many, usually paired spines (Fig. 

1E); and in a fourth group the cirri are without spines abaxially (Fig. 1G). 

FIGURE 1. A. Rachis spines usually >1 cm long, mostly adaxial or lateral, straight with briefly swollen bases; pinnae 

with an adaxial beard of spines at the bases (D. chinantlensis; Steyermark 44526). B. Rachis spines



All leaves are pinnate, with pinnae number ranging from 2–28 per side of the rachis. Pinnae are variously 

arranged; in some species they are irregularly arranged and in others they appear almost regularly arranged. In 

D. chinantlensis, and probably other species, younger leaves have strongly clustered pinnae while in larger 

plants they are almost regularly arranged. However, pinnae arrangement is difficult to record from most 

specimens, either because only a short section of the rachis is present, or because folding and pressing of the 

specimen has obscured the arrangement. Pinnae shape ranges from almost linear to broadly ovate, and again 

may depend on age of the leaf. Pinnae of younger plants of some species often appear to more ovate leaves; on 

the other hand, other species have young leaves with narrower pinnae. 

Pinnae apices and bases provide several characters. In two species (D. cirrhifer, D. stans), pinnae have 

long, filiform apices (Fig. 2A). In some species there is an adaxial ‘beard’ of spines at the bases of the pinnae 

(Fig. 1A). In other species there are spinules at the bases of the pinnae (Fig. 1I), and in one species, D. moorei, 

there are both beards and spinules at the pinnae bases. Other species have the bases of the pinnae with smooth 

surfaces adaxially (Fig. 1H). 

Inflorescences of Desmoncus range from densely spiny to almost spineless. They are usually solitary at a 

node, but in one taxon, D. mitis subsp. leptospadix, there are often 2–3 inflorescences per node. Inflorescences 

are always interfoliar and emerge from the apex of the subtending leaf sheath. They have elongate, flattened 

peduncles which are covered by the subtending sheath, and peduncles are seldom present on specimens in 

their entirety. Inflorescences are spicate or usually branched to one order; very rarely a proximal rachilla on an 

inflorescence is bifid. Rachillae number ranges from 1–43. Rachillae are variously arranged on the rachis, and 

the form of the rachises, rachillae, subtending bracteoles, and pulvini are useful taxonomically. 

Inflorescences may have a smooth rachis which is narrower than the few, distantly spaced and alternate 

rachillae, and each rachilla is usually briefly adnate proximally to the rachis with an irregular bracteole 

displaced onto the rachis, and a pulvinus may be absent (Fig. 2B); inflorescences may have an angular, 

slightly twisted rachis which is thicker than the few to numerous, closely spaced and spirally arranged 

rachillae, and each rachilla is not (or very rarely) adnate to the rachis and is subtended by an acute bracteole 

and has a well-developed axillary pulvinus (Fig. 2C); inflorescences may have an angular, slightly twisted 

rachis thicker than the closely spaced and spirally arranged rachillae, and each rachilla is not adnate to the 

rachis but has an irregular bracteole adnate to the rachilla and appearing displaced distally onto the rachilla 

(Fig. 2D), with a poorly-developed axillary pulvinus; inflorescences may have a ridged, not twisted rachis 

which is much thicker than the numerous, closely spaced and spirally or irregularly arranged rachillae, and 

each rachilla is not or only briefly adnate to the rachis and is subtended by an acute bracteole and with a welldeveloped 

axillary pulvinus (Fig. 2E); or the inflorescence may lack a rachis and be spicate (Fig. 2F). 

Rachillae are mostly glabrous but are tomentose in a few species (Fig. 2E). 

FIGURE 2. A. Pinna with long, filiform apex (D. stans; de Nevers 7760). B. Inflorescence with the rachis smooth, not twisted, 

narrower than the few, distantly spaced and alternate rachillae, each rachilla briefly adnate proximally to the rachis and with an 

irregular bracteole displaced onto the rachis, with an axillary pulvinus (D. mitis; Balslev 4775). C. Inflorescence with the rachis 

angular, slightly twisted, thicker than the few to numerous, closely spaced and spirally arranged rachillae, each rachilla not adnate to 

the rachis, subtended by an acute bracteole and with a well-developed axillary pulvinus (D. polyacanthos; Wessels Boer 1647). D. 

Inflorescence with the rachis angular, slightly twisted, thicker than the closely spaced and spirally arranged rachillae, each rachilla not 

adnate to the rachis and with an irregular bracteole adnate to the rachilla and appearing displaced distally onto the rachilla, with a 

poorly-developed axillary pulvinus (D. orthacanthos; Costa 680068). E. Inflorescence with the rachis ridged, not twisted, much 

thicker than the numerous, closely spaced and spirally or irregularly arranged rachillae, each rachilla not adnate to the rachis, 

subtended by an acute bracteole and with a well-developed axillary pulvinus; rachillae brown tomentose initially (D. horridus; 

Davidse 17831). F. Inflorescence rachis absent (inflorescence spicate) (D. stans; de Nevers 7760). G. Peduncular bract broad, the 

surfaces ribbed, brown tomentose, densely covered with long, straight or sinuous, briefly swollen-based, diagonally or vertically 

oriented spines, these flattened or triangular in cross-section, whitish-brown proximally, black or brown distally, with tomentose 

margins (D. chinantlensis; Balick 1715). H. Peduncular bract broad, the surfaces ridged, brown tomentose, densely covered with 

short, recurved, markedly swollen-based, diagonally oriented spines, these triangular in cross-section, whitish-brown proximally, 

brown distally, with tomentose margins (D. polyacanthos; Prance 29626). I. Peduncular bract narrow, elongate, ribbed, scarcely 

brown tomentose, without spines (D. mitis; Nee 34905). Scale bar = 1 cm. 


HENDERSON



Prophylls are elongate and are inserted near the base of the peduncle. Most of the length of the prophyll, 

like the peduncle, is covered by the sheath. There is a single peduncular bract, and this is inserted high up on 

the peduncle usually beyond the apex of the prophyll. The exterior surface of the peduncular bract is useful 

taxonomically and ten different states are recognized here ranging from spiny (Fig. 2G, 2H, 3A, 3C, 3F, 3G) to 

lacking spines (Fig. 2I, 3B, 3D, 3E). 

Flowers are arranged in triads, at least on the proximalmost parts of the rachillae. Commonly distal most 

parts of the rachillae bear staminate flowers only, and this part of the rachilla is often deciduous. Triads are 

surrounded by variously shaped bracteoles. Staminate flowers have a short, three-lobed calyx and much 

longer, free, lanceolate petals. The apices of the petals are caudate. Stamen number ranges from 5–12, 

although most species have six stamens. However, this is seldom recorded because staminate flowers are early 

deciduous and are seldom present on specimens. Filaments are partly and irregularly adnate to the petals. 

Pistillate flowers have a very short, lobed, cupular calyx and much longer, tubular, lobed corolla. There are 

approximately six, very small staminodes. The gynoecium is trilocular and triovulate. Where known, 

inflorescences are protogynous (see Listabarth 1994 for pollination in Desmoncus). 

Fruits are variously colored and shaped. Most are described on specimen labels as being red or bright red, 

but yellow is also recorded. Mesocarp fibers are an useful identification character in Desmoncus. These fibers, 

or their lack, give fruit surfaces a characteristic appearance ranging from uneven with numerous, 

subepidermal, short, often branching (Y-shaped) fibers (Fig. 3H); smooth without any apparent subepidermal 

fibers (Fig. 3I); uneven with numerous, subepidermal, long, branching fibers (Fig. 4A); to bumpy from the 

numerous, subepidermal, short, oblique fibers (Fig. 4B). Fruiting corollas in one species, D. giganteus, are 

almost half as long as fruits (Fig. 4C). In most species, fruiting corollas split irregularly into 3 lobes and the 

lobes often split again (Fig. 4E) but in a few species fruiting corollas do not or scarcely split and tend to 

remain cupular (Fig. 4D). Endocarp shape varies from globose to obovoid with rounded or slightly peaked 

apices; narrowly ellipsoid with rounded apices; broadly obovoid with flattened apices; to ovoid to obovoid 

with prominent, peaked apices (Fig. 4F). 

Eophylls have one pair of pinnae. However, only four specimens have been seen with eophylls 

(representing D. chinantlensis, D. leptoclonos, D. myriacanthos, and D. polyacanthos). Galeano & Bernal 

(2010) illustrated the eophylls of D. cirrhifer and D. myriacanthos (as D. orthacanthos) as bifid. According to 

Dransfield et al. (2008), the eophyll of D. costaricensis has two pairs of pinnae. 

FIGURE 3. A. Peduncular bract broad, ridged, densely covered with short, straight, swollen-based, vertically oriented 

spines, these terete, whitish-brown proximally, brown distally, without tomentum (D. parvulus; Mori 8063). B. 

Peduncular bract broad, ribbed, densely brown tomentose, without spines (D. vacivus; Vásquez 2646). C. Peduncular 

bract broad, ribbed or ridged, densely covered with felty, reddish-brown tomentum, sparsely covered with short, scarcely 

swollen-based, diagonally oriented, flattened spines, whitish-brown proximally, brown distally, with tomentose margins 

(D. cirrhifer; Moore 9471) D. Peduncular bract broad, ribbed with several more prominent, lighter colored ribs, brown, 

scarcely or not tomentose, without spines (D. loretanus; Gentry 25785). E. Peduncular bract narrow, ribbed, densely 

whitish-brown tomentose, not spiny (D. stans; Grayum 8115). F. Peduncular bract broad, the surface ribbed or ridged, 

brown tomentose or glabrous, sparsely to moderately covered with short, straight or sinuous, briefly swollen-based, 

diagonally or vertically oriented spines, these flattened or triangular in cross-section, whitish-brown proximally, black or 

brown distally, with tomentose margins (D. leptoclonos; Irwin 7194). G. Peduncular bract broad, the surface deeply 

ridged, dark brown tomentose, sparsely covered with long, straight or sinuous spines, the bases scarcely swollen but 

running directly into the ridges of the bract and lying flat against the bract surface, flattened in cross-section, brown 

proximally and distally, with tomentose margins (D. interjectus; Schultes 16233). H. Fruit surface uneven with 

numerous, subepidermal, short, often branching (Y-shaped) fibers (D. orthacanthos; Carvalho 627). I. Fruit surfaces 

smooth, without any apparent subepidermal fibers; fruiting corolla splitting irregularly into 3 lobes, the lobes splitting 

again (D. polyacanthos; Nascimento 723). Scale bar = 1 cm. 


HENDERSON



FIGURE 4. A. Fruit surface uneven with numerous, subepidermal, long, branching fibers (D. obovoideus; Morales 

6333). B. Fruit surface bumpy from numerous, subepidermal, short, oblique fibers (D. kunarius; de Nevers 4415). C. 

Fruiting corollas to half as long as fruits (D. giganteus; Henderson 1688). D. Fruiting corollas not or scarcely splitting, 

tending to remain cupular; fruiting corollas less than one quarter as long as fruits (D. myriacanthos; Cuadros 1855). E. 

Fruiting corollas splitting irregularly into 3 lobes, the lobes often splitting again (D. polyacanthos; Smith 3791). F. 

Endocarps. Top right—endocarp broadly obovoid with flattened apices, the pores lateral on or near flattened apices; (D. 

obovoideus; Aguilar 11417). Middle left—endocarp globose to obovoid with rounded or slightly peaked apices, the 

pores lateral (D. chinantlensis; Bangham 368). Middle right—endocarp narrowly ellipsoid with rounded apices, the 

pores lateral (D. myriacanthos; Croat 9314). Bottom left—endocarp ovoid to obovoid with prominent, peaked apices, 

the pores lateral (D. giganteus; Huashikat 1099). Scale bar = 1 cm. 

Morphological variation 

There are several sources of infraspecific variation in morphology and these make the taxonomy of 

Desmoncus problematic. There appears to be no evidence that this variation is due to any cytological or 

breeding system abnormality (e.g., polyploidy, apomixis). The genus has a chromosome number of 2n = 30 

(although only 3 species have been counted) and there is no evidence of polyploidy (Dransfield et al. 2008). 


HENDERSON

Plants of Desmoncus are monoecious and, where known, they are outcrossing and pollinated by beetles 

(Listabarth 1994). Observed variation may be due to only a few factors—environment, ontogeny, clines, and 

hybridization. 

Environmental variation has been mentioned in the introduction. Wessels Boer (1965) gave a detailed 

discussion and illustration of this. He showed that plants of the same species growing in different but adjacent 

habitats could exhibit very different leaf morphology and spininess. Plants of D. polyacanthos exposed to full 

sunlight had sheaths with numerous, rather straight, swollen-based spines; plants growing in dense shade had 

almost spineless sheaths; and plants growing in a riparian habitat in full sun were larger than normal and were 

densely spiny. Exactly this type of variation is observed amongst the specimens of D. polyacanthos, and it has 

misled earlier botanists (e.g., Bailey 1943, 1947, 1948, 1949). 

Ontogenetic variation in Desmoncus is also apparent. For example, in D. chinantlensis, leaves of young 

plants have rachises which often lack spines and pinnae which are shorter, ovate, strongly clustered, and lack 

spines at the bases. Adult plants have spiny rachises and longer, more linear, irregularly arranged pinnae with 

beards of spines at the bases. It was this kind of variation that led to confusion over the types of D. 

chinantlensis (Bailey 1933). On the other hand, juvenile plants of one outlying population of D. moorei have 

numerous, linear, regularly arranged pinnae and appear quite different from adult plants with fewer, ovate, 

irregularly arranged pinnae. 

There is only one known case of clinal variation in Desmoncus. In D. orthacanthos, which has a linear 

distribution along the Atlantic coast of Brazil, regression shows petiole length, rachis length, basal pinna 

length, basal pinna width, peduncular bract length, and rachilla length decrease from north to south whereas 

fruit length increases from north to south. Clinal variation seems a minor source of variation within 

Desmoncus, particularly compared with Geonoma (Henderson 2011). However, Geonoma species usually 

have much wider elevation ranges than Desmoncus and many Geonoma species exhibit variation with change 

in elevation. 

A fourth source of variation is hybridization. Interspecific hybrids in palms have been reported in only a 

few genera, and most of these are cocosoid palms (especially Attalea, Bactris, Butia, Allagoptera, Syagrus). 

Desmoncus is also a cocosoid palm, so perhaps it is not surprising to find frequent hybrids. Although it is 

difficult to recognize hybrids from herbarium specimens, they are identified here based on two criteria. First, 

that specimens exhibit intermediate morphology, and second that they occur sympatrically with both putative 

parent species. Eighteen specimens (2% of all specimens) are here postulated to be of hybrid origin. All these 

involve D. polyacanthos, the most widespread and variable species, and either D. pumilus, D. mitis or D. 

horridus. These last two, D. mitis and D. horridus, are also widespread and variable. All postulated hybrids 

come from the central and western Amazon region and none occur in other regions. 

A fifth problem causing taxonomic problems has to do with species complexes. One species, Desmoncus 

polyacanthos, the most variable in the genus, is here regarded as a species complex. Such complexes were 

defined by Henderson (2011) as widespread, variable species in which numerous local variants occur. 

Desmoncus polyacanthos is extremely variable and defies subspecific division. It is of interest to note that the 

greatest variation in this species occurs in the same area as that of several species complexes in Geonoma— 

the western Amazon basin and sub-andean foothills. Here, D. polyacanthos exhibits a myriad of local forms, 

often with more than one form occurring sympatrically. Henderson (2011) considered that the high levels of 

variability in species complexes may be based on resource (habitat) polymorphisms, and these may represent 

an intermediate step in sympatric speciation. 

These factors—environment, ontogeny, clines, hybridization, and species complexes—make a revision of 

Desmoncus difficult. The characters used here in the delimitation of species (Appendix I) are sometimes 

difficult to define and score, particularly those to do with spininess in both leaves and peduncular bracts. The 

difficulties of definition and scoring of characters also lead to problems in using these characters in 

phylogenetic reconstruction. There are also other problems with phylogeny in Desmoncus. One character in 

particular illustrates this very well. The genus has traditionally been split into two sections depending on 

whether the leaf rachis spines are straight or recurved (e.g. Burret 1934), and this character is also used here. 


Phytotaxa 35 © 2011 Magnolia Press • 15

However, there is evidence that in at least one species, D. interjectus, recurved spines appear to have been 

secondarily derived from straight spines. Such reversals complicate phylogenetic reconstruction. 

A second problem in phylogenetic reconstruction is found in choice of outgroups. If more evidence of the 

potential of morphology to mislead is needed, it is found in generic relationships. Desmoncus is placed in the 

subfamily Arecoideae, tribe Cocoseae, subtribe Bactridinae (Dransfield et al. 2008). The Bactridinae contains 

five genera (Acrocomia, Aiphanes, Astrocaryum, Bactris, and Desmoncus). Recently Eiserhardt et al. (2011) 

have proposed that Desmoncus and Acrocomia are sister genera. Such a relationship would never be 

postulated based on morphology; in fact Desmoncus appears most dissimilar to Acrocomia and most similar to 

Bactris. The final point in considering phylogeny based on morphology is that the leaf structure of Desmoncus 

is so different from potential outgroups (e.g., Acrocomia) that it is not possible to polarize character states. For 

these reasons, phylogenetic reconstruction based on morphological data is not attempted here. 

Taxonomic Treatment 

Desmoncus Martius (1824: 20) 

Type:—Desmoncus polyacanthos Martius. 

Atitara Barrère ex Kuntze (1891: 726). Type:—Atitara polyacantha (Martius) Kuntze. 

Stems solitary, or clustered. Leaf rachis spines usually >1 cm long, mostly adaxial or lateral, straight with 

briefly swollen bases, or rachis spines usually

own tomentose, sparsely to densely covered with long, straight or sinuous, briefly swollen-based, 

diagonally or vertically oriented spines, these flattened or triangular in cross-section, whitish-brown 

proximally, black or brown distally, with tomentose margins (rarely without spines), or peduncular bracts 

broad, the surfaces ridged, brown tomentose, sparsely to densely covered with short, recurved, markedly 

swollen-based, diagonally oriented spines, these triangular in cross-section, whitish-brown proximally, brown 

distally, with tomentose margins (rarely without spines), or peduncular bracts narrow, elongate, ribbed, 

scarcely brown tomentose, without spines (rarely with few spines), or peduncular bracts broad, ridged, 

densely covered with short, straight, swollen-based, vertically oriented spines, these terete, whitish-brown 

proximally, brown distally, without tomentum, or peduncular bracts broad, ribbed, densely brown tomentose, 

without spines (very rarely with a few spines), or; peduncular bracts broad, ribbed or ridged, densely covered 

with felty, reddish-brown tomentum, sparsely covered with short, scarcely swollen-based, diagonally 

oriented, flattened spines, whitish-brown proximally, brown distally, with tomentose margins, or peduncular 

bracts broad, ribbed with several more prominent, lighter colored ribs, brown, scarcely or not tomentose, 

without spines (rarely with a few spines), or peduncular bracts narrow, ribbed, densely whitish-brown 

tomentose, not spiny, or peduncular bracts broad, the surfaces ribbed or ridged, brown tomentose or glabrous, 

sparsely to moderately covered with short, straight or sinuous, briefly swollen-based, diagonally or vertically 

oriented spines, these flattened or triangular in cross-section, whitish-brown proximally, black or brown 

distally, with tomentose margins, or peduncular bracts broad, the surfaces deeply ridged, dark brown 

tomentose, sparsely covered with long, straight or sinuous spines, the bases scarcely swollen but running 

directly into the ridges of the bract and lying flat against the bract surface, flattened in cross-section, brown 

proximally and distally, with tomentose margins; rachillae brown tomentose initially, or rachillae glabrous or 

scarcely tomentose initially; stamens five to seven, or stamens eight to 12; fruit surfaces uneven with 

numerous, subepidermal, short, often branching (Y-shaped) fibers, or fruit surfaces smooth, without any 

apparent subepidermal fibers, or fruit surfaces uneven with numerous, subepidermal, long, branching fibers, 

or fruit surfaces bumpy from numerous, subepidermal, short, oblique fibers; fruiting corollas less than one 

quarter as long as fruits, or fruiting corollas to half as long as fruits; fruiting corollas splitting irregularly into 

3 lobes, the lobes often splitting again, or fruiting corollas not or scarcely splitting, tending to remain cupular; 

endocarps globose to obovoid with rounded or slightly peaked apices, the pores lateral, or endocarps narrowly 

ellipsoid with rounded apices, the pores lateral, or endocarps broadly obovoid with flattened apices, the pores 

lateral on or near flattened apices, or endocarps ovoid to obovoid with prominent, peaked apices, the pores 

lateral. 

Key to the species of Desmoncus 

1 Mexico and Central America ...................................................................................................................................... 2 

- South America, Trinidad and Tobago, and the Lesser Antilles .................................................................................. 10 

2 Pinnae with long, filiform apices; cirri absent, the rachis terminating beyond the distalmost, opposite pair of pinnae 

in a

often branching (Y-shaped) fibers, or fruit surfaces with numerous, subepidermal, long, branching fibers; proximal 

rachillae 12.1(5.5–19.0) cm long, 1.2(0.6–2.2) mm wide; widespread ........................................................................ 5 

5 Rachillae brown tomentose initially; pinnae with spinules at the bases; Caribbean coast of Nicaragua and Costa Rica 

and Nicoya Peninsula, Costa Rica .................................................................................................................D. moorei 

- Rachillae glabrous or scarcely tomentose initially; pinnae with smooth surfaces at the bases adaxially, without 

spinules or dense tomentum; widespread .................................................................................................................... 6 

6 Fruiting corollas cupular, not or scarcely splitting; endocarps narrowly ellipsoid with rounded apices; central and 

eastern Panama ................................................................................................................................... D. myriacanthos 

- Fruiting corollas splitting irregularly into 3 lobes, the lobes often splitting again; endocarps globose to obovoid with 

rounded apices and lateral pores, or endocarps broadly obovoid with flattened apices, the pores lateral on or near 

flattened apices; widespread ........................................................................................................................................ 7 

7 Fruit surfaces with numerous, subepidermal, long, branching fibers.......................................................................... 8 

- Fruit surfaces with numerous, subepidermal, short, often branching (Y-shaped) fibers ............................................ 9 

8 Basal pinna 14.0 cm long, 4.9(4.2–5.5) cm wide; central Panama with outliers in western and eastern Panama ........ 

............................................................................................................................................................... D. obovoideus 

- Basal pinna 23.2(18.0–26.0) cm long, 3.7(2.4–5.0) cm wide); Osa Peninsula and adjacent areas in Costa Rica ......... 

........................................................................................................................................................................D. osensis 

9 Petioles 10.5(8.0–15.5) cm long; pinnae without a beard of spines at the bases adaxially Caribbean coast of Costa 

Rica ...................................................................................................................................................... D. costaricensis 

- Petioles 2.2(0.5–4.7) cm long; pinnae with an adaxial beard of spines at the bases; southern Mexico, Belize, Guatemala, 

Honduras, and Nicaragua ............................................................................................................ D. chinantlensis 

10 Pinnae with long, filiform apices; cirri poorly-developed, the rachis terminating in a short cirrus with spines abaxially, 

acanthophylls absent but some small, acanthophyll-like pinnae present; western Colombia and western Ecuador 

..................................................................................................................................................................... D. cirrhifer 

- Pinnae without long, filiform apices; cirri well-developed, with acanthophylls, or poorly-developed, the rachis terminating 

in a short cirrus, acanthophylls present or absent, or absent, the rachis terminating beyond the distalmost 

pair of pinnae in a short ‘stub’; widespread ............................................................................................................... 11 

11 Leaf spines, especially on the rachis (where usually >1 cm long and mostly adaxial or lateral), straight with briefly 

swollen bases; cirri without spines abaxially ............................................................................................................. 12 

- Leaf spines, especially on the rachis (where usually

any apparent subepidermal fibers ............................................................................................................................. 20 

19 Peduncular bract surfaces ridged, brown tomentose, densely covered with short, recurved, markedly swollen-based, 

diagonally oriented spines; fruits 11.9(11.2–12.8) mm long; central Amazon region of Brazil and Colombia ............ 

....................................................................................................................................................................... D. setosus 

- Peduncular bract surfaces ribbed, densely brown tomentose, without spines (very rarely with a few spines); fruits 

20.9(18.4–23.9) mm long; western Amazon region in Colombia, Peru, and Brazil ..................................... D. vacivus 

20 Fruit surfaces smooth, without any apparent subepidermal fibers ..................................................... D. polyacanthos 

- Fruit surfaces uneven with numerous, subepidermal, short, often branching (Y-shaped) fibers .............................. 21 

21 Peduncular bract surfaces deeply ridged, dark brown tomentose, sparsely covered with long, straight or sinuous 

spines, the bases scarcely swollen but running directly into the ridges of the bract and lying flat against the bract surface; 

Colombia (Amazonas, Vaupés) ....................................................................................................... D. interjectus 

- Peduncular bracts ribbed with several more prominent, lighter colored ribs, brown, scarcely or not tomentose, without 

spines (rarely with a few spines), or peduncular bract surfaces ridged, brown tomentose, sparsely to densely covered 

with spines; widespread, excluding Colombia .................................................................................................. 22 

22 Peduncular bracts ribbed with several more prominent, lighter colored ribs, brown, scarcely or not tomentose, without 

spines (rarely with a few spines); Peru (Loreto) ................................................................................. D. loretanus 

- Peduncular bract surfaces ridged, brown tomentose, sparsely to densely covered with spines; Peru (Madre de Dios, 

Ucayali), eastern Bolivia, Brazil (Goiás, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Rondônia), and Paraguay 

(San Pedro) ................................................................................................................................................................. 23 

23 Pinnae bases with uneven surfaces at the bases adaxially, usually covered with spinules and/or dense tomentum; 

Peru (Madre de Dios, Ucayali) ................................................................................................................ D. madrensis 

- Pinnae bases with smooth surfaces adaxially, without spinules or dense tomentum ................................................ 24 

24 Endocarps globose to obovoid with rounded or slightly peaked apices; eastern Bolivia ......................... D. latisectus 

- Endocarps narrowly ellipsoid with rounded apices; Brazil (Goiás, Mato Grosso, Mato Grosso do Sul, Minas Gerais, 

Rondônia) and Paraguay (San Pedro) .................................................................................................... D. leptoclonos 

1. Desmoncus chinantlensis Liebm. ex Martius (1837–1837: 321). 

Atitara chinantlensis (Liebm. ex Martius) Kuntze (1891: 727). 

Lectotype (designated by Bailey 1933):—MEXICO. Oaxaca: Lacoba, Chinantla, no date, F. Liebmann 6595 (C, n.v., 

lectotype image!). 

Desmoncus anomalus Bartlett (1935: 84). Type:—GUATEMALA. Alta Vera Paz: Secanquím, 29 April 1904, O. Cook & 

C. Doyle 97 (holotype US!). Synon. nov. 

Desmoncus lundellii Bartlett (1935: 84). Type:—GUATEMALA. Petén: near El Paso de Petén, 26 April 1932, C. 

Lundell 1555 (holotype MICH!). 

Desmoncus quasillarius Bartlett (1935: 85). Type:—BELIZE. Corozal: San Andres, July 1933, P. Gentle 348 (holotype 

MICH!). 

Desmoncus uaxactunensis Bartlett (1935: 86). Type:—GUATEMALA. Petén: Uaxactún, 18 April 1931, H. Bartlett 

12576 (holotype MICH!). 

Desmoncus ferox Bartlett (1935: 87). Type:—GUATEMALA. Petén: Tikal, 12–15 April 1931, H. Bartlett 12584 

(holotype MICH!). 

Plants 8.1(2.5–20.0) m tall; stems 2.5(1.4–3.8) cm diameter, solitary or clustered. Leaf petioles 2.2(0.5–4.7) 

cm long; rachises 108.0(96.0–128.0) cm long, 10.5(5.7–15.0) mm wide, the spines usually >1 cm long, mostly 

adaxial or lateral, straight with briefly swollen bases; pinnae 23(18–35) per side of rachis, without long, 

filiform apices, with an adaxial beard of spines at the bases, without spinules or dense tomentum at the bases 

adaxially; basal pinna 21.6(7.5–31.0) cm long, 1.9(1.0–2.5) cm wide; cirri well-developed, with 

acanthophylls, without spines abaxially, with intermediate acanthophylls present, without a wide gap between 

pinnae and acanthophylls. Inflorescences with the rachis ridged, not twisted, much thicker than the numerous, 

closely spaced and spirally or irregularly arranged rachillae, each rachilla not or only briefly adnate to the 

rachis, subtended by an acute bracteole and with a well-developed axillary pulvinus; peduncles 7.2(5.0–13.1) 

mm wide; peduncular bracts 37.3(26.5–69.0) cm long, broad, the surfaces ribbed, brown tomentose, sparsely 

to densely covered with long, straight or sinuous, briefly swollen-based, diagonally or vertically oriented 

spines, these flattened or triangular in cross-section, whitish-brown proximally, black or brown distally, with 



tomentose margins; rachillae 38(28–43), glabrous or scarcely tomentose initially; proximal rachillae 

13.0(9.0–19.0) cm long, 1.2(0.9–1.7) mm wide; stamens 9(8–11); fruits 15.2(12.1–18.6) mm long, 12.7(10.3– 

16.2) mm wide, the surfaces uneven with numerous, subepidermal, short, often branching (Y-shaped) fibers; 

fruiting corollas less than one quarter as long as fruits, splitting irregularly into 3 lobes, the lobes often 

splitting again; endocarps globose to obovoid with rounded or slightly peaked apices, the pores lateral. 

Distribution and habitat:—From 13°03’–18°35’N and 84°58’–96°24’W in southern Mexico 

(Campeche, Chiapas, Oaxaca, Quintana Roo, Tabasco, and Veracruz), Belize, Guatemala, Honduras, and 

Nicaragua at 187(10–550) m elevation in lowland rainforest, often in limestone areas (Fig. 5). 

FIGURE 5. Distribution maps of Desmoncus chinantlensis, D. cirrhifer, D. costaricensis, and D. giganteus. 

Taxonomic notes:—There are three original collections from Oaxaca of Desmoncus chinantlensis— 

Liebmann 6594, 6595, and 6596, none of which was cited by Martius. Bailey (1933; see also Schultes 1940) 

designated Liebmann 6595 from C as lectotype, and this specimen is illustrated by Dahlgren (1959, plate 

180). Bailey noted that this specimen was somewhat different from the other two specimens, both at US. This 

is probably because the leaf represented by the lectotype is from a young plant. Leaves of young plants, which 

are frequently collected, have rachises which often lack spines and pinnae which are shorter, ovate, strongly 

clustered, and lack spines at the bases. 

Bartlett (1935) described five species from Belize and Guatemala, none of which are recognized here. 

One of these, D. anomalus has narrow pinnae and pistillate flowers with unusual staminodes. Another 


HENDERSON

specimen (Cook 53) from the same locality has normal pinnae and staminodes. Both Desmoncus quasillarius 

and D. uaxactunensis were said to have peduncular bracts ‘entirely or nearly unarmed’ or ‘almost unarmed’, 

but both are spiny as in other specimens. 

Subspecific variation:—There is no geographic disjunction except for an outlying specimen from 

Nicaragua, but this is likely to be an artifact of insufficient collecting. Two specimens from Honduras (Evans 

1702, Saunders 553) have somewhat tomentose rachillae, more like those of Desmoncus moorei. Other 

specimens from Honduras have less pronounced beards of spines at the pinnae bases, although this may be 

because they represent young leaves. 

2. Desmoncus cirrhifer Gentry & Zardini in Gentry (1988: 1436), as “cirrhifera”. 

Type:—COLOMBIA. Valle: Bahía Malaga, 0 m, 4°02'N, 76°15'W, 16 December 1985, A. Gentry, M. Monsalve, C. 

Restrepo & J. Gamboa 53392 (holotype CUVC n.v., isotypes MO!, COL n.v., K!). 

Plants 11.0(3.0–20.0) m tall; stems 2.0(1.2–3.0) cm diameter, clustered. Leaf petioles 10.8(7.0–16.5) cm 

long; rachises 105.3(91.0–117.0) cm long, 6.0(4.0–8.1) mm wide, the spines usually 1 cm long, mostly adaxial or 

lateral, straight with briefly swollen bases; pinnae 9(8–10) per side of rachis, without long, filiform apices, 



without a beard of spines at the bases adaxially, without spinules or dense tomentum at the bases adaxially; 

basal pinna 24.5(14.5–32.0) cm long, 5.7(3.0–9.0) cm wide; cirri well-developed, with acanthophylls, without 

spines abaxially, with intermediate acanthophylls present, without a wide gap between pinnae and 

acanthophylls. Inflorescences with the rachis ridged, not twisted, much thicker than the numerous, closely 

spaced and spirally or irregularly arranged rachillae, each rachilla not or only briefly adnate to the rachis, 

subtended by an acute bracteole and with a well-developed axillary pulvinus; peduncles 6.4(4.6–8.2) mm 

wide; peduncular bracts 20.5(15.0–28.5) cm long, broad, the surfaces ribbed, brown tomentose, sparsely to 

densely covered with long, straight or sinuous, briefly swollen-based, diagonally or vertically oriented spines, 

these flattened or triangular in cross-section, whitish-brown proximally, black or brown distally, with 

tomentose margins; rachillae 33, glabrous or scarcely tomentose initially; proximal rachillae 8.5 cm long, 

1.0(0.7–1.2) mm wide; stamens 6; fruits 13.8(12.7–14.5) mm long, 11.3(9.1–13.8) mm wide, the surfaces 

uneven with numerous, subepidermal, short, often branching (Y-shaped) fibers; fruiting corollas less than one 

quarter as long as fruits, splitting irregularly into 3 lobes, the lobes often splitting again; endocarps globose to 

obovoid with rounded or slightly peaked apices, the pores lateral. 

Distribution and habitat:—From 9°34’–10°02’N and 82°36’–83°26’W on the Caribbean coast of Costa 

Rica at 128(1–400) m elevation in lowland rainforest (Fig. 5). 

Taxonomic notes:—Desmoncus costaricensis is characterized by its leaves with few spines and large 

pinnae. See Grayum (1998) for a discussion of the nomenclature of this species. 

Subspecific variation:—One specimen (Mora 580) is tentatively included here. It is from the same 

locality as other specimens of Desmoncus costaricensis but the leaf, possibly from a juvenile plant, is 

considerably smaller. 

4. Desmoncus giganteus Henderson (1995: 225). 

Type:—BRAZIL. Acre: Rio Moa near mouth of Rio Azul, 7°25'S, 73°15'W, 14 February 1992, A. Henderson, F. Chávez 

& J. Guedes 1688 (holotype INPA!, isotypes K!, NY!). 

Plants 22.3(4.5–50.0) m tall; stems 4.4(4.0–4.8) cm diameter, solitary. Leaf petioles 20.5(15.0–30.0) cm 

long; rachises 157.5(125.0–190.0) cm long, 12.2(10.1–14.2) mm wide, the spines usually >1 cm long, mostly 

adaxial or lateral, straight with briefly swollen bases; pinnae 10 per side of rachis, without long, filiform 

apices, with an adaxial beard of spines at the bases, without spinules or dense tomentum at the bases 

adaxially; basal pinna 39.2(32.0–46.5) cm long, 6.5(5.7–7.3) cm wide; cirri well-developed, with 

acanthophylls, without spines abaxially, with intermediate acanthophylls present, without a wide gap between 

pinnae and acanthophylls. Inflorescences with the rachis ridged, not twisted, much thicker than the numerous, 

closely spaced and spirally or irregularly arranged rachillae, each rachilla not or only briefly adnate to the 

rachis, subtended by an acute bracteole and with a well-developed axillary pulvinus; peduncles 8.5(6.5–11.7) 

mm wide; peduncular bracts length no data, broad, the surfaces ribbed, brown tomentose, densely covered 

with long, straight or sinuous, briefly swollen-based, diagonally or vertically oriented spines, these flattened 

or triangular in cross-section, whitish-brown proximally, black or brown distally, with tomentose margins; 

rachillae number no data, glabrous or scarcely tomentose initially; proximal rachillae 16.0 cm long, 1.7 mm 

wide; stamens number no data; fruits 36.2(21.5–42.6) mm long, 16.6(12.7–18.3) mm wide, the surfaces 

smooth, without any apparent subepidermal fibers; fruiting corollas to half as long as fruits, splitting 

irregularly into 3 lobes, the lobes often splitting again; endocarps ovoid to obovoid with prominent, peaked 

apices, the pores lateral. 

Distribution and habitat:—From 0°32’–7°25’S and 70°10’–78°15’W in the western Amazon region in 

Colombia, Ecuador, Peru, and Brazil at 266(150–440) m elevation in lowland rainforest or secondary forest 

(Fig. 5). 


HENDERSON

Taxonomic notes:—Desmoncus giganteus is characterized by its large size and unusually large fruits 

with long fruiting corollas. It is the only species of Desmoncus always reported to have solitary stems; all 

other species have clustered or rarely solitary stems. It is probably more common than the few specimens 

suggest. 

Moreno Suárez and Moreno Suárez (2006) consider that this species occurs in Bolivia. However, the 

description and illustrations they give of the fruits do not match those of D. giganteus, and appear more like 

those of the large morphotype of D. polyacanthos. 

5. Desmoncus horridus Splitg. ex Martius (1844: 51). 

Atitara horrida (Splitg. ex Martius) Kuntze (1891: 727). Type:—SURINAME. Paramaribo, no date, F. Splitgerber 61 

(holotype BR, n.v., holotype image!). 

Plants 6.7(2.0–15.0) m tall; stems 2.4(1.2–4.8) cm diameter, clustered. Leaf petioles 4.6(2.0–11.5) cm long; 

rachises 123.8(32.0–220.0) cm long, 9.5(3.4–20.5) mm wide, the spines usually >1 cm long, mostly adaxial or 

lateral, straight with briefly swollen bases; pinnae 19(7–28) per side of rachis, without a beard of spines at the 

bases, without spinules or dense tomentum at the bases adaxially; basal pinna 21.6(12.0–34.0) cm long, 

1.9(1.3–3.7) cm wide; cirri well-developed, with acanthophylls, without spines abaxially, with intermediate 

acanthophylls present, without a wide gap between pinnae and acanthophylls. Inflorescences with the rachis 

ridged, not twisted, much thicker than the numerous, closely spaced and spirally or irregularly arranged 

rachillae, each rachilla not or only briefly adnate to the rachis, subtended by an acute bracteole and with a 

well-developed axillary pulvinus; peduncles 6.4(2.3–13.1) mm wide; peduncular bracts 28.3(10.5–47.0) cm 

long, broad, the surfaces ribbed, brown tomentose, sparsely to densely covered with long, straight or sinuous, 

briefly swollen-based, diagonally or vertically oriented spines, these flattened or triangular in cross-section, 

whitish-brown proximally, black or brown distally, with tomentose margins (rarely without spines); rachillae 

19(7–33), brown tomentose initially; proximal rachillae 10.8(5.0–17.5) cm long, 1.3(0.7–1.9) mm wide; 

stamens 6–7; fruits 15.4(10.6–21.1) mm long, 9.5(7.4–12.4) mm wide, the surfaces uneven with numerous, 

subepidermal, short, often branching (Y-shaped) fibers; fruiting corollas less than one quarter as long as fruits, 

not or scarcely splitting, tending to remain cupular; endocarps narrowly ellipsoid with rounded apices, the 

pores lateral. 

Taxonomic notes:—Specimens included in this species were placed by both Wessels Boer (1965) and 

Henderson (1995) within a widespread Desmoncus orthacanthos Martius. This is a mistake, and the two 

species are quite distinct, especially in their fruits. The specimen from W illustrated in Dahlgren (1959, plate 

186) is not the type of D. horridus, and is anyway now destroyed. 

Subspecific variation:—There is geographic disjunction and specimens occur in five regions—a coastal 

region stretching from Tobago and Trinidad southeast through the Guianas to Brazil (Maranhão, Pará, 

Tocantins); an Orinoco region comprising the Orinoco river and its tributaries north and west of the Guayana 

Highland in Venezuela and reaching adjacent Colombia; an upper Rio Negro region, along the river and its 

main tributaries in Brazil and adjacent Venezuela and Colombia; a Pantanal region in Brazil and adjacent 

Bolivia; and a western Amazon region in Ecuador and Brazil (Acre). For most variables there are too few data 

to test for differences amongst these different regions. However, ANOVA shows that for pair wise comparison 

probabilities, seven variables (rachis length, basal pinna width, peduncular bract length, peduncle width, 

rachilla length, number of rachillae, fruit length) differ significantly (P

Key to the subspecies of Desmoncus horridus 

1 Fruits 17.7(15.5–21.1) mm long; upper Rio Negro region of Brazil, Colombia, and Venezuela and western Amazon 

region of Ecuador and Brazil (Acre)............................................................................................................................. 2 

- Fruits 15.0(13.7–19.0) mm long; all other areas .......................................................................................................... 3 

2 Peduncles 3.2(2.3–4.8) mm wide; rachillae 11(7–17); upper Rio Negro region of Brazil, Colombia, and Venezuela.. 

.............................................................................................................................................................. subsp. palustris 

- Peduncles 9.8(6.1–12.0) mm wide; rachillae 23(18–27); western Amazon region of Ecuador and Brazil (Acre) ....... 

......................................................................................................................................................... subsp. occidentalis 

3 Rachillae 14(11–16); Pantanal region in Brazil and adjacent Bolivia ............................................... subsp. prostratus 

- Rachillae 21(10–33); all other areas ............................................................................................................................ 4 

4 Peduncular bracts 19.5(10.5–27.0) cm long; Río Orinoco and its tributaries north and west of the Guayana Highland 

in Colombia and Venezuela ................................................................................................................ subsp. apureanus 

- Peduncular bracts 32.2(20.5–47.0) cm long; coastal regions from Tobago and Trinidad southeast through the Guianas 

to Brazil (Maranhão, Pará, Tocantins) .......................................................................................... subsp. horridus 

5.1. Desmoncus horridus subsp. horridus 

Desmoncus hartii Bailey (1947: 369). Type:—TRINIDAD. Mayaro, March 1922, L. Bailey 619a (holotype BH!). 

Desmoncus brittonii Bailey (1947: 371). Type:—TRINIDAD. Arena Road, 7 March 1946, L. Bailey 172 (holotype BH!). 

Desmoncus tobagonis Bailey (1947: 371). Type:—TOBAGO. Mile End, 22 February 1911, W. Broadway 4077 (holotype 

NY!, isotypes BH!, K!). 

Desmoncus demeraranus Bailey (1949: 181). Type:—GUYANA. River Demerara, near Craig village, between 

Georgetown and Atkinson Field, 20 February 1948, L. Bailey 418 (holotype BH!). 

Inflorescences peduncular bracts 32.2(20.5–47.0) cm long; rachillae 24(15–33); fruits 14.2(10.6–17.9) mm 

long. 

Distribution and habitat:—From 11°15’N–3°48’S and 44°15’–62°22’W in Trinidad and Tobago, 

Venezuela (Delta Amacuro), Guyana, Surinam, French Guiana, and Brazil (Maranhão, Pará, Tocantins) at 

23(0–50) m elevation, mostly in coastal areas in open places, often along river margins (Fig. 6). 

5.2. Desmoncus horridus subsp. apureanus (Bailey) Henderson, comb. & stat. nov. 

Basionym: Desmoncus apureanus Bailey (1949: 183). Type:—VENEZUELA. Apure: road to Churruscao, west of San 

Fernando de Apure, 15 April 1939, C. Chardon s.n. (holotype BH!). 

Desmoncus velezii Bailey (1949: 186). Type:—VENEZUELA. Apure: Puerto Paez, 5 April 1946, I. Velez 2308 (holotype 

US!, isotype BH!). 

Desmoncus multijugus Steyermark (1951: 85). Type:—VENEZUELA. Bolívar: Tumeremo, 18 December 1944, J. 

Steyermark 60968 (holotype F!). 


long. 

Distribution and habitat:—From 5°24’–7°53’N and 61°00’–71°39’W in Venezuela (Apure, Barinas, 

Bolívar) and Colombia (Casanare, Vichada) at 107(30–305) m elevation, usually in riparian habitats 

especially gallery forest along the Río Orinoco and its tributaries or in shrubby savannas (Fig. 6). 

5.3. Desmoncus horridus subsp. occidentalis Henderson, subsp. nov. (Appendix IV Plates 1–4) 

A subspeciebus aliis fructibus maioribus, pedunculis longioribus atque rachillis magis numerosis differt. 

Type:—ECUADOR. Napo: Lago Garza–Yagu on opposite side of Río Napo from Añangu, 0°32’S, 76°26’W, 300 m, 9 

August 1985, H. Balslev, A. Barfod, A. Henderson, F. Skov & A. Argüello 60752 (holotype NY!, isotypes AAU!, 

QCA n.v.). 


HENDERSON


long. 

Distribution and habitat:—From 0°02’N–7°45’S and 72°45’–76°26’W in the western Amazon region in 

Colombia, Ecuador, and Brazil (Acre) at 267(200–300) m elevation along river or lake margins in black or 

white water areas (Fig. 6). 

Taxonomic notes:—A specimen (Bernal 2033) from Colombia (Putumayo), not included in the above 

description, has narrow, lanceolate pinnae and a densely spiny peduncular bract (like D. polyacanthos), and 

fruits with Y-shaped fibers. It is sympatric with subsp. occidentalis and D. polyacanthos, and may be a hybrid. 

FIGURE 6. Distribution maps of Desmoncus horridus subsp. horridus, D. horridus subsp. apureanus, D. horridus 

subsp. occidentalis, and D. horridus subsp. palustris. 

5.4. Desmoncus horridus subsp. palustris (Trail) Henderson, comb. & stat. nov. 

Basionym: Desmoncus palustris Trail (1876: 353). Atitara palustris (Trail) Kuntze (1891: 727). Type:—BRAZIL. 

Amazonas: Rio Padauiri, 28 June 1874, J. Trail 1087/LXXXI (holotype K!, isotype BM!). 


long. 

Distribution and habitat:—From 2°19’N–0°25’S and 62°55’–68°25’W in Colombia (Guainía), Brazil 

(Amazonas), and Venezuela (Amazonas) at 107(80–140) m elevation, along margins of the upper Rio Negro 

and its blackwater tributaries, growing in water at least part of the year (Fig. 6). 



Subspecific variation:—Rachis spines are less developed in subsp. palustris than in other subspecies, 

and in some cases are abaxial and appear short and recurved, almost like those of Desmoncus polyacanthos. 

See also notes under D. interjectus. 

Although Desmoncus riparius is an older name than Desmoncus palustris, its type specimen ((Desmoncus 

riparius Spruce (1869: 156). Atitara riparia (Spruce) Kuntz (1891: 727). Type:—VENEZUELA. Amazonas: 

Río Negro, San Carlos de Río Negro, October 1854, R. Spruce 46 (holotype K!)) may be of hybrid origin. A 

group of specimens (Clark 7982, Liesner 8940, Spruce 46, type of D. riparius) from Venezuela (Amazonas), 

not included in the above description, appear intermediate between subsp. palustris and D. polyacanthos, both 

of which occur in this area. These intermediates resemble D. polyacanthos but have scarcely spiny peduncular 

bracts and fruit surfaces with Y-shaped fibers. Another specimen (Liesner 6274) is from the same locality but 

much smaller and with a non-spiny sheath. 

5.5. Desmoncus horridus subsp. prostratus (Lindman) Henderson, comb. & stat. nov. 

Basionym: Desmoncus prostratus Lindman (1900: 8). Atitara prostrata (Lindman) Barbosa Rodrigues 1902: 75. 

Type:—BRAZIL. Mato Grosso: Santa Cruz da Barra, June 1894, C. Lindman 2827 (holotype S!). 

Inflorescences peduncular bracts no data; rachillae 14(11–16); fruits 14.7(14.6–14.7) mm long. 

Distribution and habitat:—From 14°45’–15°20’S and 57°11’–62°20’W in Bolivia (Santa Cruz) and 

Brazil (Mato Grosso) at 178(150–250) m elevation, mostly in the Pantanal region in gallery forest, forest 

patches in savanna, shrubby forest, or semideciduous forest, often in flooded areas (Fig. 7). 

Two specimens, not included in the above description, are unusual and may be hybrids. One (Krapovickas 

31940) appears intermediate between subsp. prostratus and possibly D. polyacanthos. The second (Cid 4636) 

is from the north of the main range of subsp. prostratus and is reported to come from “campo natural cerrado”. 

6. Desmoncus interjectus Henderson, sp. nov. (Appendix IV Plate 5) 

A speciebus aliis spinis secus costis bracteae peduncularis praedita differt. 

Type:—COLOMBIA. Amazonas: Río Yarí, cerca a la desembocadura de la quebrada El Mochilero, 120–200 m, 21 April 

1986, G. Galeano, J. Torres, J. Huitoto & B. Plazas 1092 (holotype COL!, isotype NY!). 

Plants 6 m tall; stems 2.1(1.7–2.8) cm diameter. Leaf petioles 2.9(1.0–10.5) cm long; rachises 125.0 cm long, 

8.2(6.2–11.0) mm wide, the spines usually

Distribution and habitat:—From 0°00’–0°37’S and 70°11–72°26’W in Colombia (Amazonas, Vaupés) 

at 194(160–250) m elevation along margins of blackwater streams (Fig. 7). 

Taxonomic notes:—Except for its short, recurved, swollen-based rachis spines and peduncular bracts, D. 

interjectus strongly resembles D. horridus subsp. palustris. This subspecies, as noted above, has rachis spines 

which are less developed than in other subspecies, and in some cases are abaxial and appear short and 

recurved, almost like those of D. polyacanthos. In fact, D. interjectus appears more similar to D. horridus than 

to other species with recurved rachis spines, and its recurved rachis spines may be secondarily derived from 

straight spines. 

FIGURE 7. Distribution maps of Desmoncus horridus subsp. prostratus, D. interjectus, D. kunarius, and D. latisectus 

7. Desmoncus kunarius de Nevers ex Henderson, sp. nov. (Appendix IV Plates 6–12) 

A speciebus aliis crusta fructuum fibris subepidemalibus numerosis, brevibus atque obliquis tuberculata differt. 

Type:—PANAMA. Panama: El Llano–Cartí road, 16–18½ km by road N of Pan. Am. Hwy. at El Llano, 400–450 m, 28 

March 1974, M. Nee & E. Tyson 10982 (holotype PMA!, isotype MO!). 

Plants height no data; stems 3.3(3.1–3.6) cm diameter. Leaf petioles 4.5(3.0–5.5) cm long; rachises 100.0 cm 

long, 14.2(11.6–18.5) mm wide, the spines usually >1 cm long, mostly adaxial or lateral, straight with briefly 

swollen bases; pinnae 19 per side of rachis, without long, filiform apices, with an adaxial beard of spines at 

the bases, without spinules or dense tomentum at the bases adaxially; basal pinna 29.5 cm long, 4 cm wide; 



cirri well-developed, with acanthophylls, without spines abaxially, with intermediate acanthophylls present 

(i.e., distalmost pair of pinnae reflexed as acanthophylls and with swollen bases and/or proximalmost 

acanthophylls like vestigial pinnae), without a wide gap between pinnae and acanthophylls. Inflorescences 

with the rachis ridged, not twisted, much thicker than the numerous, closely spaced and spirally or irregularly 

arranged rachillae, each rachilla not or only briefly adnate to the rachis, subtended by an acute bracteole and 

with a well-developed axillary pulvinus; peduncles 8.8(7.8–10.6) mm wide; peduncular bracts 48.5 cm long, 

broad, the surfaces ribbed, brown tomentose, densely covered with long, straight or sinuous, briefly swollenbased, 


proximally, black or brown distally, with tomentose margins; rachillae 21(20–22), glabrous or scarcely 

tomentose initially; proximal rachillae 18.5(17.5–19.5) cm long, 2.5(2.1–2.9) mm wide; stamens 12; fruits 

25.1(23.9–27.4) mm long, 21.7(20.1–24.1) mm wide, the surfaces bumpy from numerous, subepidermal, 

short, oblique fibers; fruiting corollas less than one quarter as long as fruits, splitting irregularly into 3 lobes, 

the lobes often splitting again; endocarps obovoid with rounded apices, the pores lateral. 

Distribution and habitat:—From 9°17’–9°21’N and 78°55’–79°47’W in central Panama at 270(150– 

425) m elevation in lowland rainforest (Fig. 7). 

Taxonomic notes:—Desmoncus kunarius is characterized by its large size and unusually large fruits. 

8. Desmoncus latisectus Burret (1934: 215). 

Lectotype (here designated):—BOLIVIA. Beni: Trinidad, Missiones Guarayos, 300 m, September 1926, E. Werdermann 

2508 (S!, isolectotype MO!; the holotype in B was destroyed). 

Desmoncus kuhlmannii Burret (1938: 267). Lectotype (here designated):—BOLIVIA. Beni: Riberalta, 28 November 

1923, J. Kuhlmann 522 (R!; the holotype of B was destroyed). 

Plants 2.8(2.0–4.0) m tall; stems 0.9(0.5–1.4) cm diameter. Leaf petioles 2.0(1.0–3.0) cm long; rachises 

34.1(29.0–40.0) cm long, 3.1(2.2–3.8) mm wide, the spines usually

latisectus has narrower stems, shorter petioles, narrower rachises, fewer divisions, narrower basal pinnae), 

and their habitat also appears different. Desmoncus latisectus grows mostly at lower elevations in northeastern 

Bolivia in savanna-forest margins, whereas D. leptoclonos grows at higher elevations in gallery forest in the 

Cerrado in Brazil and Paraguay. 

9. Desmoncus leptoclonos Drude (1881: 315). 

Atitara leptoclona (Drude) Barbosa Rodrigues, 1902: 76. Type:—BRAZIL. Goiás: between Goiás and Cuiabá, 

November–December 1844, H. Weddell 2900 (holotype P!). 


rachises 47.9(17.0–77.0) cm long, 5.7(4.2–7.5) mm wide, the spines usually

Plants height no data; stems 1.2(0.7–1.6) cm diameter. Leaf petioles 1.5(0.7–2.2) cm long; rachises 


11. Desmoncus madrensis Henderson, sp. nov. (Appendix IV Plate 15) 

A Desmonco loretano bractea pedunculari spinescenti differt; a D. polyacanthos fructuum crusta fibris Y-formis differt. 

Type:—PERU. Madre de Dios: Manu, Pakitza, Parque Nacional de Manu, trocha Pacal, 11˚53’S, 70˚58’W, 400 m, 24 

February 1990, F. Chávez 626 (holotype NY!, isotypes CUZ n.v., USM n.v.). 



acts 24.2(14.6–37.0) cm long, narrow, elongate, ribbed, scarcely brown tomentose, without spines (rarely 

with few spines); rachillae 5(3–7), glabrous or scarcely tomentose initially; proximal rachillae 5.3(2.0–9.0) 

cm long, 0.8(0.5–1.5) mm wide; stamens 6; fruits 10.9(8.9–15.7) mm long, 7.1(5.5–10.9) mm wide, fruit 

surfaces uneven with numerous, subepidermal, short, often branching (Y-shaped) fibers; fruiting corollas 

splitting irregularly into 3 lobes, the lobes often splitting again; endocarps narrowly ellipsoid with rounded 

apices, the pores lateral. 

Subspecific variation:—Specimens occur in the western and central Amazon region in Colombia, 

Ecuador, Peru, Bolivia, and Brazil. Based on geography, as well as sheaths, number of pinnae, and cirri 

development, it is possible to recognize four subgroups—one from subAndean regions of southern Peru and 

Bolivia having finely and densely spiny sheaths, numerous, linear or lanceolate pinnae, and well-developed 

cirri; the second from the western Amazon region, mostly from the southwestern Amazon region of Brazil but 

also in adjacent Peru and Bolivia having non- or scarcely spiny sheaths, few, ovate pinnae, and poorlydeveloped 

cirri; the third from the western Amazon region of Peru and in adjacent Colombia and Brazil 

having non-spiny sheaths, few, ovate pinnae, and usually well-developed cirri; and the fourth from the 

western Amazon region in Colombia, Ecuador, and Peru, having non-spiny sheaths, numerous, lanceolate 

pinnae, and well-developed cirri. ANOVA shows that for pair wise comparison probabilities, six variables 

(plant height, petiole length, rachis length, rachis width, basal pinna length, basal pinna width) differ 

significantly (P

and in Martius (1823–1837) as “fluvii Solimoës”, presumably in Brazil (Amazonas). All the specimens cited 

here come from further west than the most westward point of Martius’ journey in modern-day Colombia, with 

the exception of Aguirre-Galviz 1152 from Colombia (Amazonas). 

12.2. Desmoncus mitis subsp. ecirratus Henderson, subsp. nov. (Appendix IV Plate 16) 

A subspeciebus aliis pinnis paucis ovatis atque cirro normaliter absenti differt. 

Type:—BRAZIL. Rondônia: road to casserite mine, north bank of Rio Madeira in Serra dos Tres Irmãos 8 km above 

Mutumparaná, 5 July 1968, G. Prance, E. Forero, L. Coêlho, J. Ramos & L. Farias 5636 (holotype INPA!, isotype 

NY!). 

Leaf sheaths non-spiny, rarely sparsely spiny; rachises 20.9(8.0–41.0) cm long; pinnae ovate, 3(2–5) per side 

of rachis; cirri usually absent, the rachis terminating beyond the distalmost pair of pinnae in a short ‘stub’. 

Distribution and habitat:—From 3°30’–10°58’S and 56°00’–73°14’W in the south–central Amazon 

region of Peru (Ucayali), Brazil (Acre, Amazonas, Mato Grosso, Rondônia) and Bolivia (Beni, Pando) with 

an outlier in Peru (Loreto) at 177(120–300) m elevation in lowland rainforest (Fig. 9). 

Taxonomic notes:—This taxon was mistakenly referred to by Henderson (1995) as Desmoncus mitis var. 

leptoclonos, based on D. leptoclonos Dammer. However, this name is pre-occupied by D. leptoclonos Drude. 

Subspecific variation:—The outlying specimen (Asplund 14687) from Loreto, Peru, has much wider 

pinnae than other specimens. Some specimens are described on labels as being non–climbers. On some 

specimens (e.g., Moreno 163, Schunke 15033, Nee 34990, Forero 6384) there are some leaves with weakly 

developed cirri as well as leaves with the more usual poorly-developed cirri; on a few other specimens there 

are weakly developed cirri. Specimens from the eastern part of the range, from eastern Acre, Rondônia, Mato 

Grosso, and adjacent Bolivia usually have 2 pinnae per side of the shorter rachis, and these have very few 

spinules at the bases. Specimens from the western part of the range, from western Acre and adjacent Peru tend 

to have 3 pinnae per side of the longer rachis, and usually have spinulose pinnae bases. These specimens 

overlap with those of D. mitis subsp. leptospadix. 

12.3. Desmoncus mitis subsp. leptospadix (Martius) Henderson, comb. & stat. nov. 

Basionym: Desmoncus leptospadix Martius (1844: 52). Atitara leptospadix (Martius) Kuntze (1891: 727). Desmoncus 

mitis var. leptospadix (Martius) Henderson (1995: 228). Type:—PERU. Loreto: Maynas, Yurimaguas, no date, E. 

Poeppig 2207 (holotype G n.v., holotype image!). Epitype (here designated):—PERU. Loreto: Santa Rosa, lower 

Río Huallaga below Yurimaguas, ca. 125 m, 1–5 September 1929, E. Killip & A. Smith 28807 (epitype NY!, 

isoepitype US!). 

Leaf sheaths non-spiny, rarely sparsely spiny; rachises 35.0(17.2–76.0) cm long; pinnae ovate, 6(3–11) per 

side of rachis; cirri usually well-developed. 


Colombia (Amazonas), Peru (Amazonas, Cuzco, Huánuco, Loreto, Ucayali), and Brazil (Acre, Amazonas) at 

166(100–400) m elevation in lowland rainforest (Fig. 9). 

Taxonomic notes:—Henderson (1995) referred to this taxon as Desmoncus mitis var. leptospadix 

(Martius) Henderson, based on D. leptospadix Martius. The type of this is sterile and an epitype is therefore 

designated. 

Subspecific variation:—Most specimens from Maynas, Loreto, Peru have multiple (2–3) inflorescences 

at a node. Two specimens (Prance 12310, 12490), both from the same locality in western Acre, Brazil have 

larger fruits than usual; others from the same area have normal fruits. Most specimens from western Acre 

have smaller leaves than usual. A few specimens from the western part of the range have spiny peduncular 



acts (e.g., Schunke 15629) or spiny sheaths (e.g., Huashikat 2309). Three specimens from Loreto, Peru 

(Tessmann 5236, Vásquez 8325, 8329) have poorly-developed cirri. 

One unusual specimen (Simpson 698) from Loreto, possibly a mixed collection, has leaves like those of 

subsp. mitis and infructescences like those of D. polyacanthos. Another specimen from Loreto (Kvist 1156) 

appears intermediate between subsp. leptospadix and D. polyacanthos and may be a hybrid. A specimen from 

Huánuco (Listabarth 1110589) may also be a hybrid. The easternmost specimen (Rabelo 79), from from the 

Rio Urubu near Manaus, is outside the main range of subsp. leptospadix and has completely smooth 

peduncular bracts and slender rachillae with some adnation. It may belong here; it is not included in the above 

description but is mapped. 

FIGURE 9. Distribution maps of Desmoncus mitis subsp. ecirratus, D. mitis subsp. letpospadix, D. mitis subsp. 

rurrenabaquensis, and D. moorei. 

12.4. Desmoncus mitis subsp. rurrenabaquensis (Henderson) Henderson, comb. & stat. nov. 

Basionym: Desmoncus mitis var. rurrenabaquensis Henderson (1995: 228). Type:—PERU. Madre De Dios: Explorer's 

Inn on Río Tambopata, 12°50'S, 69°17'W, ca. 250 m, 6 November 1991, A. Henderson & F. Chávez 1640 (holotype 

USM!, isotypes, CUZ!, NY!). Epitype (here designated):—BOLIVIA. La Paz: Franz Tamayo, Parque Nacional 

Madidi, río Hondo, arroyo Negro, pica hacia la serranía de Toregua, 14°39’S 67°48’W, 340 m, 25 March 2002, A. 

Fuentes, M. Villanueva & B. Guili 4081 (epitype NY!, isoepitypes LPB n.v., MO!). 


HENDERSON

Leaf sheaths densely and finely spiny; rachises 45.1(25.0–69.0) cm long; pinnae linear or lanceolate, 19(16– 

22) per side of rachis; cirri well-developed. 

Distribution and habitat:—From 12°50’–17°24’S and 64°30’–70°50’W in subAndean areas of the 

southwestern Amazon region of Peru (Cuzco, Madre de Dios, Puno) and Bolivia (Beni, Cochabamba, La Paz, 

Santa Cruz) at 421(250–920) m elevation in lowland rainforest (Fig. 9). 

Taxonomic notes:—This taxon was referred to by Henderson (1995) as Desmoncus mitis var. 

rurrenabaquensis. The type of this is sterile and an epitype is therefore designated. 

13. Desmoncus moorei Henderson, sp. nov. (Appendix IV Plate 17) 

A Desmonco chinantlensi, quod similis, rachillis tomentosis atque staminibus paucioribus differt. 

Type:—COSTA RICA. Limón: P. N. Tortuguero, Pococi, Llanura de Tortuguero, Estación Cuatro Esquinas, 10˚32’N, 

83˚30’W, 2 m, 18 October 1989, J. Solano 21 (holotype INB!, isotype MO!). 


96.1(70.0–113.0) cm long, 9.6(6.7–12.0) mm wide, the spines usually >1 cm long, mostly adaxial or lateral, 

straight with briefly swollen bases; pinnae 18(15–20) per side of rachis, without long, filiform apices, with an 

adaxial beard of spines at the bases, with uneven surfaces at the bases adaxially, covered with spinules; basal 

pinna 18.4(16.0–20.0) cm long, 1.5(1.2–1.8) cm wide; cirri well-developed, with acanthophylls, without 

spines abaxially, with intermediate acanthophylls present (i.e., distalmost pair of pinnae reflexed as 

acanthophylls and with swollen bases and/or proximalmost acanthophylls like vestigial pinnae), without a 

wide gap between pinnae and acanthophylls. Inflorescences with the rachis ridged, not twisted, much thicker 

than the numerous, closely spaced and spirally or irregularly arranged rachillae, each rachilla not or only 

briefly adnate to the rachis, subtended by an acute bracteole and with a well-developed axillary pulvinus; 

peduncles 5.8(3.2–7.7) mm wide; peduncular bracts 37.8(25.5–54.5) cm long, broad, the surfaces ribbed, 

brown tomentose, sparsely to densely covered with long, straight or sinuous, briefly swollen-based, 


proximally, black or brown distally, with tomentose margins; rachillae 30(19–38), brown tomentose initially; 

proximal rachillae 14.2(11.5–18.0) cm long, 1.2(0.8–1.4) mm wide; stamens 6; fruits 12.6(11.1–14.3) mm 

long, 11.4(10.2–13.8) mm wide, the surfaces uneven with numerous, subepidermal, short, often branching (Yshaped) 

fibers; fruiting corollas less than one quarter as long as fruits, splitting irregularly into 3 lobes, the 

lobes often splitting again; endocarps globose to obovoid with rounded or slightly peaked apices, the pores 

lateral. 

Distribution and habitat:—From 9°35’–11°55’N and 82°36’–85°25’W on the Caribbean slope of 

Nicaragua and Costa Rica, with an outlier in the Nicoya Peninsula in Costa Rica at 138(1–700) m elevation in 

lowland rainforest and forest margins along rivers (Fig. 9). 

Taxonomic notes:—This species is named for Harold E. Moore (1917–1980) of Cornell University, 

preeminent student of palms. Hammel et al. (2003) identified the specimens included here as Desmoncus 

schippii Burret. The type of this is no longer extant and it is therefore treated here as an Excluded Name. 

Desmoncus moorei is similar to D. chinantlensis but differs in its spinulose pinnae bases, tomentose rachillae, 

and fewer stamens (6 versus 8–11). Specimens dry a distinctive dark brown color compared to D. 

chinantlensis. 

Subspecific variation:—Intermediate acanthophylls and adaxial beards of spines at the pinnae bases are 

much less well-developed than in similar species. The specimen from the Nicoya Peninsula in Costa Rica is 

tentatively included here. It has narrower pinnae than other specimens. Sterile, juvenile plants from the same 

locality have numerous, linear pinnae and appear quite distinct. However, there is only one fertile specimen 

from Nicoya. 



14. Desmoncus myriacanthos Dugand (1943: 75). 

Type:—COLOMBIA. Bolívar: Norosí–Tiquísio trail, lands of Loba, 150–600 m, 24 April 1916, H. Curran 174 

(holotype US!). 

Desmoncus isthmius Bailey (1943: 211). Type:—PANAMA. Darién: Marraganti and vicinity, 5 April 1908, R. Williams 

691 (holotype NY!, isotype US!). Synon. nov. 




with an adaxial beard of spines at the bases, without spinules or dense tomentum at the bases adaxially; basal 

pinna 22.0(18.0–27.0) cm long, 2.2(1.2–4.2) cm wide; cirri well-developed, with acanthophylls, without 

spines abaxially, with intermediate acanthophylls present (i.e., distalmost pair of pinnae reflexed as 

acanthophylls and with swollen bases and/or proximalmost acanthophylls like vestigial pinnae), without a 

wide gap between pinnae and acanthophylls. Inflorescences with the rachis ridged, not twisted, much thicker 

than the numerous, closely spaced and spirally or irregularly arranged rachillae, each rachilla not or only 

briefly adnate to the rachis, subtended by an acute bracteole and with a well-developed axillary pulvinus; 

peduncles 5.6(3.2–8.6) mm wide; peduncular bracts 26.7(20.0–34.0) cm long, broad, the surfaces ribbed, 

brown tomentose, sparsely to densely covered with long, straight or sinuous, briefly swollen-based, 


proximally, black or brown distally, with tomentose margins; rachillae 23(16–32), glabrous or scarcely 

tomentose initially; proximal rachillae 10.7(5.5–15.8) cm long, 1.3(0.8–2.2) mm wide; stamens 8–9; fruits 

15.1(12.6–21.0) mm long, 8.2(7.0–10.3) mm wide, the surfaces uneven with numerous, subepidermal, short, 

often branching (Y-shaped) fibers; fruiting corollas less than one quarter as long as fruits, not or scarcely 

splitting, tending to remain cupular; endocarps narrowly ellipsoid with rounded apices, the pores lateral. 

Distribution and habitat:—From 6°13’–11°18’N and 72°00’–80°18’W in central and eastern Panama, 

northwestern Colombia, and just reaching Venezuela (Zulia) at 73(10–125) m elevation in lowland rainforest 

(Fig. 10). 

Taxonomic notes:—Although published in the same year, 1943, Desmoncus myriacanthos predates D. 

isthmius by several months. 

Subspecific variation:—Specimens are from scattered areas but this is likely to be an artifact of 

insufficient collecting. The southernmost specimen (Forero 9080) from the Chocó region comes from one of 

the wettest areas in Colombia, in contrast to the much dryer habitat of the other specimens from northwestern 

Colombia. 

15. Desmoncus obovoideus Henderson, sp. nov. (Appendix IV Plates 18–19) 

A speciebus affinibus crusta fructuum aspera fibris subepidemalibus numerosis, longis atque brachiatis differt. 

Type:—PANAMA. Colón: Santa Rita ridge east of Transisthmian Highway, 300–500 m, 20 September 1972, A. Gentry 

6115 (holotype PMA!, isotype MO!). 

Plants 4.9(3.5–7.0) m tall; stems 1.7(1.4–2.0) cm diameter, branching no data. Leaf petioles 3.1(2.0–4.5) cm 

long; rachises length no data, 6.3(6.0–6.6) mm wide, the spines usually >1 cm long, mostly adaxial or lateral, 

straight with briefly swollen bases; pinnae number no data, without long, filiform apices, with an adaxial 

beard of spines at the bases, without spinules or dense tomentum at the bases adaxially; basal pinna 14.0 cm 

long, 4.9(4.2–5.5) cm wide; cirri well-developed, with acanthophylls, without spines abaxially, with 

intermediate acanthophylls present, without a wide gap between pinnae and acanthophylls. Inflorescences 




HENDERSON

with a well-developed axillary pulvinus; peduncles 4.3(2.8–5.7) mm wide; peduncular bracts length no data, 

broad, the surfaces ribbed, brown tomentose, sparsely to densely covered with long, straight or sinuous, 



21(20–21), glabrous or scarcely tomentose initially; proximal rachillae 8.5 cm long, 1.0 mm wide; stamens no 

data; fruits 15.8(14.0–17.9) mm long, 14.2(12.8–16.3) mm wide, the surfaces uneven with numerous, 

subepidermal, long, branching fibers; fruiting corollas less than one quarter as long as fruits, splitting 

irregularly into 3 lobes, the lobes often splitting again; endocarps broadly obovoid with flattened apices, the 

pores lateral on or near flattened apices. 

Distribution and habitat:—From 8°40’–9°30’N and 77°25’–80°29’W in central Panama, with outliers 

to the west and east, at 173(25–400) m elevation in lowland rainforest (Fig. 10). 

FIGURE 10. Distribution maps of Desmoncus myriacanthos, D. obovoideus, D. osensis, D. orthacanthos, and D. 

parvulus. 

Taxonomic notes:—Desmoncus obovoideus is similar to D. osensis and shares all character states except 

that D. obovoideus lacks data for stamen number. Desmoncus obovoideus also has shorter and wider basal 

pinna compared to D. osensis (mean of 14.0 cm long and 4.9 cm wide versus 23.2 cm long and 3.7 cm wide). 

Based on this, and geographic separation, the two are recognized as separate species. 

A few specimens have abaxial rachis spines which appear almost recurved, more like those of D. 

polyacanthos. There are two outliers, both of which are tentatively included here. The one from eastern 



Panama (McPherson 6961) is without fruits. The other (Hammel 1863), from central Panama, also has abaxial 

rachis spines which appear almost recurved. 

Subspecific variation:—In most specimens the adaxial beard of spines at the bases of the pinnae is 

poorly-developed. One specimen (Garwood 2178) is smaller than the others. 

17. Desmoncus orthacanthos Martius (1823–1837: 87). 

Atitara orthacantha (Martius) Barbosa Rodrigues (1902: 76). Type:—BRAZIL. Bahia: Rio Mucuri, no date, M. Neuwied 

s. n. (holotype M!). 

Desmoncus lophacanthos Martius (1844: 50). Atitara lophacantha (Martius) Barbosa Rodrigues (1902: 76). Type:— 

BRAZIL. Bahia: Ilhéus, 20 March 1837, B. Luschnath s.n. (holotype M!). 




without a beard of spines at the bases, without spinules or dense tomentum at the bases adaxially; basal pinna 

14.3(7.6–25.0) cm long, 2.6(1.0–4.8) cm wide; cirri well-developed, with acanthophylls, without spines 

abaxially, with intermediate acanthophylls present (i.e., distalmost pair of pinnae reflexed as acanthophylls 

and with swollen bases and/or proximalmost acanthophylls like vestigial pinnae), without a wide gap between 

pinnae and acanthophylls. Inflorescences with the rachis angular, slightly twisted, thicker than the closely 

spaced and spirally arranged rachillae, each rachilla not adnate to the rachis and with an irregular bracteole 

adnate to the rachilla and appearing displaced distally onto the rachilla, with a poorly- to well-developed 

axillary pulvinus; peduncles 3.9(2.0–7.2) mm wide; peduncular bracts 26.4(19.0–39.0) cm long, broad, the 

surfaces ribbed, brown tomentose, rarely with long, straight or sinuous, briefly swollen-based, diagonally or 

vertically oriented spines, these flattened or triangular in cross-section, whitish-brown proximally, black or 

brown distally, with tomentose margins; rachillae 14(8–22), glabrous or scarcely tomentose initially; proximal 

rachillae 10.4(5.0–20.5) cm long, 0.8(0.3–1.2) mm wide; stamens 5–7; fruits 14.4(11.4–18.4) mm long, 

10.9(8.4–15.2) mm wide, the surfaces uneven with numerous, subepidermal, short, often branching (Yshaped) 

fibers; fruiting corollas less than one quarter as long as fruits, splitting irregularly into 3 lobes, the 

lobes often splitting again; endocarps globose to obovoid with rounded or slightly peaked apices, the pores 

lateral. 

Distribution and habitat:—From 8°45’–23°01’S and 35°06’–43°28’W in the Atlantic Coastal Forest of 

Brazil at 105(0–700) m elevation, usually near the sea in restinga or scrub forest (Fig. 10). 

Taxonomic notes:—Both Wessels Boer (1965) and Henderson et al. (1995) considered Desmoncus 

orthacanthos to be much more widespread. This was based on a misinterpretation, and it is here considered to 

be confined to the Atlantic Coastal Forest of Brazil (see notes under Desmoncus horridus). Desmoncus 

orthacanthos has an unique inflorescence structure with each rachilla with an irregular, adnate bracteole that 

appears displaced distally onto the rachilla (Fig. 2D). Pinnae spines are also distinctive. All specimens have at 

least some pinnae with one or rarely two large spines right at the very base of the pinnae on the abaxial 

surface. 

Subspecific variation:—There is no geographic disjunction, except for two outlying specimens from 

Pernambuco. There is geographical variation in this species. Regression shows there are significant (P

16. Desmoncus osensis Henderson, sp. nov. (Appendix IV Plates 20–21) 

A Desmonco obovoideo pinnis basalibus longioribus atque angustioribus diifert. 

Type:—COSTA RICA. Puntarenas: Golfito, Distrito Golfito, Refugio de Vida Silvestre Golfito, camino a La Gamba, 

8°40’N 83°11’W, 94 m, 9 October 2008, R. Aguilar 11417 (holotype NY!, isotypes CR n.v., USJ n.v.). 

Plants height no data; stems 1.6(1.2–2.6) cm diameter, branching no data. Leaf petioles 2.7(2.0–3.5) cm long; 

rachises 85 cm long, 7.1(6.4–7.8) mm wide, the spines usually >1 cm long, mostly adaxial or lateral, straight 

with briefly swollen bases; pinnae 13 per side of rachis, without long, filiform apices, with an adaxial beard of 

spines at the bases, without spinules or dense tomentum at the bases adaxially; basal pinna 23.2(18.0–26.0) 

cm long, 3.7(2.4–5.0) cm wide; cirri well-developed, with acanthophylls, without spines abaxially, with 

intermediate acanthophylls present, without a wide gap between pinnae and acanthophylls. Inflorescences 



with a well-developed axillary pulvinus; peduncles 4.5(4.1–4.8) mm wide; peduncular bracts length no data, 

broad, the surfaces ribbed, brown tomentose, sparsely to densely covered with long, straight or sinuous, 



19(18–19), glabrous or scarcely tomentose initially; proximal rachillae length no data, 1.6 mm wide; stamens 

9; fruits 15.1(13.9–17.3) mm long, 13.6(11.0–15.9) mm wide, the surfaces uneven with numerous, 

subepidermal, long, branching fibers; fruiting corollas less than one quarter as long as fruits, splitting 

irregularly into 3 lobes, the lobes often splitting again; endocarps broadly obovoid with flattened apices, the 

pores lateral on or near flattened apices. 

Distribution and habitat:—From 8°36’–9°31’N and 83°11’–84°06’W in the Osa Peninsula and adjacent 

areas in Costa Rica at 145(50–350) m elevation in lowland rainforest (Fig. 10). 

Taxonomic notes:—Desmoncus osensis is similar to D. obovoideus and shares all character states except 

that D. obovoideus lacks data for stamen number. Desmoncus osensis also has longer and narrower basal 

pinna compared to D. obovoideus (mean of 23.2 cm long and 3.7 cm wide versus 14.0 cm long and 4.9 cm 

wide). Based on this, and geographic separation, the two are recognized as separate species. 

Subspecific variation:—Hammel et al. (2003) had only two specimens (Aguilar 290, Morales 6333) 

available from the Osa Peninsula, and identified them as Desmoncus sp. A. They considered that these two 

specimens were different from one another and possibly not conspecific. In the present study, seven 

specimens have been examined. They are indeed variable, and in some the adaxial beard of spines at the bases 

of the pinnae are poorly-developed. Some of this variation may be due to the age of the leaves represented by 

the specimens. There is also variation in fruit size. Without more specimens it is not possible to resolve this 

problem. 

18. Desmoncus parvulus Bailey (1948: 115). 

Type:—GUYANA. Potaro–Siparuni: Tumatumari, 18 June–8 July 1921, H. Gleason 164 (holotype NY! isotype US!). 

Desmoncus kaieteurensis Bailey (1948: 115). Type:—GUYANA. Potaro-Siparuni: trail from Tukeit to Kaiatuk Plateau, 

29 April 1944, B. Maguire & D. Fanshawe 23093 (holotype BH!, isotypes K!, NY!). 



acanthophylls, with few spines abaxially throughout, with no intermediate acanthophylls present, usually with 

a wide gap between pinnae and acanthophylls (i.e., gap wider than that between adjacent acanthophylls). 

Inflorescences with the rachis angular, slightly twisted, thicker than the few to numerous, closely spaced and 

spirally arranged rachillae, each rachilla not (or very rarely) adnate to the rachis, subtended by an acute 

bracteole and with a well-developed axillary pulvinus; peduncles 1.6(1.0–3.1) mm wide; peduncular bracts 

12.5(8.5–17.0) cm long, broad, ridged, densely covered with short, straight, swollen-based, vertically oriented 

spines, these terete, whitish-brown proximally, brown distally, without tomentum; rachillae 9(7–14), glabrous 

or scarcely tomentose initially; proximal rachillae 4.6(3.0–6.2) cm long, 0.7(0.5–0.9) mm wide; stamens 6; 

fruits 11.1(8.9–14.3) mm long, 9.4(7.1–11.2) mm wide, the surfaces uneven with numerous, subepidermal, 

long, branching fibers; fruiting corollas less than one quarter as long as fruits, splitting irregularly into 3 lobes, 

the lobes often splitting again; endocarps globose to obovoid with rounded or slightly peaked apices, the pores 

lateral. 

Distribution and habitat:—From 4°05’S–8°20’N and 45°19’–72°12’W in the eastern and central 

Amazon region in Brazil, the Guianas, Venezuela, and Colombia at 209(5–650) m elevation in lowland 

rainforest (Fig. 10). 

Taxonomic notes:—The name Desmoncus macroacanthos Martius was used by Wessels Boer (1965) for 

specimens included here. This name is here treated under Excluded Names; the sterile type has spiny leaf 

sheaths and non–spiny cirri and does not appear to represent this species. The name Desmoncus 

phoenicocarpus Barbosa Rodrigues was used for this species by Henderson (1995). The type is an illustration, 

also having spiny leaf sheaths and non–spiny cirri, and is also here treated under Excluded Names. 

Subspecific variation:—Specimens come from widely scattered localities but this may be an artifact of 

insufficient collecting. Two specimens from the Rio Tapajos in Brazil are unusual. One (Maciel 167) has very 

short petiole, numerous, small pinnae, and cirrus without spines abaxially throughout; the second (Anderson 

10577) has very large pinnae. These may be of hybrid origin. 

19. Desmoncus polyacanthos Martius (1823–1837: 85). 

Atitara polyacantha (Martius) Kuntze (1891: 726). Lectotype (here designated):—BRAZIL. Bahia: without locality, no 

date, C. Martius s.n. (M!). 

Desmoncus oxyacanthos Martius (1823–1837: 88). Desmoncus polyacanthos var. oxyacanthos (Martius) Drude (1881: 

314). Atitara oxyacantha (Martius) Kuntze (1891: 727). Type:—BRAZIL. Rio de Janeiro: Sebastianopolis, no date, 

M. Neuwied s.n. (holotype M!). 

Desmoncus ulei Dammer (1907: 129). Lectotype (here designated):—BRAZIL. Amazonas: Manaus, Rio Negro, 

February 1901, E. Ule 5388 (MG!). 

Desmoncus campylacanthus Burret (1934: 210). Lectotype (here designated):—BRAZIL. Rio de Janeiro: Serra de Bica, 

Cascadura, 10 December 1882, A. Glaziou 14366 (P!, holotype B, isolectotypes C n.v., K!; the holotype at B was 

destroyed). 

Desmoncus prestoei Bailey (1943: 215). Type:—TRINIDAD. “Hort. Trin.”, no date, W. Broadway 5568 (holotype BH!). 

Desmoncus peraltus Bailey (1947: 373). Type:—TRINIDAD. Arena, 17 January 1946, L. Bailey 101 (holotype BH!). 

Desmoncus maguirei Bailey (1948: 108). Type:—SURINAME. Kwakoegron, 19 October 1947, B. Maguire & G. Stahel 

25011 (holotype NY!, isotypes BH!, U!). 

Desmoncus mirandanus Bailey (1949: 183). Type:—VENEZUELA. Miranda: Cárdenas (Guinand Estate), Siquire 

Valley, 400–800 m, 6 April 1917, H. Pittier 7078 (holotype BH!, isotype US!). 



arranged rachillae, each rachilla subtended by an acute bracteole and with an axillary pulvinus; peduncles 

3.5(1.6–12.4) mm wide; peduncular bracts 26.3(16.5–34.0) cm long, broad, sparsely to densely covered with 

short, markedly swollen-based, diagonally oriented spines, these triangular in cross-section, whitish-brown 

proximally, brown distally, with tomentose margins, rarely spines few or absent; rachillae 15(5–37), glabrous 

or scarcely tomentose initially; proximal rachillae 7.4(3.3–13.0) cm long, 1.1(0.6–2.0) mm wide; stamens 5– 

6; fruits 16.4(11.2–23.5) mm long, 11.9(7.9–17.9) mm wide, the surfaces smooth, without any apparent 

subepidermal fibers; fruiting corollas less than one quarter as long as fruits, splitting irregularly into 3 lobes, 

the lobes often splitting again; endocarps globose to obovoid with rounded apices, the pores lateral, not 

equidistant, the sterile pores closer latitudinally. 

Distribution and habitat:—From 10°37’N–21°31’S and 35°09’–78°38’W in Trinidad, Venezuela, the 

Guianas, Brazil (including the Atlantic Coastal Forest), Colombia, Ecuador, Peru, and Bolivia at 205(0–1000) 

m elevation in a variety of habitats including lowland rainforest on terra firme, flooded forest, campina, 

restinga, or scrub forest near the sea (Fig. 11). Read (1979) also included the Lesser Antillean island of St. 

Vincent in the distribution of this species, but only one, sterile specimen from there has been seen. There is 

another specimen at P labelled “Martinique” but without more precise locality. 

FIGURE 11. Distribution maps of Desmoncus polyacanthos, D. prunifer, D. pumilus, and D. setosus. 

Taxonomic notes:—Desmoncus polyacanthos Martius was emended by Drude (1881) and excluded by 

Burret (1934). However there seems no reason for these designations. Martius’s (1823–1837) description and 

illustration, based on an extant type collected by Martius himself, perfectly represent this widespread species. 



Subspecific variation:—There is geographic disjunction and specimens occur in two regions—the 

Atlantic Coastal Forest of Brazil, and all other areas. Specimens from the Atlantic Coastal Forest differ 

significantly from others from Brazil in only three variables (rachis length, number of pinnae, rachilla 

width)(t-test, P

acts and are referred to the ulei morphotype. 

In northwestern Peru (Amazonas) and just reaching extreme southern Ecuador (Zamora-Chinchipe), 

specimens are large with ovate pinnae, densely spiny peduncular bracts, and large fruits. They are referred to 

as the amazonas morphotype. 

In central Peru (San Martín) specimens are extremely variable. A single specimen from southern San 

Martín (Schunke 6927) is larger, with densely spiny sheaths and long, almost linear pinnae (sanmartin 

morphotype). A second specimen from the northern part of the department (Williams 6661) is completely 

different, with smaller, ovate pinnae and a finely spiny peduncular bract (ulei morphotype). 

In south-central Peru (Pasco) there are two specimens (Henderson 3009, Smith 3791) which are 

considerably larger than others and have large stems, long, lanceolate pinnae, and large inflorescences with 

densely spiny peduncular bracts (pasco morphotype). A specimen from Ucayali (Gentry 25466) is similar, but 

another from adjacent Junín (Macbride 5470) has more ovate pinnae and is referred to the large morphotype. 

In western Acre, Brazil, and adjacent Peru (Ucayali) specimens are of the large morphotype. One 

specimen (Henderson 1675) from western Acre on the border with Amazonas is of the ulei morphotype. Five 

specimens from eastern Acre (Coêlho 28, Daly 9387, 11986, Figueiredo 342, 615) resemble those of the ulei 

morphotype except for their pinnae with spinulose bases. They may represent hybrids with the sympatric D. 

mitis subsp. ecirratus. 

In Madre de Dios, Peru and western Bolivia specimens are all of ulei morphotype, although there is much 

variation in fruit size. 

Specimens from the Atlantic Coastal Forest of Brazil (polyacanthos morphotype) occur in two areas, with 

northern outliers in Ceará, Alagoas, and Pernambuco. There are too few specimens from there to test for 

differences between these areas. Along the Atlantic coast, D. polyacanthos occurs slightly more inland than 

the sympatric D. orthacanthos. 

20. Desmoncus prunifer Poepp. ex Martius (1823-1837: 148). 

Atitara prunifera (Poepp. ex Martius) Kuntze (1891: 727). Desmoncus polyacanthos var. prunifer (Martius) Henderson, 

1995: 233. Type:—PERU. Loreto: Maynas, Yurimaguas, December 1830, E. Poeppig 2148 (holotype G n.v., 

holotype image!, isotypes GOET n.v., P!). Epitype (here designated):—PERU. Loreto: Yurimaguas, lower Río 

Huallaga, ca. 135 m, 22 August–9 September 1929, E. Killip & A. Smith 28045 (epitype US!). 

Plants height no data; stems 0.8(0.7–0.8) cm diameter, branching no data. Leaf petioles 3.8(2.2–5.4) cm long; 

rachises 46.5 cm long, 3.2(2.8–3.6) mm wide, the spines usually

Taxonomic notes:—This name, as Desmoncus polyacanthos var. prunifer (Poeppig ex Martius) 

Henderson, based on D. prunifer Poeppig ex Martius, was mistakenly used by Henderson (1995) for 

specimens here identified as Desmoncus vacivus. The type of D. prunifer was given by Henderson as Poeppig 

2148, although this specimen was not cited by Martius. Three sheets of Poeppig 2148 are extant at G but 

comprise leaves only. Two other specimens were at W but are both now destroyed, although images are 

extant. Neither is labeled as Poeppig 2148, although both appear to have been collected by Poeppig at the type 

locality. One is illustrated by Dahlgren (1959, plate 194, negative 31323) and appears to be the same species 

as Poeppig 2148. The second specimen at W, an image of which is present at NY (negative 31325), also 

appears to be the same species, and has a densely spiny peduncular bract. 

The images of these five specimens strongly resemble one another and three recently-collected specimens 

(Croat 18115, Killip 28045, Williams 785) from the same general locality, including one from Yurimaguas 

itself. Based on this, they are all regarded as conspecific, and the Yurimaguas specimen (Killip 28045) is 

designated as epitype of D. prunifer (because the type is sterile). Unfortunately these specimens still lack 

fruits (described by Martius as “magnitudine juglandum” but apparently lost). Desmoncus prunifer shares all 

character states with Desmoncus polyacanthos, but differs it its much more densely spiny peduncular bract 

and spinulose pinnae bases. A fourth specimen (Galeano 2113) from some distance away in Colombia 

(Amazonas) may also belong here. This specimen has a single fruit, but this is not unusually large and appears 

similar to those of D. polyacanthos. This specimen is not included in the above description but is mapped. 

21. Desmoncus pumilus Trail (1876: 353). 

Atitara pumila (Trail) Kuntze, 1891: 727. Type:—BRAZIL. Amazonas: Rio Padauiri, 26 June 1874, J. Trail 1086/LXXV 

(holotype K!, isotypes BM!, GH!, P!). 

Plants height 4.0(3.0–5.0) m; stems 0.5(0.3–0.7) cm diameter. Leaf petioles 1.7(0.5–8.0) cm long; rachises 


Two specimens (dos Santos 75, Rabelo 87) have pinnae bases with small spines but these appear different 

from the pinnae bases of D. mitis (pinnae bases with uneven surfaces at the bases adaxially, usually covered 

with spinules and/or dense tomentum). Some specimens (Calderon 2709, Carvalho 2149, Cid 10996, dos 

Santos 757) have distal pinnae intermediate between acanthophylls and more usual pinnae, almost like those 

of D. chinantlensis and similar species. 

22. Desmoncus setosus Martius (1823–1837: 89). 

Atitara setosa (Martius) Kuntze (1891: 727). Lectotype (here designated):—BRAZIL. Amazonas: Rio Japurá, near Tefé, 

no date, C. Martius s. n. (M!). Epitype (here designated):—BRAZIL. Amazonas: Mun. Tefé, near mouth of Rio 

Tefé, 19 January 1991, A. Henderson & J. Guedes 1588 (epitype INPA!, isoepitype NY!). 





lateral, straight with briefly swollen bases; pinnae 5(4–6) per side of rachis, with long, filiform apices, without 

a beard of spines at the bases, with uneven surfaces at the bases adaxially, usually covered with spinules and/ 

or dense tomentum; basal pinna 14.2(10.7–19.0) cm long, 3.7(2.4–4.3) cm wide; cirri absent, the rachis 

terminating beyond the distalmost, opposite pair of pinnae in a

or dense tomentum; basal pinna 13.1(7.0–17.7) cm long, 3.1(1.6–4.7) cm wide; cirri well-developed, with 

acanthophylls, with few spines abaxially, mostly on proximal part only, with no intermediate acanthophylls 

present, usually with a wide gap between pinnae and acanthophylls (i.e., gap wider than that between adjacent 

acanthophylls). Inflorescences with the rachis angular, slightly twisted, thicker than the few, closely spaced 

and spirally arranged rachillae, each rachilla not (or very rarely) adnate to the rachis, subtended by an acute 

bracteole and with a well-developed axillary pulvinus; peduncles 2.0(1.0–3.0) mm wide; peduncular bracts 

18.8(16.0–24.5) cm long, broad, ribbed, densely brown tomentose, without spines (very rarely with a few 

spines); rachillae 5(4–9), glabrous or scarcely tomentose initially; basal rachilla 5.1(3.5–6.8) cm long, 

1.5(1.0–2.3) mm wide; stamens 6; fruits 20.9(18.4–23.9) mm long, 13.3(11.1–15.3) mm wide, the surfaces 

uneven with numerous, subepidermal, long, branching fibers; fruiting corollas less than one quarter as long as 

fruits, splitting irregularly into 3 lobes, the lobes often splitting again; endocarps globose to obovoid with 

rounded or slightly peaked apices, the pores lateral. 


Colombia, Peru, and Brazil at 174(112–550) m elevation in lowland rainforest, often on sandy soils (Fig. 12). 

Taxonomic notes:—This distinctive species, with its small leaves and large fruits, was referred to by 

Henderson (1995) as Desmoncus polyacanthos var. prunifer (Poeppig ex Martius) Henderson, based on D. 

prunifer Poeppig ex Martius. This is a mistake, as explained under D. prunifer. 

Undescribed species 

Four specimens from the same locality on eastern Andean slopes in Ecuador are unusual in their straight leaf 

spines, narrow pinnae with long, filiform apices and with an adaxial beard of spines at the bases, and high 

elevation habitat (700–1350 m). They are all sterile but appear to represent an undescribed species. 

ECUADOR. Napo: Hollin-Loreto road to Coca 11 km from take-off from Baeza-Tena road, 1350 m, 

0°42’S 77°43’W, 1 Oct 1995, Balslev et al. 6424 (AAU); Santa Rita west of Archidona, 800–1130 m, 0°55’S 

77°52’W, 12 Aug 1996, Balslev et al. 6447 (AAU); Llanganates, ca. 700 m, 23 Nov 2010, Cornejo et al. 8398 

(NY); km 2, carretera nueva Cotundo-Coca, 1130 m, 5 Aug 1984, Dodson et al. 15042 (MO). 

Acknowledgements 

I thank the many people who have helped with this work. The staff of the herbarium of the New York 

Botanical Garden processed the many 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; Michael Bevans of the Herbarium Digitization Laboratory provided the type images; Dr. Scott 

Mori helped with photography; and Xavier Cornejo helped with information on Ecuadorean specimens. The 

curators of the following herbaria sent these loans, or allowed me to study specimens in their care: A, AAU, 

BH, BM, CEN, COAH, COL, CR, F, GH, HEPH, IBGE, INB, INPA, K, M, MG, MICH, MO, NY, P, PMA, R, 

S, SPF, UB, and US. I thank Drs. Nelson Zamora and Barry Hammel in Costa Rica; Dr Mireya Correa and 

Carmen Galdames in Panama; Paula Leitman in Rio de Janeiro, Drs. José Pirani and Renata Pardini in São 

Paulo, Dr. Aldicir Scariot and Renata Martins in Brasília, and Drs. Michael Hopkins and Alberto Vincentini in 

Manaus. Dr. Rodrigo Bernal wrote the Latin diagnoses. 

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Appendix I. Qualitative Variables—Characters 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/Desmoncus.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. 

1. Rachis spines usually >1 cm long, mostly adaxial or lateral, straight with briefly swollen bases (1); rachis spines 

usually

tomentose margins (rarely without spines) (2); peduncular bracts narrow, elongate, ribbed, scarcely brown tomentose, 

without spines (rarely with few spines) (3); peduncular bracts broad, ridged, densely covered with short, straight, 

swollen-based, vertically oriented spines, these terete, whitish-brown proximally, brown distally, without tomentum (4); 

peduncular bracts broad, ribbed, densely brown tomentose, without spines (very rarely with a few spines) (5); peduncular 

bracts broad, ribbed or ridged, densely covered with felty, reddish-brown tomentum, sparsely covered with short, 

scarcely swollen-based, diagonally oriented, flattened spines, whitish-brown proximally, brown distally, with tomentose 

margins (6); peduncular bracts broad, ribbed with several more prominent, lighter colored ribs, brown, scarcely or not 

tomentose, without spines (rarely with a few spines) (7); peduncular bracts narrow, ribbed, densely whitish-brown 

tomentose, not spiny (8); peduncular bracts broad, the surfaces ribbed or ridged, brown tomentose or glabrous, sparsely 

to moderately covered with short, straight or sinuous, briefly swollen-based, diagonally or vertically oriented spines, 

these flattened or triangular in cross-section, whitish-brown proximally, black or brown distally, with tomentose margins 

(9); peduncular bracts broad, the surfaces deeply ridged, dark brown tomentose, sparsely covered with long, straight or 

sinuous spines, the bases scarcely swollen but running directly into the ridges of the bract and lying flat against the bract 

surface, flattened in cross-section, brown proximally and distally, with tomentose margins (10). (peduncbract) 

10. Rachillae brown tomentose initially (1); rachillae glabrous or scarcely tomentose initially (2). Recorded only from 

inflorescences with flowers because tomentum is early deciduous. (tomentum) 

11. Stamens five to seven (1); stamens eight to 12 (2). (stamenno) 

12. Fruit surfaces uneven with numerous, subepidermal, short, often branching (Y-shaped) fibers (1); fruit surfaces 

smooth, without any apparent subepidermal fibers (2); fruit surfaces uneven with numerous, subepidermal, long, 

branching fibers (3); fruit surfaces bumpy from numerous, subepidermal, short, oblique fibers (4). (fruitsurf) 

13. Fruiting corollas less than one quarter as long as fruits (1); fruiting corollas to half as long as fruits (2). (frcorolla) 

14. Fruiting corollas splitting irregularly into 3 lobes, the lobes often splitting again (1); fruiting corollas not or scarcely 

splitting, tending to remain cupular (2). (splitting) 

15. Endocarps globose to obovoid with rounded or slightly peaked apices, the pores lateral (1); endocarps narrowly 

ellipsoid with rounded apices, the pores lateral (2); endocarps broadly obovoid with flattened apices, the pores lateral on 

or near flattened apices (3); endocarps ovoid to obovoid with prominent, peaked apices, the pores lateral (4). (endocarps) 

Traits 

Abbreviations in parentheses at the end of each variable are the column labels in the Data Matrix. 

1. Stems solitary (1); stems clustered (2). (stems) 



Appendix II. Quantitative variables 

Abbreviations in parentheses at the end of each variable are the column labels in the Data Matrix. 

1. Plant height (m); data taken from specimen labels. (plheight) 

2. Stem diameter (cm); data taken from specimens only, not from labels, measured just above petiole insertion, and 

including sheaths. (stemdiameter) 

3. Petiole length (cm); measured from leaf sheath to first pinna. (petiole) 

4. Rachis length (cm); measured from proximal to distal pinna, excluding cirri. (rachislen) 

5. Rachis width (mm); measured at base of leaf blade. (rachiswid) 

6. Number of pinnae per side of rachis, excluding acanthophylls. (nodivisions) 

7. Basal pinna length (cm). (baspinlen) 

8. Basal pinna width (cm); measured at the widest part. (baspinwid) 

9. Peduncular bract length (cm). (pedbract) 

10. Peduncle width (mm); measured just below the peduncular bract scar. (pedunclewid) 

11. Proximal rachillae length (cm). (rachillen) 

12. Proximal rachilla width (mm); measured at middle of rachilla, between triad scars. (rachillwid) 

13. Number of rachillae. (norachillae) 

14. Number of stamens. (stamen) 

15. Fruit length (mm). (fruitlen) 

16. Fruit diameter (mm). (fruitdiam) 


HENDERSON

Appendix III. Excluded Names 

This list of names comprises published names that have been included in Desmoncus but for which no type specimens 

are available, or the type specimens are insufficient for identification (i.e., the specimen lacks reproductive structures), or 

the type host institution is unknown, or the name has been transferred to another genus, or the specimen is an illustration. 

In particular, no type specimens of Barbosa Rodrigues have survived and all his types are drawings. These have not 

proved possible to identify with any certainty and all Barbosa Rodrigues’ names are treated here as Excluded Names. 

Note that both Glassman (1972) and Wessels Boer (1965) designated lectotypes for many of Barbosa Rodrigues’ (1875, 

1888, 1902) species; however, these should have been neotypes because they were not designated from the original 

material. 

Atitara drudiana Kuntze (1891: 727). Type not designated. 

Atitara paraensis Barbosa Rodrigues (1902: 76). Desmoncus paraensis (Barbosa Rodrigues) Barbosa Rodrigues (1903: 

57). Type not designated. Neotype (designated by Wessels Boer 1965):—Barbosa Rodrigues 1903: t. 57. 

Bactris tenerrima Drude (1881: 328). Desmoncus tenerrimus (Drude) Drude ex Burret (1934: 236). Desmoncus mitis 

var. tenerrimus (Drude) Henderson (1995: 228). Type:—COLOMBIA. Amazonas: Río Caquetá, Araracuara, no 

date, C. Martius s. n. (holotype M!). 

Desmoncus aculeatus Wendland (1854: 20). Atitara aculeata (Wendland) Kuntze (1891: 727). Type not designated. 

Desmoncus aereus Drude (1881: 307). Atitara aerea (Drude) Barbosa Rodrigues (1902: 75). Type:—BRAZIL. 

Amazonas: Rio Negro, Ayrão, 4 July 1874, J. Trail 1088/LXXXVIII (holotype K!). 

Desmoncus americanus Lodd. ex Loudon (1830: 382). Atitara americana (Lodd. ex Loudon) Kuntze (1891: 727). Type 

not designated. 

Desmoncus andicola Pasquale (1867: 36). Type not designated. 

Desmoncus angustisectus Burret (1930: 1025). Type:—BRAZIL. Amazonas: Rio Negro, Trinidade, 12 September 1928, 

P. Luetzelburg 22171 (holotype B, destroyed, isotype M!). 

Desmoncus ataxacanthus Barbosa Rodrigues (1875: 25). Atitara ataxacantha (Barbosa Rodrigues) Kuntze (1891: 727). 

Type not designated. Neotype (designated by Wessels Boer 1965):—Barbosa Rodrigues, 1903: t. 55. 

Desmoncus brevisectus Burret (1934: 215). Type:—BRAZIL. Pará: without locality, 15 November 1932, W. Hopp 35 

(holotype B, destroyed). 

Desmoncus caespitosus Barbosa Rodrigues (1888: 37). Atitara caespitosa (Barbosa Rodrigues) Barbosa Rodrigues 

(1902: 76). Type:—BRAZIL. Rio de Janeiro: Serra do Rodeio, no date, J. Barbosa Rodrigues s. n. (holotype 

destroyed). Neotype (designated by Glassman 1972):—Barbosa Rodrigues 1903: t. 63. 

Desmoncus cuyabaensis Barbosa Rodrigues (1898: 30). Atitara cuyabaensis (Barbosa Rodrigues) Barbosa Rodrigues 

(1902: 75). Type:—BRAZIL. Mato Grosso: Rio Cuiabá, no date, J. Barbosa Rodrigues s. n. (holotype 

destroyed). Neotype (designated by Wessels Boer 1965):—Barbosa Rodrigues 1903: t. 54A. 

Desmoncus dasyacanthus Burret (1934: 213). Type:—VENEZUELA. Carabobo: Puerto Cabello, no date, P. Preuss 

1567 (holotype W, destroyed). 

Desmoncus duidensis Steyermark (1951: 85). Type:—VENEZUELA. Amazonas: Caño Negro, SE of Cerro Duida, 23 

August 1944, J. Steyermark 57944 (holotype F!). 

Desmoncus dubius Lodd. ex Loudon (1830: 382). Atitara dubia (Lodd. ex Loudon) Kuntze (1891: 727). Type not 

designated. 

Desmoncus granatensis Bull (1875: 5). Atitara granatensis (Bull) Kuntze (1891: 727). Type not designated. 

Desmoncus grandifolius Linden (1881: 16). Type not designated. 

Desmoncus huebneri Burret (1934: 200). Type:—BRAZIL. Roraima: Serra do Frechal, Rio Branco, no date, G. Huebner 

80 (holotype B, destroyed). 

Desmoncus inermis Barbosa Rodrigues (1901: 17). Atitara inermis (Barbosa Rodrigues) Barbosa Rodrigues (1902: 76). 

Type:—BRAZIL. Rio de Janeiro: Serra do Mar, no date, J. Barbosa Rodrigues s. n. (holotype not known). 

Neotype (designated by Glassman 1972):—Barbosa Rodrigues 1903: t. 60. 

Desmoncus intermedius Martius ex Wendland in Kerchove de Dentergarten (1878: 243). Type not designated. 

Desmoncus latifrons Bull. Atitara latifrons (Bull) Kuntze (1891: 727). Place of original publication not known. 



Desmoncus leiorhachis Burret (1934: 203). Type:— BELIZE. Toledo: Rio Grande, 29 March 1933, W. Schipp S.517a 


Desmoncus leptochaete Burret (1934: 204). Type:—COSTA RICA. Puntarenas: Río Abrojo, 12 January 1898, H. Pittier 

11969 (holotype B, destroyed). 

Desmoncus leptoclonos Dammer (1907: 129). Desmoncus mitis var. leptoclonos Henderson (1995: 227). Superfluous 

names. 

Desmoncus longifolius Martius (1844: 52). Type:—PERU. Pasco: Pozuzo, 1784, J. Pavón s. n. (holotype MA n.v., 

isotypes F!, G n.v., K!). 

Desmoncus longisectus Burret (1934: 212). Type:—BRAZIL. Alagoas: Serra do Porco, 300 m, March 1932, E. 

Werdermann 2984 (holotype B, destroyed). 

Desmoncus luetzelburgii Burret (1930: 1025). Type:—BRAZIL. Amazonas: Rio Papuri, Rio Uaupés, Trinidade, 13 

December 1928, P. Luetzelburg 23833 (holotype B, destroyed, isotypes, M! R!). 

Desmoncus macroacanthos Martius (1823-1837: 86). Atitara macroacantha (Martius) Kuntze (1891: 727). Type:— 

BRAZIL. Pará: without locality, no date, C. Martius s.n. (holotype M!). 

Desmoncus macrocarpus Barbosa Rodrigues (1888: 34). Atitara macrocarpa (Barbosa Rodrigues) Barbosa Rodrigues 

(1902: 75). Type not designated. Neotype (designated by Wessels Boer 1965):—Barbosa Rodrigues 1903: t. 53. 

Desmoncus macrodon Barbosa Rodrigues (1888: 39). Atitara macrodon (Barbosa Rodrigues) Barbosa Rodrigues (1902: 

75). Type not designated. Neotype (designated by Glassman 1972):—Barbosa Rodrigues 1903: t. 59. 

Desmoncus major Crueg. ex Grisebach (1864: 519). Atitara major (Crueg. ex Grisebach) Kuntze, 1891: 727. Type:— 

TRINIDAD. Caroni, no date, H. Crueger s.n. (holotype not known). 

Desmoncus melanacanthos Drude (1881: 306). Type not designated. 

Desmoncus nemorosus Barbosa Rodrigues (1888: 36). Atitara nemorosa (Barbosa Rodrigues) Barbosa Rodrigues (1902: 

75). Type not designated. Neotype (designated by Glassman 1972):—Barbosa Rodrigues 1903: t. 56. 

Desmoncus oligacanthus Barbosa Rodrigues (1875: 24). Atitara oligacantha (Barbosa Rodrigues) Kuntze (1891: 727). 

Type not designated. Neotype (designated by Glassman 1972) :—Barbosa Rodrigues 1903, t. 58. 

Desmoncus orthacanthos var. trailiana Drude (1881: 311). Type:—BRAZIL. Amazonas: Rio Purus, Jauaria, 10 

September 1874, J. Trail 1079/CXXVI (holotype K!). 

Desmoncus orthacanthos var. mitis Drude (1881: 311). Type:—BRAZIL. “Sertão Amaroleité”, September–October 

1844, H. Weddell 2493 (holotype P!). 

Desmoncus panamensis Linden (1881: 16). Type not designated. 

Desmoncus phengophyllus Drude (1881: 314). Atitara phengophylla (Drude) Kuntze (1891: 727). Type:—BRAZIL. 

Amazonas: Rio Jutahi, 27 January 1875, J. Trail 1075/CCV (holotype K!). 

Desmoncus philippianus Barbosa Rodrigues (1888: 38). Atitara philippiana (Barbosa Rodrigues) Barbosa Rodrigues 

(1902: 76). Type not designated. Neotype (designated by Glassman 1972):—Barbosa Rodrigues 1903: t. 61. 

Desmoncus phoenicocarpus Barbosa Rodrigues (1875: 24). Atitara phoenicocarpa (Barbosa Rodrigues) Kuntze (1891: 

727). Type not designated. Neotype (designated by Glassman 1972):—Barbosa Rodrigues 1875 t. 1, f. 3–4a–d. 

Desmoncus polyacanthos var. cuspidata Drude (1881: 314). Type not designated. 

Desmoncus polyacanthos var. angustifolia Drude. This name appears in Glassman (1972) and the citation of Drude 

(1881: 313) is given. The type specimen is said to be Spruce 627 at M. This specimen still exists, with this name written 

on the label. However, this name is not found on page 313 of Drude (1881), nor on any other page. 

Desmoncus polyphyllus Poit. ex Desfontaines (1829: 30). Atitara polyphylla (Poit. ex Desfontaines) Kuntze (1891: 727). 

Type not designated. 

Desmoncus pycnacanthos Martius (1823-1837: 89). Atitara pycnacantha (Martius) Kuntze (1891: 727). Type:— 

BRAZIL. Goias: Villa de Palma, Rio Pillões, no date, J. Pohl 2488 (holotype W, destroyed). 

Desmoncus pycnacanthos var. sarmentosus Drude (1881: 313). Type:—BRAZIL. Rio de Janeiro, no date, A. Glaziou 

8068 (holotype C n.v., holotype image!, isotype P!). 

Desmoncus rudentum Martius (1844: 48). Atitara rudenta (Martius) Barbosa Rodrigues (1902: 75). Type:—BOLIVIA. 

Santa Cruz: Río Piray, near Palometa, no date, A. d’Orbigny 26 (holotype P!). 

Desmoncus schippii Burret (1934: 202). Type:—BELIZE. Toledo: Rio Grande, 29 March 1933, W. Schipp S.517 



HENDERSON

Desmoncus setosus var. mitescens Drude (1881: 316). Type not designated. 

Desmoncus wallisii Linden (1881:16). Type not designated. 

Desmoncus werdermannii Burret (1933: 114). Type:—BRAZIL. Bahia: Feira Santa Ana, ca. 100 m, April 1932, E. 

Werdermann 3201 (holotype B, destroyed). 

Appendix IV. Plates of Type Images 



Appendix IV, Plates 1–4. Isotype of Desmoncus horridus subsp. occidentalis (H. Balslev, A. Barfod, A. Henderson, F. 

Skov & A. Argüello 60752, NY). Plates 2–3 on next 3 pages. 


HENDERSON

Appendix IV, Plate 2. 





HENDERSON




Appendix IV, Plate 5. Isotype of Desmoncus interjectus (G. Galeano, J. Torres, J. Huitoto & B. Plazas 1092, NY). 


HENDERSON

Appendix IV, Plates 6–12. Isotype of Desmoncus kunarius (M. Nee & E. Tyson 10982, MO). Plates 7–12 on the 

following 6 pages. 





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HENDERSON






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Appendix IV, Plates 13–14. Holotype of Desmoncus loretanus (A. Gentry, C. Diaz, J. Aronson & N. Jaramillo 25785, 

NY). Plate 14 on the next page. 


HENDERSON




Appendix IV, Plate 15. Isotype of Desmoncus madrensis (F. Chávez 626, NY). 


HENDERSON

Appendix IV, Plate 16. Isotype of Desmoncus mitis subsp. ecirratus (G. Prance, E. Forero, L. Coêlho, J. Ramos & L. 

Farias 5636, NY). 



Appendix IV, Plate 17. Isotype of Desmoncus moorei (J. Solano 21, MO). 


HENDERSON

Appendix IV, Plates 18–19. Isotype of Desmoncus obovoideus (A. Gentry 6115, MO). Plate 19 on the next page. 





HENDERSON

Appendix IV, Plates 20–21. Isotype of Desmoncus osensis (R. Aguilar 11417, NY). Plate 21 on the next page. 





HENDERSON

Appendix V. Numerical List of Taxa and Specimens Examined 

Numerical List of Taxa 

1. Desmoncus chinantlensis 

2. Desmoncus cirrhifer 

3. Desmoncus costaricensis 

4. Desmoncus giganteus 

5. Desmoncus horridus 

5.1. Desmoncus horridus subsp. horridus 

5.2. Desmoncus horridus subsp. apureanus 

5.3. Desmoncus horridus subsp. occidentalis 

5.4. Desmoncus horridus subsp. palustris 

5.5. Desmoncus horridus subsp. prostratus 

6. Desmoncus interjectus 

7. Desmoncus kunarius 

8. Desmoncus latisectus 

9. Desmoncus leptoclonos 

10. Desmoncus loretanus 

11. Desmoncus madrensis 

12. Desmoncus mitis 

12.1. Desmoncus mitis subsp. mitis 

12.2. Desmoncus mitis subsp. ecirratus 

12.3. Desmoncus mitis subsp. leptospadix 

12.4. Desmoncus mitis subsp. rurrenabaquensis 

13. Desmoncus moorei 

14. Desmoncus myriacanthos 

15. Desmoncus obovoideus 

16. Desmoncus osensis 

17. Desmoncus orthacanthos 

18. Desmoncus parvulus 

19. Desmoncus polyacanthos 

20. Desmoncus prunifer 

21. Desmoncus pumilus 

22. Desmoncus setosus 

23. Desmoncus stans 

24. Desmoncus vacivus 

Specimens Examined 

Specimens are arranged by collector (with first initial, when known) in alphabetical order, followed by collector’s 

number in increasing order (s. n. = without number), followed by species number in parentheses. (*) = suspected hybrid. 

Acevedo, P. 6667 (19); 8070 ((5.4)); 14585 (19); 14657 (18) 

Agostini, G. 2663 (19) 

Aguilar, R. 286 (23); 290 (16); 2672 (23); 4676 (3); 5655 (13); 5656 (3); 5711 (16); 6606 (16); 11417 (16); 11975 (16) 

Aguilar, G. 703 (1) 

Aguirre-Galviz, L. 1152 (12.1) 

Aizprua, R. 1443 (14) 

Allen, P. 5117 (14); 5787 (16) 



Altamirano, S. 3294 (8) 

Amaral, I. 368 (22); 706 (19) 

Anderson, A. 296 (12 .2); 1175 (19); 2041 (19); 2205 (5.1) 

Anderson, W. 9625 (9); 10577 (18) 

Antonio, T. 1785 (15) 

Arango, J. 4579 (14) 

Araújo, A. 386 (5.1) 

Arbeláez, M. 652 (21) 

Archer, W. 75 (9); 8353 (5.1) 

Arias, J. 1187 (24); 1373 (18); 1552 (12.3) 

Asplund, E. 14047 (24); 14247 (24); 14687 (12.2) 

Aulestia, M. 3553 (4) 

Baca, V. 190 (19) 

Bailey, L. 101 (19); 172 (5.1); 418 (5.1); 619 (5.1) 

Balée, W. 2301 (18) 

Balick, M. 951 (19); 1493 (19) , 1715 (1); 2201 (1); 3275 (1) 

Balslev, H. 4775 (12.1); 6265 (19); 6599 (19); 6620 (19); 6630 (19); 6640 (19); 6649 (12.1); 6650 (12.1); 6656.1 (19); 

6657 (12.1); 6681 (12.1); 6684 (12.1); 6688 (19); 6689 (19); 6690 (19); 6843 (24); 6859 (12.3); 6925 (24); 7122 (10); 

7328 (19); 7342 (19); 7388 (19); 7404 (19); 7408 (19); 7430 (12.1); 7441 (19); 7462 (10); 7463 (19); 7950 (19); 8085 

(19); 8097 (19); 8200 (1); 60752 (5.3); 62042 (4); 62050 (19); 62057 (5.3); 62438 (19) 

Bangham, N. 368 (1) 

Barbour, P. 5067 (19) 

Barfod, A. 60073 (2) 

Bartlett, H. 12576 (1); 12584(1); 16728 (14) 

Bastos, M. 1905 (19); 2238 (19) 

Beck, H. 2272 (2); 10191 (8) 

Berlin, B. 743 (4) 

Bernal, R. 507 (14); 523 (14); 2033 (*); 2083 (5.3); 2538 (12.3); 4408 (5.4) 

Bernardi, A. 6522 (19) 

Betancur, J. 13982 (6) 

Blum, K. 2105 (14) 

Bond, F. 63 (5.1) 

Borchsenius, F. 285 (2) 

Britton, N. 1035 (5.1); 1559 (19) 

Broadway, W. 309 (5.1); 3498 (5.1); 4077 (5.1) 

Bunting, G. 10219 (14) 

Byg, A. 47 (19) 

Byron 1066 (18) 

Cabrera, R. 3054 (12.3) 

Calderón, C. 2709 (21); 2851 (12.2) 

Calonje, M. 7004 (7) 

Campo, J. 450 (5.2) 

Cárdenas, D. 4534 (6); 4642 (22); 6801 (6); 22353 (6); 23658 (6); 24503 (5.4) 

Carvalho, F. 627 (17); 2149 (21); 4522 (17); 6849 (19) 

Cascante, A. 988 (13) 

Castillo, A. 2084 (18) 

Castro, E. 150 (3) 

Cavalcante, P. 1611 (19) 

Chagas, F. 132 (9) 

Chagas, J. 1103 (18); 3853 (21) 


HENDERSON

Chardon, C. s.n (5.2) 

Chávez, F. 626 (11); 683 (11) 

Churchill, H. 5662 (7) 

Cid, C. 37 (19); 2892 (12.2); 2929 (12.2); 3299 (12.3); 4636 (*); 10996 (21) 

Clark, J. 4223 (2) 

Clark, H. 7982 (*) 

Clarke, H. 12189 (19) 

Coêlho, D. 688 (22) 

Coêlho, L. s.n. (19); 28 (*); 28 (18); 1523 (19); 2078 (22); 36032 (19) 

Colque, O. 572 (12.4); 594 (12.4) 

Contreras, E. 868 (1); 976 (1); 2581 (1) 

Cook, O. 11 (1); 42 (14); 53 (1); 97 (1); 142 (2) 

Coradin, L. 7331 (9) 

Cordeiro, I. 199 (19) 

Cornejo, F. 2946 (12.4) 

Costa, J. 680066 (19); 680068 (17); 680069 (17); 680083 (19) 

Couvreur, T. 291 (8) 

Cowan, C. 3378 (1) 

Cremers, G. 11838 (18); 12139 (19); 14340 (19) 

Croat, T. 7759 (14); 9314 (14); 10047 (14); 14561 (14); 16806 (2); 18115 (20); 18439 (24); 18767 (24); 18793 (10); 

19176 (24); 19584 (24); 20039 (24); 21619 (19); 23338 (1); 24746 (1); 40249 (1); 62352 (12.2); 88912 (19) 

Cuadros, H. 1855 (14) 

Cuatrecasas, J. 3977 (5.2) 

Curran, H. 174 (14); 533 (19) 

Dahlgren, B. 437 (19); 523 (19) 

Daly, D. 535 (19); 6494 (12.4); 9387 (*); 10779 (19); 10894 (12.3); 11467 (12.2); 11627 (12.3); 11734 (12.2); 11800 

(12.2); 11986 (*) 

Davidse, G. 12721 (5.2); 13822 (5.2); 14755 (5.2); 15872 (5.2); 17831 (5.1); 17988 (19); 21852 (19); 27672 (19); 27712 

(5.4); 30963 (13) 

De La Cruz, J. 3360 (18) 

Delascio, F. 7964 (5.2) 

Díaz, C. 252 (10) 

Diaz, W. 319 (19) 

Dodson, C. 7190 (2) 

Dorr, L. 7779 (19) 

Dransfield, J. 4861 (2) 

Duke, J. 9806 (14) 

Duque, J. 7391 (12.3) 

Dwyer, J. 10958 (1); 11352 (1) 

da Vinha, S. 76 (19) 

de Bruijn, J. 1235 (19) 

de Castro, A. 573 (19) 

de Granville, J.-J. 2259 (19); 11232 (18); 13678 (18) 

de Nevers, G. 4415 (7); 4716 (2); 4784 (7); 7760 (23) 

de Paula, J. 1213 (19) 

dos Santos, J. 757 (21) 

Eiten, G. 9209 (9); 10208 (19) 

Ek, R. 835 (18) 

Espinelli, F. 406 (21) 

Estupiñán, C. 464 (14) 



Evans, R. 1702 (1); 2742 (19) 

Fendler, A. 732 (19); 2468 (19) 

Fernandes, H. 90 (19); 2090 17 

Fernández, A. 2410 (19); 3942 (19); 11383 (19) 

Ferreira, E. 145 (19); 321 (12.2); 518 (9) 

Fiaschi, P. 244 (17); 256 (17); 723 (17) 

Figueiredo, C. 342 (*); 615 (*) 

Fleck, D. 508 (12.3); 795 (4); 815 (12.1) 

Fleury, M. 1435 (19) 

Folli, D. 673 (17); 1696 (17) 

Fonnegra, R. 272 (2) 

Fonsêca, S. 163 (19) 

Forero, E. 6384 (12.2); 9080 (14) 

Forest Dept. 3579 (19); 5212 (5.1) 

Foster, R. 8814 (12.3) 

França, G. 303 (19) 

Freitas, J. 216 (12.2) 

Fróes, R. 2001 (5.1); 11765 (19); 31198 (19) 

Fuchs, H. 21833 (2) 

Fuentes, A. 4081 (12.4) 

Furtado, P. 76 (9) 

Galeano, G.1092 (6); 2113 (20) 

Garwood, N. 2178 (15) 

Gentle, P. 348 (1); 528 (1); 2453 (1); 4750 (1) 

Gentry, A. 4999 (14); 6115 (15); 8035 (1); 13351 (18); 14819 (19); 16560 (12.3); 19108 (10); 20734 (24) , 22007 (19); 

25466 (19); 25785 (10); 27084 (11); 27743 (12.3); 29230 (12.3); 30363 (2); 31736 (24); 35355 (2); 38752 (19); 39709 

(12.3); 41015 (2); 42260 (12.3); 43657 (11); 43738 (19); 46310 (5.2); 47293 (5.4); 47988 (2); 49428 (17); 49499 (17); 

50223 (5.1); 50347 (19); 53319 (2); 53392 (2); 53795 (2); 54546 (19); 54608 (12.3); 56146 (19); 58515 (12.3); 60885 

(12.3); 61654 (19); 61699 (19); 61978 (12.1); 68598 (11); 74275 (19); 76380 (19); 78307 (11); 78784 (23) 

Gillespie, L. 812 (19); 1050 (18); 1226 (5.1) 

Gines, H. 5248 (5.1) 

Gleason, H. 164 (18); 269 (18) 

Goldenberg, R. 1347 (19) 

González, A. 813 (19) 

Gottsberger, I. 111482 (5.1) 

Graham, J. 4699 (12.3) 

Grández, C. 434 (19); 1069 (24); 1782 (10); 2336 (24) 

Grayum, M. 4463 (3); 8746 (3); 11117 (13) 

Guánchez, F. 1731 (19) 

Gudiño, E. 397 (19) 

Guillén, R. 1517 (5.5); 1559 (5.5); 3169 (5.5); 3841 (8) 

Hage, J. 400 (19); 858 (17) 

Hahn, W. 5134 (18) 

Hammel, B. 1863 (15); 19136 (23); 21197 (13); 21940 (3) 

Harley, R. 11230 (9); 17904 (19); 17969 (17); 18151 (17) 

Harmon, W. 2543 (1) 

Hassler, E. 11293 (9) 

Hatschbach, G. 47732 (17); 57017 (17) 

Hayes, S. 359 (14) 

Hemmendorff, E. 349 (17) 


HENDERSON

Henderson, A. 36 (19); 62 (13); 178 (22); 300 (*); 336 (19); 875 (24); 977 (19); 1077 (18); 1101 (12.2); 1588 (22); 1607 

(19); 1609 (18); 1610 (19); 1617 (17); 1640 (12.4); 1642 (19); 1675 (19); 1688 (4); 1693 (12.2); 1696 (5.3); 1811 (23); 

3009 (19) 

Heringer, E. 15163 (19) 

Hernández, H. 373 (1); 629 (1) 

Herrera, G. 1107 (13); 4596 (23) 

Hoffman, B. 799 (19); 6380 (19) 

Hoffman, D. 141 (5.4) 

Holdridge, L. 5126 (13) 

Hollowell, T. 412 (5.1) 

Holm, R. 931 (13) 

Howard, R. 10405 (5.1) 

Huashikat, V. 979 (4); 1098 (4); 1099 (4); 2309 (12.3) 

Idrobo, J. 506 (19) , 4707 (12.3) 

Irwin, H. 7194 (9) 

Jangoux, J. 85007 (12.2) 

Jansen-Jacobs, M. 2539 (19) 

Jaramillo, J. 4344 (12.1); 4607 (12.1) 

Jaramillo, N 1290 (19) 

Jardim, J. 89 (17); 1538 (19) 

Jativa, C. 644 (2) 

Jenman, G. 7726 (5.1) 

Juncosa, A. 1570 (2) 

Kahn, F. 504 (19); 540 (*); 563 (19); 3503 (19); 3506 (19) 

Kelloff, C. 631 (19); 917 (18) 

Kennedy, H. 2800 (15) 

Killip, E. 27071 (12.3); 28045 (20); 28807 (12.3); 30573 (19) 

Kinupp, V. 1410 (22); 2334 (9) 

Knab-Vispo, C. 368 (19) 

Kohn, E. 1163 (4) 

Kramer, K. 2472 (19); 2708 (5.1); 3146 (19) 

Krapovickas, A. 23228 (17); 31940 (*) 

Krieger, L. 18905 (19) 

Krukoff, B. 4996 (12.2) 

Kuhlmann, J. 522 (8) 

Kvist, L. 1156 (*); 1196 (19); 1545 (19) 

La Rotta, C. 137 (24) 

Lanna, J. 973 (17) 

Leal, E. 140 (5.1) 

Lemos, R. 6321 (19) 

Lewis, M. 813 (17); 1059 (19); 37933 (19) 

Liebmann, F. 6596 (1); 6956 (1) 

Liesner, R. 5694 (19); 6274 (*); 8940 (*) , 9216 (19); 13632 (19); 25728 (18) 

Lima, J. 796 (19) 

Lindman, C. 2827 (5.5); 3273 (9) 

Listabarth, C. 1110589 (*) 

Little, D. 16639 (5.1) 

Lleras, E. 17184 (10) 

Lobato, L. 34 (5.1) 

Loomis, H. 54 (19); 63 (5.1) 



López, R. 10994 (19) 

Loureiro, A. s. n. (*) 

Luetzelburg, P. 20026 (17) 

Lundell, C. 1555 (1); 2646 (1); 3421 (1); 4750 (1); 16237 (1) 

Maas, P. 3559 (18); 12879 (12.2); 13130 (12.2) 

Macbride, J. 5470 (19) 

Macedo, A. 705 (9) 

Maciel, U. 167 (18) 

Madison, P. 31 (19); 361 (21) 

Madriñán, S. 931 (5.4) 

Maguire, B. 23093 (18); 25011 (19) 

Malagón, W. 85 (4) 

Malave, I. 23 (5.1) 

Manriquez, G. 2535 (1); 3482 (1) 

Marín, J. 194 (23); 1983 (24) 

Martínez, E. 6344 (1); 27903 (1) 

Martius, C. s. n. (12.1) 

Mata 4 (19); 18 (19) 

Matos, F. 244 (19) 

Matuda, E. 3196 (1) 

Mauro, J. 50 (19) 

Maxon, W. 6573 (14) 

Mayfield, M. s. n.. (23) 

McDaniel, S. 14208 (19); 21624 (24); 32182 (10) 

McDowell, T. 3109 (18); 3730 (19); 4104 (18) 

McPherson, G. 6961 (15) 

Mendes, V. 251 (9) 

Mendoza, A. 2733 (1) 

Miraña, J. 8 (24) 

Miranda, F. 470 (*) 

Molina, A. 1808 (13) 

Monteiro, O. 1317 (19) 

Moore, H. 6361 (1); 8273 (1); 8432 (24); 8449 (24); 8591 (12.4); 9471 (2); 10121 (13); 10349 (5.1); 10360 (19); 10377 

(5.1); 10380 (5.1); 10510 (14) 

Mora, E. 580 (3); 626 (23) 

Moraes, M. 845 (12.4) 

Moraga, C. 975 (13) 

Morales, J. 6333 (16); 9284 (13) 

Moreno, L. 162 (8); 163 (12.2); 175 (12.4); 185 (12.2); 192 (8) 

Moreno, P. 12428 (13); 24093 (1) 

Mori, S. 5184 (15); 7737 (2); 8063 (18); 8100 (18); 8455 (18); 9128 (19); 15604 (18); 17735 (19); 22378 (22) 

Morillo, G. 8873 (19) 

Mowbray, T. 1441 (13) 

Murillo, J. 1940 (19) 

Murray, N.1550 (24) 

Mutchnick, P. 829 (19) 

Nascimento, J. 723 (19) 

Nee, M. 10982 (7); 14003 (14); 17811 (19); 34905 (12.2); 34990 (12.2); 39611 (19); 42329 (18) 

Noblick, L. 1796 (17); 4695 (17); 4721 (17); 4724 (19); 4771 (19); 4791 (17); 4804 (19) 

Nogueira, P. 688 (5.1) 


HENDERSON

Núñez, P. 10015 (11); 10483 (11); 19077 (12.3) 

Oldeman, R. 1269 (19) 

Oliveira, J. 732 (19) 

Ortega, F. 3290 (5.2) 

Pabon, M. 329 (5.4) 

Palacios, W. 932 (19) 

Pardini, R. 39 (19); 72 (19) 

Peixoto, A. 3010 (17); 3402 (17); 3491 (17) 

Pereira-Silva, G. 4648 (9); 4766 (9); 9567 (19) 

Persaud, A. 77 (18) 

Pessôa, C. 254 (12.2); 319 (12.2) 

Pinto, E. 1278 (5.2) 

Pipoly, J. 7442 (18); 7451 (19); 9236 (18); 9956 (18); 15001 (24); 16068 (12.3) 

Pirani, J. 2330 (19); 2418 (19); 2485 (17); 2961 (17); 3062 (17) 

Pires, O. 222 (22) 

Pires, M. 484 (19); 2602 (19); 47538 (19) 

Pittier, H. 416 (2); 2606 (14); 7078 (19); 7795 (19) 

Plowman, T. 3720 (14); 9547 (18); 9577 (*); 9742 (*); 9881 (*); 12572 (19) 

Poncy, O. 249 (19) 

Poole, J. 2005 (5.4) 

Posada, A. 2244 (12.3) 

Prance, G. 5636 (12.2); 10067 (19); 12310 (12.3); 12490 (12.3); 13946 (12.2); 23993 (24); 29511 (19); 29626 (19) 

Prévost, M. 1852 (5.1); 2729 (19) 

Quesada, F. 801 (23) 

Quinet, A. 1343 (12.2) 

Rabelo, A. 79 (12.3); 87 (21); 88 (19) 

Rabelo, B. 2533 (18) 

Ratter, J. 5735 (19) 

Read, R. 3564 (19); 3570 (17) 

Redden, K. 5943 (19) 

Renteria, E. 3554 (14); 4812 (14) 

Revilla, J. 341 (10); 957 (24); 1078 (19); 1166 (10); 1568 (24) 

Rimachi, M. 10809 (10); 11783 (12.3) 

Robles, R. 1381 (13) 

Rocha, A. 258 (18); 688 (19); 725 (19); 991 (5.1) 

Rodrigues, W. 9790 (21) 

Rodriguez, A. 364 (13) 

Rosa, N. 665 (19) 

Rosero, L. 2007 (18) 

Rubio, D. 1422 (2) 

Rudas, A. 2297 (19); 2868 (12.3); 3103 (12.3) 

Rueda, R. 1726 (13); 3544 (13); 4097 (13); 4718 (13) 

Rusby, H. 98 (19) 

Sacco, J. 43 (11) 

Saldias, M. 2624 (5.5) 

Santos, N. 5162 (17) 

Sasaki, D. 14 (12.2) 

Saunders, J. 553 (1) 

Scariot, A. 617 (19); 621 (*) 

Schinini, A. 29368 (9) 



Schultes, R. 3941 (24); 15158 (6); 16233 (6) 

Schunke, J. 2644 (11); 6927 (19); 14877 (19); 15033 (12.2); 15269 (12.3); 15726 (12.3) 

Sevilha, A. 2410 (19) 

Shattuck, O. 787 (14) 

Sidney [= Fonsêca, S.] 249 (9) 

Siebert, S. 2 (1) 

Silva, S. 390 (9) 

Silva, M. 1666 (5.4); 2463 (5.1) 

Silveira, M. 417 (19); 1636 (12.2) 

Simpson, D. 698 (*) 

Sinaca, S. 1230 (1) 

Smith, C. 6041 (5.2) 

Smith, D. 3791 (19); 13959 (19) 

Smith, H. 2339 (14) 

Smith, L. 6390 (17) 

Smith, S. 1392 (19); 2127 (19) 

Soejarto, D. 780 (12.3) 

Solano, J. 21 (13) 

Solomon, J. 3444 (24); 7814 (12.2); 17098 (12.2) 

Sothers, C. 337 (19) 

Sousa, M. 12040 (1) 

Souza 266 (19) 

Souza, S. 1032 (19) 

Spellman, L. 1882 (1) 

Spruce, R. 46 (*) 

Stahel, G. s. n. (5.1) 

Stergios, B. 6040 (5.2) 

Stevens, W. 23935 (13) 

Steyermark, J. 38646 (1); 44526 (1); 45990 (1); 49232 (1); 60968 (5.2); 87273 (19); 95458 (19); 101287 (19); 102000 

(5.2); 114309 (5.1); 114693 (5.1); 116785 (19); 122127 (19); 122179 (19); 122423 (18); 123408 (14) 

Strudwick, J. 5013 (19) 

Sutton, D. 68 (1) 

Taylor, D. 24 (1) 

Tessmann, G. 5236 (12.3) 

Thomas, W. 6026 (17); 6777 (19); 12143 (19); 14265 (17) 

Timaná, M. 1272 (11); 2629 (11) 

Tipaz, G. 1274 (2); 1483 (2) 

Trail, J. 1083 (12.3); 1086 (21); 1087 (5.4) 

Trujillo, B. 1706 (24); 17454 (5.1) 

Tunqui, S. 340 (12.1) 

Ule, E. 5388 (19) 

Urego, L. 1064 (12.1) 

Valenzuela, L. 2254 (11) 

Vargas, E. 2102 (19) 

Vargas, G. 1564 (23) 

Vargas, L. 867 (12.2); 1046 (8) 

Vásquez 1143 (19); 2300 (*); 2646 (24); 2956 (24); 3119 (24); 3381 (19); 5153 (12.3); 5662 (24); 5695 (100, 5905 

(12.3); 6544 (24); 8329 (12.3); 8352 (12.3); 9170 (24); 10686 (19); 11246 (19); 12139 (12.3); 3529 (24); 13559 (19); 

13823 (24); 14108 (12.3); 16148 (19); 16319 (24); 18203 (24); 18335 (12.3); 18978 (19); 24109 (19) 

van der Wal, M. 284 (12.1) 


HENDERSON

Velez, I. 2308 (5.2) 

Vester, H. 71 (12.1) 

Vicentini, A. 620 (19) 

Vieira, M. 1233 (19) 

Vormisto, J. 530 (24); 612 (19); 622 (12.3) 

Webber, A. 1155 (19) 

Wendt, T. 5776 (1) 

Werdermann, E. 2508 (8) 

Wessels Boer, J. 170 (5.1); 174 (19); 193 (19); 197 (5.1); 526 (5.1); 575 (19); 619 (19); 970 (19); 1037 (19); 1322 (19); 

1335 (5.1); 1625 (19); 1632 (5.1); 1639 (5.1); 1647 (19); 1648 (5.1); 1649 (19); 1658 (5.1); 2082 (19); 2377 (21) 

Wilbert, W. 178 (5.1); 68327 (5.1) 

Williams, R. 691 (14); 785 (20) 

Williams, L. 3451 (10); 3907 (12.3); 6661 (19); 9041 (1); 12724 (5.2) 

Wurdack, J. 43486 (5.4) 

Young, K. 97 (11) 

Yuncker, T. 4612 (1); 8809 (1) 

Zarucchi, J. 1715 (4); 2007 (18) 

Zuluaga, S. 20 (14) 



Appendix VI. Index of Names 

Accepted taxa are bold face. 

Atitara . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .16 

Atitara aculeata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Atitara aerea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Atitara americana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Atitara ataxacantha . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Atitara caespitosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Atitara chinantlensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .19 

Atitara costaricensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .21 

Atitara cuyabaensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Atitara drudiana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Atitara dubia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Atitara granatensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Atitara horrida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .23 

Atitara inermis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Atitara latifrons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Atitara leptoclona . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .29 

Atitara leptospadix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .29 

Atitara lophacantha . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .38 

Atitara macroacantha . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Atitara macrocarpa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Atitara macrodon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Atitara major . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Atitara mitis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .31 

Atitara nemorosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Atitara oligacantha . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Atitara orthacantha . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .38 

Atitara oxyacantha . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .40 

Atitara palustris . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .25 

Atitara paraensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Atitara phengophylla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Atitara philippiana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Atitara phoenicocarpa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Atitara polyacantha . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Atitara polyphylla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Atitara prostrata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .26 

Atitara prunifera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .43 

Atitara pumila . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .44 

Atitara pycnacantha . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Atitara riparia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .26 

Atitara rudenta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Atitara setosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .45 

Bactris tenerrima . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Desmoncus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .16 

Desmoncus aculeatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Desmoncus aereus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Desmoncus americanus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Desmoncus andicola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Desmoncus angustisectus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Desmoncus anomalus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .19 

Desmoncus apureanus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .24 


HENDERSON

Desmoncus ataxacanthus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Desmoncus brevisectus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Desmoncus brittonii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .24 

Desmoncus caespitosus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Desmoncus campylacanthus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .40 

Desmoncus chinantlensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .19 

Desmoncus cirrhifer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .21 

Desmoncus costaricensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .21 

Desmoncus cuyabaensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Desmoncus dasyacanthus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Desmoncus demeraranus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .24 

Desmoncus dubius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Desmoncus duidensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Desmoncus ferox . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .19 

Desmoncus giganteus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .22 

Desmoncus granatensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Desmoncus grandifolius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Desmoncus hartii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .24 

Desmoncus horridus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .23 

Desmoncus horridus subsp. apureanus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .24 

Desmoncus horridus subsp. horridus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .24 

Desmoncus horridus subsp. occidentalis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .24 

Desmoncus horridus subsp. palustris . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .25 

Desmoncus horridus subsp. prostratus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .26 

Desmoncus huebneri . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Desmoncus inermis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Desmoncus interjectus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .26 

Desmoncus intermedius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Desmoncus isthmius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .36 

Desmoncus kaieteurensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .39 

Desmoncus kuhlmannii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .28 

Desmoncus kunarius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .27 

Desmoncus latifrons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Desmoncus latisectus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .28 

Desmoncus leiorachis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Desmoncus leptochaete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Desmoncus leptoclonos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .29 

Desmoncus leptoclonos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Desmoncus leptospadix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .33 

Desmoncus longifolius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Desmoncus longisectus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Desmoncus lophacanthos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .38 

Desmoncus loretanus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .29 

Desmoncus luetzelburgii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Desmoncus lundellii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .19 

Desmoncus macroacanthos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Desmoncus macrocarpus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Desmoncus macrodon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Desmoncus madrensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .31 

Desmoncus maguirei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .40 

Desmoncus major . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Desmoncus melanacanthos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Desmoncus mirandanus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .40 

Desmoncus mitis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .31 



Desmoncus mitis subsp. ecirratus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .33 

Desmoncus mitis subsp. leptospadix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .33 

Desmoncus mitis subsp. mitis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .32 

Desmoncus mitis subsp. rurrenabaquensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .34 

Desmoncus mitis var. leptospadix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .33 

Desmoncus mitis var. rurrenabaquensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .34 

Desmoncus mitis var. tenerrimus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Desmoncus moorei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .35 

Desmoncus multijugus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .24 

Desmoncus myriacanthos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .36 

Desmoncus nemorosus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Desmoncus obovoideus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .36 

Desmoncus oligacanthus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Desmoncus orthacanthos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .38 

Desmoncus orthacanthos var. mitis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Desmoncus orthacanthos var. trailiana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Desmoncus osensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .39 

Desmoncus oxyacanthos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .40 

Desmoncus palustris . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .25 

Desmoncus panamensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Desmoncus paraensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Desmoncus parvulus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .39 

Desmoncus peraltus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .40 

Desmoncus phengophyllus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Desmoncus philippianus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Desmoncus phoenicocarpus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Desmoncus polyacanthos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .40 

Desmoncus polyacanthos var. angustifolia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Desmoncus polyacanthos var. cuspidata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Desmoncus polyacanthos var. oxyacanthos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .40 

Desmoncus polyacanthos var. prunifer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .43 

Desmoncus polyphyllus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Desmoncus prestoei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .40 

Desmoncus prostratus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .26 

Desmoncus prunifer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .43 

Desmoncus pumilus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .44 

Desmoncus pycnacanthos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Desmoncus pycnacanthos var. sarmentosus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Desmoncus quasillarius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .19 

Desmoncus riparius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .26 

Desmoncus rudentum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54 

Desmoncus schippii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .55 

Desmoncus setosus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .45 

Desmoncus setosus var. mitescens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .55 

Desmoncus stans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .45 

Desmoncus tenerrimus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53 

Desmoncus tobagonis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .24 

Desmoncus uaxactunensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .19 

Desmoncus ulei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .40 

Desmoncus vacivus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .46 

Desmoncus velezii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .24 

Desmoncus wallisii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .55 

Desmoncus werdermannii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .55 


HENDERSON

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