Attalea Revision 2021

Phytotaxa 444 (1): 001–076 ISSN 1179-3155 (print edition)
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Monograph PHYTOTAXA
Copyright © 2020 Magnolia Press ISSN 1179-3163 (online edition)
https://doi.org/10.11646/phytotaxa.444.1.1
PHYTOTAXA
444
A revision of Attalea
(Arecaceae, Arecoideae, Cocoseae, Attaleinae)
ANDREW HENDERSON
The New York Botanical Garden, Bronx, NY 10458-5126, U.S.A.
ahenderson@nybg.org; https://orcid.org/0000-0001-6334-9435
Magnolia Press
Auckland, New Zealand
Accepted by Carlos Lehnebach: 23 Mar. 2020; published: 25 May 2020 1

ANDREW HENDERSON
A revision of Attalea (Arecaceae, Arecoideae, Cocoseae, Attaleinae)
(Phytotaxa 444)
76 pp.; 30 cm.
25 May 2020
ISBN 978-1-77670-933-5 (paperback)
ISBN 978-1-77670-934-2 (Online edition)
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ISSN 1179-3155 (print edition)

ISSN 1179-3163 (online edition)
2 • Phytotaxa 444 (1) © 2020 Magnolia Press HENDERSON

Table of contents
Abstract ...........................................................................................................................................................................................................3
Introduction .....................................................................................................................................................................................................3
Materials and Methods ....................................................................................................................................................................................6
Results ............................................................................................................................................................................................................8
Taxonomic Treatment ...................................................................................................................................................................................13
References .....................................................................................................................................................................................................63
Appendix I. Quantitative Variables ...............................................................................................................................................................68
Appendix II. Qualitative Variables................................................................................................................................................................68
Appendix III. Excluded and Doubtful Names ..............................................................................................................................................70
Appendix IV. Index of Names.......................................................................................................................................................................72
Acknowledgements .......................................................................................................................................................................................77
Abstract
A taxonomic revision of the neotropical palm genus Attalea based on morphological data was carried out. Nine hundred
and two herbarium specimens were scored for 21 quantitative and 33 qualitative variables. Using the Phylogenetic Species
Concept, qualitative variables were used to recognize 30 species. These are widely distributed in Central and South America
from Mexico to Bolivia and Paraguay, with one species in Haiti. Nomenclature, descriptions, and distribution maps are pro-
vided for each species. Qualitative variables are illustrated, and illustrations of 22 living plants are also provided.
Key words: Maximiliana, Neotropics, Orbignya, Palmae, Scheelea
Introduction
Attalea is placed in the subfamily Arecoideae, tribe Cocoseae, subtribe Attaleinae (Dransfield et al. 2008). The Attaleinae
contains 11 genera. Seven of these are neotropical (Allagoptera, Attalea, Butia, Jubaea, Parajubaea, Polyandrococos,
Syagrus), two are from Madagascar (Beccariophoenix, Voanioala), one is from southern Africa (Jubaeopsis), and the
pantropical Cocos is possibly of neotropical origin (Meerow et al. 2009). Attalea is common and widespread in the
neotropics in Central and South America, with one species occurring in the Caribbean.
Perhaps no other genus of palms has had such a long and complex taxonomic history as Attalea. In its almost
two hundred year history, from the first description to the latest monograph (Glassman 1999), no fewer than 16
genera (including two of hybrid origin) and 167 species have been proposed. One species, here called Attalea maripa
(Aublet) Martius, has been placed in six different genera. Most recent estimates of the number of taxa range from five
genera and 65 species (Glassman 1999) to one genus and 27 species (Henderson et al. 1995). The whole, complicated
taxonomic history has been told by Glassman (1999) and will not be repeated here.
Glassman’s (1999) treatment is the most recent, and is the starting point for the present study. Glassman recognized
five genera (Attalea, Maximiliana, Orbignya, Scheelea, Ynesa) and 65 species. He examined only about 450 specimens
(excluding images of specimens and cultivated plants) compared with 902 in this study. He relied on staminate flowers
to distinguish the four genera, but, as discussed below, these genera are not monophyletic. He also relied on quantitative
data to distinguish species.
Recently there has been renewed interest in the taxonomy and phylogeny of Attalea. Pintaud (2008) and Rodríguez
del Castillo et al. (2016) have given discussions of the problems of nomenclature and morphology, and Pintaud et al.
(2016) have discussed Attalea in the western Amazon region. There have been several studies on phylogeny (Meerow
et al. 2009; Meerow et al. 2015; Rodríguez del Castillo et al. 2016; Freitas et al. 2016), as discussed below. These
phylogenetic studies have now firmly established that only one genus, Attalea, should be recognized.
Problems in Attalea
Probably no other neotropical genus is as difficult taxonomically as Attalea. There are several reasons for this. The
plants themselves are often extremely large, with some species having stems up to 30 m tall and leaves up to 10 m
long. This means they are difficult to collect and consequently specimens in herbaria tend to be few and fragmentary.
A REVISION OF ATTALEA Phytotaxa 444 (1) © 2020 Magnolia Press • 3

Inflorescences and infructescences are large with woody bracts and are seldom collected and preserved in their entirety.
Inflorescences are also often unisexual, so that either staminate or pistillate parts are collected, but not both. Much
previous taxonomic work in the group has been based on staminate flowers, and without these, which are seldom
present, it was difficult to identify specimens.
Type specimens are often fragmentary, destroyed, or otherwise equivocal, and a consequence of this is numerous
doubtful names. The type localities of many of the earlier names are often vague and difficult to establish. The
Brazilian botanist João Barbosa Rodrigues described approximately 20 taxa, but the types are no longer extant and
only represented by drawings. The German botanist Max Burret described approximately 30 species but many of his
types were destroyed in Berlin.
The natural habitats of most species have been greatly modified by man, and this makes collecting problematic,
especially recollecting species with fragmentary type specimens. Conversely, some species appear to have spread into
disturbed habitats (e.g., Pires-O’Brien 1993) and sometimes this brings together species that were previously separate
geographically.
The group has suffered from a parochial approach in which botanists from one country ignore species in adjacent
countries.
There are several widespread and complex species. These do not appear to be based on any cytological or breeding
system abnormality (e.g., apomixis). Although only seven species have been counted, all have a chromosome number
of 2n = 32, and there is no evidence of polyploidy (Dransfield et al. 2008). Plants of Attalea are monoecious and,
where known, they are outcrossing and pollinated by beetles (Henderson 2002). There is limited evidence for any
kind of clinal variation (but see discussion under A. funifera). Two of these widespread and complex species, Attalea
butyracea and A. phalerata, have given the most taxonomic problems in the genus, and have been split into numerous,
little different species, often based on quantitative variables. There still seems a tendency to recognize such species
(e.g., Pintaud 2008; Rodríguez del Castillo et al. 2016; Pintaud et al. 2016). However, many populations of widespread
and complex species are different quantitatively. Without an explicit species concept, and using quantitative variables,
any number of ‘species’ in these complexes can be recognized.
As noted above, previously at least five genera (Attalea, Maximiliana, Orbignya, Scheelea, Ynesa) were commonly
recognized. Now that only one monophyletic genus is recognized, Attalea nomenclature is bedeviled by confusion over
the many published species names. Often the same specific epithet was used for species in the five different genera.
However, by far the greatest impediment to understanding Attalea appears to be hybridization.
Hybridization in Attalea
In at least half of the 30 species recognized here there is evidence of hybridization. However, while hybrids can be
hypothesized based on morphological data, they cannot be known with any certainty, and detecting hybrids from
herbarium specimens is problematic. Only a few field studies of hybrids have been carried out (e.g., Balick et al. 1987a,
1987b; Anderson & Balick 1988). Nevertheless, there seems overwhelming evidence of extensive hybridization in the
genus, and clearly hybridization has been important in the evolution of Attalea. In the present study, hybrids make
scoring of some variables problematic, as discussed in the Materials and Methods section.
Hybrids occur in scattered localities (Fig. 1). While several of these hybrids appear to be isolated plants (e.g.,
Attalea × voeksii, A. salvadorensis) known from restricted areas, others form hybrid zones. For example, Markleya
dahlgreniana occurs over a wide area from Surinam to Pará and Maranhão in Brazil. Orbignya × teixeriana occurs
across the southern margins of the Amazon region from Maranhão in Brazil to Santa Cruz in Bolivia. The presence of
numerous hybrids in the northeastern part of Brazil, particularly Maranhão (Fig. 1) may reflect a long history of human
disturbance in these regions.
As noted above, several species are known to be invasive. Little is known about the association between
hybridization and invasiveness but there is likely to be a correlation (Ellstrand & Schierenbeck 2000).
Pinheiro (1997) gave an extensive discussion on hybrids in Attalea. Using anatomical sections of leaves from
75 herbarium specimens, representing 30 species, he scored 40 quantitative and 47 qualitative characters. Pinheiro
used only 1–3 specimens per species, so he had a small sample size. He considered that as many as 16 species were
either putative hybrids or ‘intermediates’, species with confusing patterns of character variation. Based on multivariate
analysis of his data, Pinheiro concluded the following species were of probable hybrid origin (names follow Pinheiro’s
nomenclature i.e., Attalea s.s., Orbignya, Scheelea): A. nucifera, A. racemosa, A. tessmannii, × A. minarum, × A.
piassabossu, O. sagotii, O. polysticha, O. spectabilis, S. butyracea, and S. phalerata. These are discussed under the
relevant species below.
FIGURE 1. Distribution of all Attalea specimens examined (hybrids, white dots).
Phylogeny
There have been several previous attempts to understand species relationships in Attalea. Pinheiro (1997) studied
generic boundaries within Attalea using phylogenetic methods and anatomical data from leaf sections from 75 herbarium
specimens, representing 30 species. He was able to score 47 qualitative characters. Using parsimony analysis and
qualitative characters, with three species of Syagrus as outgroup, the following relationships were resolved (Pinheiro’s
Fig. 32; names follow his nomenclature i.e., Attalea s.s., Maximiliana, Orbignya, Scheelea). Maximiliana maripa
was resolved as sister species to all others; three species (S. insignis, S. attaleoides, A. nucifera—the Scheelea group)
were sister to the remaining species. Four species (O. cuatrecasana, O. cohune, O. guacuyule, O. luetzelburgii) were
unresolved. The remaining species formed two clades; one with four species (O. eichleri, O. phalerata, O. oleifera,
O. brejinhoensis—the Brazilian Orbignya group) and the other with nine species (A. oleifera, A. dubia, A. funifera,
A. humilis, A. geraensis, A. pindobassu, A. × piassabossu, as well as O. crassispatha and A. colenda—the Attalea
group).
The second study is that of Meerow et al. (2009; see also Meerow et al. 2015). These authors used molecular
data to study the position of Cocos in relation to other genera of the Attaleinae, and included 16 species of Attalea.
They found high support for a monophyletic Attalea. Within Attalea, they found a highly supported Attalea clade (A.
brasiliensis, A. humilis, A. oleifera, A. pindobassu, A. funifera, and A. seabrensis), and the Caribbean A. crassispatha
as sister to two remaining clades; A. cohune and A. guacuyule as sister species and sister to a Scheelea clade (A.
anisitsiana, A. phalerata, A. butyracea) and an Orbignya clade (A. brejinhoensis, A. eichleri, A. speciosa). Rodríguez
del Castillo et al. (2016) obtained similar results using different markers.
The most inclusive study of phylogeny using similar molecular markers as Meerow et al. (2009) is that of Freitas
et al. (2016), who included 37 species. They recovered three main clades. The first clade was termed the Scheelea-like
clade (comprising A. allenii, A. amygdalina A. anisitsiana, A. butyracea, A. cohune, A. colenda, A. cuatrecasana, A.
ferruginea, A. guacuyule, A. insignis, A. nucifera, A. phalerata, A. plowmanii, A. tessmannii – names as used by Freitas
et al.). It included species previously included in Scheelea but also several species previously included in Orbignya

(A. cohune, A. cuatrecasana, A. guacuyule) and some in Attalea s.s. (A. allenii, A. ferruginea). The second clade was
termed the Orbignya-like clade (comprising A. allenii, A. attaleoides A. barreirensis, A. brejinhoensis, A. crassispatha,
A. eichleri, A. luetzelburgii, A. maripa, A. microcarpa, A. moorei, A. polysticha, A. speciosa, A. spectabilis, A. vitrivir).
This clade contained species previously included in Orbignya, as well as A. crassispatha, A. luetzelburgii, and A.
maripa. Attalea moorei was included here, although this species was previously included in Scheelea. The third, termed
the Attalea-like clade (comprising A. brasiliensis, A. burretiana, A. funifera, A. geraensis, A. humilis, A. oleifera, A.
pindobassu, A. seabrensis) was strongly supported.
In summary, there seem to be several areas needing more research in Attalea. A more robust phylogeny is needed,
based on the most recent methodology (e.g., Barrett et al. 2016) and including all species. Species complexes should be
studied throughout their range, using, for example, the field methods of Rodríguez del Castillo et al. (2016) for Attalea
phalerata, and an explicit species/subspecies concept should be applied to the results. Such sampling will reveal either
quantitatively different populations or, more likely, continuous variation. The extent to which hybrids will complicate
this research is unknown, but clearly more studies are needed of hybrids, and hybrid zones, in Attalea.
Materials and Methods
Species concept
This study is based on the Phylogenetic Species Concept (PSC). 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). The terms character and trait are used in the sense of Nixon & Wheeler (1990). Characters
are qualitative variables of which only one state is found in all comparable individuals within a species. In contrast,
traits are qualitative variables of which more than one state occurs within a species (i.e., they are polymorphic within
a given species).
According to Davis and Nixon (1992), phylogenetic species are delimited by successive rounds of aggregation
of local populations (population aggregation analysis—PAA), based on analysis of characters and traits. However,
herbarium 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, this kind of analysis is
problematic. Because of this, Snow et al. (2003) proposed a slightly different method to delimit phylogenetic species,
that they called specimen aggregation analysis (SAA). However, with both PAA and SAA, populations or individual
specimens still need to be identified to species before the analysis can begin. Other methods, such as those used by
Henderson (2011), are based on circular reasoning—species are defined by characters, but characters are defined by
species. These methods also lead to the elimination of traits for the whole genus, but clearly some qualitative variables
could function as traits in some species or groups of species, and characters in others. In this study all specimens were
determined a priori to what are termed preliminary species, and the method proceeds from there in several stages, as
explained below.
Glassman (1999) recognized 65 species in five genera (Attalea, Maximiliana, Orbignya, Scheelea, Ynesa) and these
are the preliminary species of this study. They function as hypotheses that are tested as having a unique combination
of qualitative variable states. If the unique combination is upheld, then they are recognized as phylogenetic species. If
not, then they are further tested. New hypotheses of species (and characters) are proposed as a result.
The first stage was to determine all specimens examined to preliminary species according to either determinations
on the specimens themselves or using Glassman’s (1999) key. Sterile specimens were usually excluded from this
study because of the difficulty of identifying them. Types based on illustrations were not included in the data matrix.
Approximately 30% of names with types are illustrations, and these are placed based on their protologues.
The next stage was to score all specimens for as many qualitative variables as possible. These variables (Appendix
II) were taken either from existing treatments (e.g., Henderson 1995, Glassman 1999), or from direct observation of
the specimens. Thirty-three qualitative binary or multistate variables were scored. At the same time, location (latitude,
longitude, elevation) and quantitative variables (Appendix I) were also scored. Latitude, longitude, and elevation
were taken from the specimen label, when given. Since most specimens lack coordinates, these were estimated from
the collection locality using electronic gazetteers, Google Earth, or other sources. However, there are many problems
with the retrospective assignment of coordinates; label locality data, especially on older specimens, is often vague
or inaccurate and place names may be misspelled and difficult to find in gazetteers. Different gazetteers sometimes

provide different coordinates for what is apparently the same place name. The distribution maps, produced by Arcview
GIS 3.2 (Environmental Systems Research Institute, Inc.), where each dot represents at least one specimen, should
therefore be viewed as an approximation of species distributions rather than a completely accurate one. At the scale
of the maps, the dots are anyway several kilometers across. Quantitative variables were measured from specimens or
taken from specimen labels (where, in case of ranges, median values were used). Variables were counted or measured
with a ruler or digital calipers. Twenty-one quantitative variables were found and scored. Ten are vegetative and 11
from reproductive structures.
Completion of these stages led to a data matrix of specimens determined to preliminary species (using Glassman’s
names) and scored for location, quantitative, and qualitative variables. An additional column recorded the herbarium
in which the specimens examined are housed. In several cases, duplicates of specimens in different herbaria were
examined and scored. Herbarium abbreviations are from Thiers (2020, continuously updated). One other column gives
the country of origin of the specimens. The final matrix contains 902 specimens from 21 herbaria (https://www.nybg.
org/content/uploads/2020/03/Attalea.xlsx).
The next stage was to examine, preliminary species by preliminary species, the states of each qualitative
variable. State distributions were visualized using Winclada (Nixon 2004). There were two possible outcomes of this
examination.
First, the states of all qualitative variables were consistent within a preliminary species. These species, with
unique combinations of qualitative variables, were treated as phylogenetic species (and variables were then treated as
characters). Where two or more preliminary species had the same unique combination they were combined into one
phylogenetic species.
Second, one or more qualitative variable varied within a preliminary species. These were then split into two or
more preliminary species such that variables were consistent within each one. However, most polymorphic preliminary
species were based on a few, vegetative variables (e.g., petioles short or absent/petioles well-developed, or pinnae
regularly arranged/pinnae irregularly arranged), and splitting these into several, non–polymorphic species was found
to give improbable results, and no corroborating evidence that these were separate taxa was found (no geographic
separation, no quantitative differences). These variables were usually treated a priori as traits and were not used to split
preliminary species. Then, in most cases, none or only one variable was polymorphic within a preliminary species.
These polymorphic preliminary species were not split and recognized as polymorphic phylogenetic species, with some
qualitative variables treated as traits. In some cases, t-tests were run to test for differences between populations of
phylogenetic species.
Hybrids complicated this scoring. Thirty-six specimens were presumed to be hybrids, based on intermediate
morphology. Other specimens, possibly representing hybrids, were included in their relevant species. Some qualitative
variables, particularly the arrangement of rachillae on the rachis and flowers on the rachillae, were difficult to score,
possibly because of hybridization. Presumed hybrids and named hybrid species are discussed under the ‘Subspecific
variation’ section of the relevant species.
The result of these four stages was that preliminary species were tested as hypotheses and either supported or
not; and new hypotheses of characters and traits (i.e., phylogenetic species) were presented and were used in the
treatment.
Taxonomic treatment
A genus description of Attalea is given, based on qualitative and quantitative variables (see also section on Morphology
for an illustrated discussion of morphology). This is followed by a key to all species, based on quantitative variables,
qualitative variables, and geography.
For each species, arranged in alphabetical order, complete synonymy is given. Most types (or images of types)
of names of Attalea have been examined for this study and these are followed by a “!”. Those which have not been
examined are followed by “n.v.”. In a few cases epitypes are designated. Excluded and Doubtful Names are listed in
Appendix III. An index of all names is given (Appendix IV).
Descriptions of species are compilations of quantitative and the relevant qualitative variables. For quantitative
variables (in descriptions and keys), the mean of the variable is given followed by the range (e.g., stems 8.8(1.5–30.0)
m long, 43.7(19.0–75.0) cm diameter). If only a single value is given it is because only one value was recorded.
Descriptions are followed by notes on distribution and habitat, other notes, and discussion of subspecific variation, if
applicable. Under habitat, elevation data is given as the mean followed by the range (e.g., at 198(15–950) m).
In order to avoid confusion over generic names, when referring to taxa that Glassman (1999) and others recognized,
his generic names are used, with Attalea suffixed as s.s. (sensu stricto), hence Attalea s.s., Maximiliana, Orbignya, and
Scheelea.
In the References section, the work on palms by Martius, Historia Naturalis Palmarum, is cited by part rather
than volume. Although three volumes are found in libraries, these were not published in chronological order, and each
volume is made up of different parts and different editions published over a period of 30 years (Stafleu & Cowan
1981).
Results
Application of the Phylogenetic Species Concept to the 65 preliminary species recognized by Glassman (1999) resulted
in recognition of 30 phylogenetic species.
A discussion of Distribution and Habitat and Morphology of these species precedes the Taxonomic Treatment
section.
Distribution and Habitat
Attalea species are distributed from 21°32’N (Mexico) to 25°37’S (Paraguay) and 105°16’W (Mexico) to 35°37’W
(Brazil) (Fig. 1). The country with the highest number of species is Brazil with 20 followed by Colombia with 15. The
area with the highest number of species is the western Amazon basin of Peru in the Iquitos region.
Species of Attalea occur from sea level to 1,470 m (A. amygdalina in Colombia). However, most specimens are
from lower elevations. Of the 412 specimens with elevation data, 343 (83%) occur at ≤500 m.
Figure 1 shows the distribution of all specimens examined. What is striking is the paucity of specimens from the
Brazilian Amazon region. Apart from Acre, the area around Manaus, and the extreme eastern part of the region, there
are large areas of the Brazilian Amazon where there are no collections of Attalea. However, this lack of specimens
from the Amazon does emphasize a biogeographic division in Attalea between species from eastern Brazil (Cerrado
and Atlantic Coastal Forest; mostly Attalea s.s. and Orbignya) and those from the western Amazon region (mostly
Scheelea).
Species of Attalea are found in a variety of habitats. They occur in lowland or, rarely, in montane rainforest, mostly
in non–flooded areas but sometimes along river margins; in cerrado vegetation; in open savannas; or on stabilized sand
dunes. Some species seem to actively invade disturbed areas (e.g., A. butyracea, some forms of A. sagotii) and many
species persist and seem to flourish in disturbed areas.
Morphology
The genus description given below in the Taxonomic Treatment section is based on characters and traits (Appendix
II) and quantitative variables (Appendix I). In the following discussion, morphology is treated in more detail, and the
morphology of several attributes of Attalea not used in delimiting species is discussed. Most characters and states are
illustrated. A detailed generic description can also be found in Dransfield et al. (2008).
Stems of Attalea take two forms. One is a tall, aerial stem, here referred to as caulescent, and the other is a short,
subterranean stem here referred to as acaulescent. Caulescent species can have large stems up to 30.0 m long and
75.0 cm diameter. About half the species have caulescent stems and half acaulescent. However, the difference is not
absolute, and some species with otherwise acaulescent stems may have individuals or populations with caulescent
stems (e.g., A. amygdalina) and vice versa (e.g., A. funifera). Stems of Attalea appear to be always solitary, although in
a few instances labels describe stems as clustered. It is not clear if these are truly clustered or are perhaps examples of
two or more individual stems growing close together and misinterpreted as clustered. Pires-O’Brien (1993) described
plants of A. sagotii spreading by rhizomes.
Leaves of Attalea are always pinnate and consist of sheath, petiole, rachis, and pinnae. Sheaths are large and
woody on caulescent species, but are usually subterranean in acaulescent species and thus seldom present or recorded
on specimens. Margins of sheaths of caulescent species have short, flexible fibers. In one species, Attalea funifera,
long and rigid fibers are present.
There is a problem distinguishing sheaths from petioles. Petioles in a few caulescent species appear to be either
very short or absent (e.g., Attalea butyracea). However, sometimes labels state that petioles are up to a meter or more
in length. This appears to be based on a misinterpretation, because the apex of the sheath appears to be ‘petiolar’. In
such cases, what appears to be the petiole is in fact the sheath, with fibers along the margins. This difference between

sheaths and petioles is difficult to score from herbarium specimens. It also seems likely in some species that younger
plants (that are more often collected or measured) have longer petioles and older, taller plants have short or absent
petioles. Other caulescent and all acaulescent species appear to have well-developed petioles (without fibers on the
margins) and are here scored as such (with the exception of some plants of A. sagotii). However, even here there may
be problems. In A. amygdalina, for example, petioles, recorded on labels as being from 3.0–224.0 cm long, appear in
cultivated plants to be very short or absent, and what appears to be the petiole is in fact an extension of the sheath.
Rachises are well-developed in all species, ranging in length from 106.0–1,250.0 cm long. Pinnae are numerous
(48–318 per side of the rachis) and are mostly regularly arranged. Some species have both regularly and irregularly
arranged pinnae (e.g., Attalea butyracea), and in these pinnae can be irregularly arranged proximally on the rachis for
varying distances and then regularly arranged distally. In fact, most species appear to have the distalmost few pinnae
regularly arranged. In a few species (e.g., A. allenii, A. insignis) pinnae are arranged in distant clusters. Juvenile plants
of species with the adults having regularly arranged pinnae can also exhibit this arrangement (e.g., A. butyracea). The
proximalmost few pinnae in some caulescent species are closely spaced, stiff and fibrous, and in some acaulescent
species are distantly spaced and the pinnae elongate and pendulous. In a few species pinnae may have dense patches of
brown hairs on the abaxial and proximal part of mid-vein. In other species the abaxial mid-vein may have a continuous
cover of brown hairs.
Pinnae shape varies very little in Attalea, unlike most other palm genera of equal or greater size, and pinnae are
narrowly to broadly linear-lanceolate. Pinnae in most, if not all, species have a line of brown tomentum along the
abaxial, distal margin, and this appears broader at the pinnae apex. Apices of pinnae are briefly and asymmetrically
bifid. Such tomentose, asymmetric apices are found in other genera of the Attaleinae, and do not appear to be useful
taxonomically.
In a few species, pinnae have auricles at the bases. The auricle is here defined as a short extension (a few millimeters
in length) of the proximal margin of the pinna which is free from the abaxial surface of the rachis, instead of running
into the rachis. It is consistently present in a few species previously placed in the genus Scheelea (A. butyracea, A.
cephalotus, A. insignis, A. phalerata, A. plowmanii) and also in A. tessmannii. In some species with pinnae arranged in
distant clusters, an auricle is present only on the proximalmost pinnae of a cluster. However, the auricle is difficult to
score for several reasons. It often breaks off from herbarium specimens; it appears to be only present on the proximal
few pinnae; and it is not consistently present within some species. The majority of species do not have auricles, or
rather the proximal margin of the pinnae runs into the rachis.
In some acaulescent species the distalmost few pinnae remain joined at their apices, sometimes with ‘windows’
between them proximally. Some species (e.g., A. allenii, A. cuatrecasana, A. insignis) consistently exhibit this variable,
but others are inconsistent. In other acaulescent and all caulescent species the distalmost few pinnae are free from one
another.
Inflorescences in Attalea are said to be of three kinds, all of which are borne on the same plant (Henderson 2002);
with staminate flowers only, with staminate and pistillate flowers, or with pistillate flowers only (and then usually with
some sterile staminate flowers also present). The first and the third kinds are the most common, and here inflorescences
are scored as predominantly unisexual. However, in at least two species, A. crassispatha and A. maripa, bisexual
inflorescences commonly occur.
Inflorescences consist of a short prophyll, a much longer, woody peduncular bract, a short to long peduncle,
and a rachis with few to numerous rachillae. Staminate inflorescences usually have many more, longer rachillae than
pistillate inflorescences. Staminate rachillae are often covered with abundant raphides (Fig. 2A). There is a tendency
for rachillae to be absent from the adaxial surface of the rachis, although this character is difficult to score. Staminate
rachillae are absent from the adaxial surface of the rachis (Fig. 2A) in five species (A. amygdalina, A. eichleri, A.
nucifera, A. racemosa, A. septuagenata) and pistillate rachillae are absent from the adaxial surface of the rachis in five
species (A. amygdalina, A. eichleri, A. insignis, A. racemosa, A. septuagenata). All these except A. septuagenata are
acaulescent.
There is also a tendency for staminate and pistillate flowers to be absent from the adaxial surface of the rachillae.
Staminate flowers are absent from the adaxial surface of the rachillae (Fig. 2A) in most species, and only a few species
(e.g., A. cohune, A. cuatrecasana) have the staminate flowers spirally arranged around the rachillae. However, this
variable is not easy to score in some species (e.g., A. sagotii). Pistillate flowers are absent from the adaxial surface
of the rachillae in several species, especially those having longer pistillate rachillae (e.g., A. butyracea). Staminate
flowers are variously arranged in monads or dyads along the rachillae and are subtended by small bracteoles which
may be either ribbed and the same texture as the rachilla and continuous with it abaxially or crustaceous and different
in texture from the rachilla and appearing separate from it by an abaxial, proximal margin (Fig. 2B).
FIGURE 2. A. Staminate inflorescences: staminate rachillae absent from the adaxial surface of the rachis; rachillae with abundant
raphides; and staminate flowers mostly in monads, superficial on rachillae, loosely and singly arranged along abaxial surface of rachillae
(Attalea amygdalina, Moore 10199). B. Bracteoles: left side—staminate dyad bracteoles not ribbed, crustaceous, different in texture from
the rachilla and appearing separate from it by an abaxial, proximal margin; staminate flowers slightly sunken in rachillae, spirally arranged
all around rachillae (A. cohune, Moore 6199); right side—staminate monad/dyad bracteoles ribbed, the same texture as the rachilla and
continuous with it abaxially; staminate flowers in monads or dyads, superficial on rachillae, loosely and spirally or somewhat irregularly
arranged all around rachillae (sometimes fewer or absent from adaxial, proximal surface of rachillae) (A. tessmannii, Henderson 1691). C.
Staminate flowers: left—stamens much shorter than the petals; staminate petals linear, free, not curved at the apices, flat in cross-section
(A. butyracea, Galeano 2068); top right—stamens exserted from the petals; staminate petals linear, free, not curved at the apices, concave
in cross-section (A. maripa, Jansen-Jacobs 1060); bottom right—stamens neither exserted from nor much shorter than petals; staminate
petals linear, joined at the bases, curved over at the apices, flat in cross-section (A. luetzelburgii, Wessels Boer 2374). D. Staminate petals:
left—surfaces of petals smooth and coriaceous; petals irregularly shaped, joined at the bases, sometimes fused, curved over and split at the
apices, flat in cross-section (A. speciosa, Balick 1684); right—surfaces of petals ribbed and not coriaceous; petals lanceolate, acuminate,
flattened, free, not or only slightly curved at the apices, flat in cross-section (A. funifera, Noblick 4245). E. Staminate petal margins: top
left—margins of staminate petals smooth (A. iguadummat, de Nevers 7656); top right—margins of staminate petals minutely denticulate
(arrowed) (A. oleifera, Noblick 4590); bottom left—margins of staminate petals minutely serrate (arrowed) (A. allenii, Bernal 1024);
bottom right—margins of staminate petals tomentose (arrowed) (A. racemosa, Wessels Boer 1906). F. Anthers: top—coiled and twisted
(A. sagotii, Henderson 1165); bottom—straight (A. oleifera, Noblick 4537). G. Pistillate petals: left—cuspidate (A. cuatrecasana, Balick
1658); right—not cuspidate (A. plowmanii, Blicher 3). H. Pistillate petals: top left—with elaborate, lobed margins (arrowed) (A. allenii,
King 664); top right—with elaborate, lobed margins (A. barreirensis, Anderson 36868); bottom—with irregular margins (A. cuatrecasana,
Balick 1658). I. Staminodial rings: left—distal margins of staminodial ring smooth, membranous, not tomentose, with minute teeth
(arrowed), not apparently joined early in development to the ovary and usually not leaving a faint, circular, distal scar on fruit (A. funifera,
Mori 10879); right—distal margins of staminodial ring tomentose, without minute teeth, joined early in development to the ovary and
usually leaving a faint, circular, distal scar on fruit (A. cohune, Balick 1720). Scale bar = 1 cm.
Flowers in Attalea are always unisexual. Staminate flowers have three small sepals, three (rarely more or fewer)
long petals, six or more stamens (rarely fewer), and a small pistillode. The previous division of Attalea into several
genera based on the petals and stamens of their staminate flowers is simplistic and the situation is a great deal more
complex. In fact, staminate flower morphology is much more variable in Attalea than in most other palm genera. In
this study, staminate flowers provide seven characters divided into 33 states. Stamens range from 5–70 in number with
12 species consistently having six stamens. Stamens may be exserted from or much shorter than the variously shaped
petals (Fig. 2C). The surfaces of staminate petals may be smooth or ribbed (Fig. 2D) and the margins smooth, minutely
denticulate, minutely serrate, or tomentose (Fig. 2E). Anthers may be coiled and twisted or straight (Fig. 2F).
Pistillate flowers are larger and much less variable than staminate flowers. They consist of three large, cuspidate
sepals, three large, cuspidate or non-cuspidate petals (Fig. 2G), a prominent, cupular staminodial ring (in all but A.
crassispatha), and a gynoecium of 3-several connate carpels with prominent stigmas. There appears to be little variation
in sepals and petals, although in a few species petals have elaborate, lobed margins (Fig. 2H). Staminodial rings are
always present. They may be well-developed, cupular with entire margins (Fig. 2I), or well-developed, cupular with
lobed margins, or poorly developed and forming a low ring. However, they tend to split irregularly as fruits develop.
In some species, particularly those previously included in Orbignya, staminodial rings are somewhat tomentose at the
apex and appear joined early in development to the ovary and there leave a faint, circular scar (Fig. 2I). Other species,
particularly those previously included in Attalea s.s. have staminodial rings with smooth margins, often with minute
teeth, and are not joined to the ovary early in development (Fig. 2I). However, these variables are rather difficult to
score and may not be useful taxonomically.

FIGURE 3. A. Persistent pistillate perianth in fruit (Attalea speciosa, Balick 1696). B. Endocarp surfaces with stout, rounded, adherent
fibers (A. speciosa, Noblick 4614). C. Endocarp surfaces: left—surfaces uneven, longitudinally ridged (A. oleifera, Noblick 4590); surfaces
smooth, not ridged (hole near top of endocarp is a bruchid beetle exit hole) (A. maripa, Wessels Boer 474). D. Endocarps: top—surfaces
proximally with clear outlines of the three locule divisions, these marked by irregular, small pits (A. crassispatha, Henderson 805);
bottom—endocarps beaked at the apices (A. guianensis, Moore 9535). E. Endocarps: left—bases regularly rounded (A. maripa, Wessels
Boer 474); right—bases irregularly pointed, the endocarp becoming thinner and flanged (A. insignis, Henderson 856). F. Opercula:
top—superficial, not obscured by fibers (A. colenda, Little 21208); bottom—sunken in endocarps and obscured by persistent fibers
(arrowed) (A. racemosa, Cuello 575). G. Endocarps in cross-section: top—with abundant, scattered fibers (A. oleifera, Noblick 4779);
bottom—with abundant fibers in regular clusters (A. phalerata, Zardini 54047). H. Endocarps: top—1-seeded, the abortive locules visible
in cross-section appressed to fertile locule (arrowed); endocarps in cross-section without fibers (A. colenda, Little 21208); bottom—1-
several-seeded, the abortive locules visible in cross-section but not appressed to fertile locules (A. dubia, Pardini 83). I. Seeds: top—with
a well-developed central cavity in cross-section (A. colenda, Little 21208); bottom—seeds solid in cross-section (A. eichleri, Moreno 179).
Scale bar = 1 cm.
Fruit size in Attalea ranges from 29.9–143.1 mm long and 18.5–99.3 mm diameter. Fruit color is most commonly
given on labels as brown, but a few species have fruits that mature orange or yellow. In this study, fruits, and especially
endocarps, provide numerous characters. Most of these taxonomically useful characters have been previously
overlooked, presumably because they are obscured by the tough and persistent epicarp, mesocarp, and fruiting perianth
(Fig. 3A). The fruiting perianth in some specimens can cover up to half or more of the fruit (Fig. 3A). Fruits of
all Attalea species are covered in a thin layer of brown tomentum, and in A. septuagenata fruits are grayish-white
tomentose at the apices. Fruits in A. cephalotus are elongate and angled by mutual pressure and are borne on very
short rachillae with only one fruit per rachilla. Endocarps are covered with stout, adherent fibers (Fig. 3B) which are
difficult to remove. Endocarps are thick and woody and usually uneven externally with longitudinal ridges, although
a few species have smooth endocarp surfaces (Fig. 3C). Endocarps may be beaked or rounded at the apices (Fig. 3D)
and at the base may have the outlines of three locule divisions marked by small, irregular pits (Fig. 3D). Most species
of Attalea have similar endocarp bases which are irregularly pointed, with the endocarp becoming thinner and flanged
toward the base (Fig. 3E), although in a few species endocarp bases are regularly rounded or lobed (Fig. 3E). Opercula
are mostly sunken in the endocarps and obscured by the persistent endocarp fibers (Fig. 3F), but in a few species the
opercula are superficial (Fig. 3F). In cross-section, endocarps usually have abundant fibers (Fig. 3G) and in a few
species these fibers are regularly clustered (Fig. 3G).
Seed number ranges from one to seven (up to eight according to Noblick 1991). In three species (A. barreirensis,
A. colenda, A. crassispatha), endocarps are 1-seeded and the abortive locules are visible in cross-section appressed to
the fertile locule (Fig. 3H); in all other species endocarps are 1-several-seeded and the abortive locules, when present,
are visible but not appressed to fertile locules (Fig. 3H). In one species (A. colenda) seeds have a well-developed
central cavity, visible in cross-section (Fig. 3I); in all other species they are solid or sometimes with a small, irregular
central cavity. Germination is remote and eophylls are undivided.
Taxonomic Treatment
Attalea Kunth in Humboldt, Bonpland & Kunth (1816: 309). Type:—Attalea amygdalina Kunth in Humboldt, Bonpland & Kunth
Maximiliana Martius (1826: 131), conserved name. Type:—Maximiliana regia Martius
Orbignya Martius ex Endlicher (1836–1840: 257). Type:—Orbignya phalerata Martius
Lithocarpus Targioni Tozzetti ex Steudel (1841: 56), illegitimate name. Type:—Lithocarpus cocciformis Targioni Tozzetti
Scheelea Karsten (1856: 264). Type:—Scheelea regia Karsten
Englerophoenix Kuntze (1891: 728). Type:—Englerophoenix regia (Martius) Kuntze
Pindarea Barbosa Rodrigues (1896: 17). Type:—Pindarea concinna Barbosa Rodrigues
Bornoa Cook (1939: 264), invalidly published name.
Temenia Cook (1939: 276), invalidly published name.
Heptandra Cook (1939: 277), invalidly published name.
Ethnora Cook (1940: 297), invalidly published name.

Parascheelea Dugand (1940: 10). Type:—Parascheelea anchistropetala Dugand
Sarinia Cook (1942: 78). Type:—Sarinia funifera (Martius) Cook
Ynesa Cook (1942: 71). Type:—Ynesa colenda Cook
Markleya Bondar (1957: 50). × Maximbignya (Bondar) Glassman (1999: 199). Type:—Markleya dahlgreniana Bondar
× Attabignya Balick, Anderson & Medeiros-Costa (1987a: 27). Type:— × Attabignya minarum Balick, Anderson & Medeiros-Costa
Plants 11.4(1.0–32.5) m tall. Stems 8.3(0.3–30.0) m long, 36.4(15.0–75.0) cm diameter, caulescent or acaulescent.
Leaves 14(4–40) per stem; sheaths 117.0(15.0–550.0) cm long; petioles 89.9(0.0–400.0) cm long, short or absent or
well-developed; rachises 531.3(106.0–1,250.0) cm long; pinnae 145(48–318) per side of rachis, regularly or irregularly
arranged, or arranged in distant clusters, with or without auricles at the bases, sometimes distalmost few pinnae remaining
joined at their apices and with ‘windows’ between them. Inflorescence peduncular bracts 139.1(31.0–354.0) cm long;
peduncles 69.7(3.3–215.0) cm long; staminate rachillae 207(19–1,000), 14.5(2.5–53.0) cm long, arranged all around
rachis or absent from adaxial surface, with abundant, few or no raphides; staminate monad/dyad bracteoles usually
ribbed, the same texture as the rachilla and continuous with it abaxially or not ribbed, crustaceous, different in texture
from the rachilla and appearing separate from it by an abaxial, proximal margin; staminate flowers 11.8(5.2–23.8) mm
long, variously arranged in monads, rarely dyads; staminate petals variously shaped, lanceolate or linear, sometimes
curved over at the apices, the surfaces smooth, coriaceous, ribbed, sometimes granular tomentose, the margins smooth
or minutely denticulate or minutely serrate; stamens 11(5–70), exserted from petals, or much shorter than petals, or
neither exserted from nor much shorter than petals; anthers straight or coiled and twisted; pistillate rachillae 143(14–
640), 10.0(0.2–40.0) cm long, arranged all around rachis or absent from adaxial surface, at least proximally; pistillate
petals cuspidate or not, with elaborate, lobed margins or with aculeate or irregular margins; staminodial rings poorly
developed, forming a low, membranous ring or usually well-developed, forming a prominent cupule, deeply lobed
or not lobed although often splitting irregularly as fruits develop; fruits 71.3(29.9–143.1) mm long, 41.1(18.5–99.3)
mm diameter, sometimes elongate and angled by mutual pressure, borne on very short rachillae with only one fruit
per rachilla, or not elongate and angled and borne on short to long rachillae with 1-several fruits per rachilla, brown
tomentose, sometimes grayish-white tomentose at the apices; endocarp surfaces with stout, rounded, adherent fibers
or smooth or with few, thin fibers, proximally usually with clear outlines of three locule divisions, these marked by
irregular, small pits, uneven, longitudinally ridged or smooth, not ridged, the bases irregularly pointed, the endocarp
becoming thinner and flanged, or regularly rounded, or regularly lobed; endocarps in cross-section with abundant fibers
or without or with few fibers, fibers, when present, in regular clusters or scattered or rarely in a few, irregular clusters,
1-seeded, the abortive locules visible in endocarp cross-section appressed to fertile locule or 1-several-seeded, the
abortive locules, when present, visible in endocarp cross-section but not appressed to fertile locules; opercula sunken
in endocarps and obscured by persistent fibers or opercula superficial or slightly sunken in endocarps, not obscured by
fibers; seeds 2(1–7), usually solid in cross-section, sometimes with a well-developed central cavity in cross-section.
Key to the species of Attalea
1. Endocarps surfaces smooth or with few, thin fibers; endocarps in cross-section without or with few fibers; opercula superficial or
slightly sunken in endocarps, not obscured by fibers ........................................................................................................................2.
- Endocarp surfaces uneven, longitudinally ridged, with stout, rounded, adherent fibers; endocarps in cross-section usually with
abundant fibers; opercula sunken in endocarps and obscured by persistent fibers ...........................................................................6.
2. Staminate petals lanceolate, joined at the bases, curved outward at the apices, flat in cross-section; staminodial rings poorly devel-
oped, forming a low ring; southwestern Haiti ............................................................................................................. A. crassispatha
- Staminate petals not as above; staminodial rings well-developed, cupular ......................................................................................3.
3. Staminate flowers in dyads, slightly sunken in rachillae, irregularly arranged on abaxial surface of rachillae; staminate petals lin-
ear, joined at the bases, curved over at the apices, flat in cross-section; staminodial rings with lobed margins; Colombia (Guainía,
Vaupés), Venezuela (Amazonas), and Brazil (Amazonas) ........................................................................................ A. luetzelburgii
- Staminate flowers not as above; staminate petals not as above; staminodial rings with entire, rarely lobed margins .....................4.
4. Leaves arranged in five vertical ranks; stamens exserted from petals; Amazon region and adjacent areas of Colombia, Venezuela,
Guyana, Surinam, French Guiana, Trinidad, Brazil, Ecuador, Peru, and Bolivia ............................................................... A. maripa
- Leaves not arranged in five vertical ranks; stamens not exserted from petals ................................................................................. 5.
5. Stems acaulescent; petioles well-developed; northeastern Amazon region of Surinam, French Guiana, and Brazil (Amazonas)......
....................................................................................................................................................................................... A. guianensis
- Stems caulescent; petioles short or absent; Pacific coast of southern Colombia and Ecuador ...........................................A. colenda
6. Surfaces of staminate petals smooth, coriaceous; anthers coiled and twisted ...................................................................................7.
- Surfaces of staminate petals ribbed, not coriaceous; anthers straight .............................................................................................11.
7. Stems caulescent, 8.3(1.5–25.0) m long ............................................................................................................................................8.

- Stems acaulescent, 1.0 m long ...........................................................................................................................................................9.
8. Staminate flowers in dyads, spirally arranged all around rachillae; staminate petals spathulate, free, curved over at the apices;
Pacific coast of Mexico (Colima, Guerrero, Nayarit, Oaxaca), the Caribbean slope of Mexico, Guatemala, Belize, and Honduras,
and the Magdalena valley of Colombia (Boyacá, Caldas) ...................................................................................................A. cohune
- Staminate flowers mostly in monads, rarely dyads, alternately arranged along abaxial surface of rachillae; staminate petals irregu-
larly shaped, joined at the bases, sometimes fused, curved over and often split at the apices; Surinam, Brazil (Amazonas, Bahia,
Ceará, Maranhão, Mato Grosso, Minas Gerais, Pará, Piauí, Rondônia), and Bolivia (Beni, Pando, Santa Cruz) ............ A. speciosa
9. Pinnae irregularly arranged and spreading in different planes; staminate rachillae absent from adaxial surface of rachis; pistillate
rachillae absent from adaxial surface of rachis, at least proximally; Brazil (Bahia, Goiás, Maranhão, Mato Grosso, Pará, Tocantins)
and Bolivia (Beni) ............................................................................................................................................................... A. eichleri
- Pinnae regularly arranged; staminate rachillae arranged all around rachis; pistillate rachillae arranged all around rachis............10.
10. Abaxial mid-vein with a continuous cover of brown hairs; staminate flowers in dyads, spirally arranged all around rachillae; fruits
119.9(111.5–143.1) mm long, 77.0(58.6–88.0) mm diameter; endocarps in cross-section with abundant fibers; Pacific coast of
Colombia (Cauca, Chocó, Nariño, Valle del Cauca) ..................................................................................................A. cuatrecasana
- Abaxial mid-vein without a continuous cover of brown hairs; staminate flowers mostly in monads, rarely dyads, spirally arranged
but tending to be absent from adaxial surface of rachillae, especially proximally; fruits 38.3(29.9–56.9) mm long, 23.7(20.1–31.2)
mm diameter; endocarps in cross-section without or with few fibers; northern Amazon region of Colombia, Venezuela, the Gui-
anas, Peru, and Brazil ........................................................................................................................................................... A. sagotii
11. Staminate flowers mostly in monads, rarely dyads, superficial on rachillae, alternately arranged along abaxial surface of rachillae;
pistillate petals with elaborate, lobed margins.................................................................................................................................12.
- Staminate flowers not as above; pistillate petals with irregular margins ........................................................................................14.
12. Staminate rachillae with abundant raphides; surfaces of staminate petals not granular tomentose; margins of staminate petals
smooth; endocarps in cross-section without or with few fibers; endocarps 1-seeded, the abortive locules visible in endocarp cross-
section appressed to fertile locule; Brazil (Bahia, Goiás) ............................................................................................ A. barreirensis
- Staminate rachillae without or with few raphides; surfaces of staminate petals granular tomentose; margins of staminate petals
minutely serrate; endocarps in cross-section with abundant fibers; endocarps 1-several-seeded, the abortive locules, when present,
visible in endocarp cross-section but not appressed to fertile locules .............................................................................................13.
13. Pinnae arranged in distant clusters and spreading in different planes; central and eastern Panama and western Colombia ...............
............................................................................................................................................................................................... A. allenii
- Pinnae regularly arranged; Colombia (Antioquia, Bolívar, Cundinamarca, Santander) in the Magdalena valley.............A. nucifera
14. Sheath margins with long, rigid fibers; Atlantic Coastal Forest of Brazil (Alagoas, Bahia, Sergipe) ................................A. funifera
- Sheath margins without long, rigid fibers .......................................................................................................................................15.
15. Staminate rachillae absent from adaxial surface of rachis; staminate flowers mostly in monads, superficial on rachillae, loosely and
singly arranged along abaxial surface of rachillae; pistillate rachillae absent from adaxial surface of rachis, at least proximally .....
.........................................................................................................................................................................................................16.
- Staminate rachillae arranged all around rachis, rarely absent from adaxial surface of rachis; staminate flowers not as above; pistil-
late rachillae arranged all around rachis, rarely absent from adaxial surface of rachis, at least proximally ...................................18.
16. Stems caulescent, 9.5 m long; fruits grayish-white tomentose at the apices; Colombia (Amazonas) .......................A. septuagenata
- Stems acaulescent or to 3 m long; fruits not grayish-white tomentose at the apices ......................................................................17.
17. Abaxial mid-vein with a continuous cover of brown hairs; staminate rachillae with abundant raphides; margins of staminate petals
smooth; Cauca valley in Colombia (Antioquia, Valle del Cauca) ................................................................................ A. amygdalina
- Abaxial mid-vein without a continuous cover of brown hairs; staminate rachillae without or with few raphides; margins of sta-
minate petals tomentose; Amazon region in Colombia (Amazonas, Caquetá, Guainía, Guaviare), Venezuela (Amazonas), Peru
(Loreto), and Brazil (Amazonas) ......................................................................................................................................A. racemosa
18. Stamens 6, much shorter than petals ...............................................................................................................................................19.
- Stamens usually more than 6, not much shorter than petals............................................................................................................23.
19. Stems usually acaulescent; distalmost few pinnae remaining joined at their apices, sometimes with ‘windows’ between them;
pistillate petals cuspidate .................................................................................................................................................................20.
- Stems usually caulescent; distalmost few pinnae free from one another; pistillate petals not cuspidate ........................................21.
20. Pinnae arranged in distant clusters and spreading in different planes; pistillate rachillae absent from adaxial surface of rachis, at
least proximally; western Amazon region of Colombia (Amazonas, Caquetá, Casanare, Guaviare, Meta), Brazil (Acre, Amazonas),
Ecuador (Napo), and Peru (Loreto) .....................................................................................................................................A. insignis
- Pinnae usually regularly arranged; pistillate rachillae arranged all around rachis; Colombia (Amazonas) and Peru (Loreto) ...........
........................................................................................................................................................................................ A. plowmanii
21. Staminate rachillae 26.4(10.5–53.0) cm long, the flowers tending to be absent from adaxial, proximal surface of rachillae; endo-
carp fibers scattered; Mexico, Guatemala, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Trinidad and Tobago, Brazil,
Ecuador, Peru, and Bolivia .............................................................................................................................................. A. butyracea
- Staminate rachillae 9.6(6.0–15.0) cm long, the flowers tending to be absent from adaxial surface of rachillae; endocarp fibers in
regular clusters.................................................................................................................................................................................22.
22. Fruits 87.4(63.6–110.6) mm long, elongate and angled by mutual pressure, borne on 1.3(0.5–2.0) cm long rachillae, with only 1
fruit per rachilla; Colombia (Amazonas), Brazil (Acre), Peru (Huánuco, Loreto, Madre de Dios, San Martín), and Bolivia (Pan-
do) ...................................................................................................................................................................................A. cephalotus
- Fruits 61.3(48.4–79.1) mm long, not elongate and angled, borne on 4.6(2.0–7.7) cm long rachillae with 1-several fruits per rachil-
la; Brazil (Acre, Goiás, Mato Grosso, Mato Grosso do Sul, Maranhão, Pará, São Paulo, Tocantins), Peru (Junín, Madre de Dios),
Bolivia (Beni, La Paz, Pando, Santa Cruz), and Paraguay (Amambay, Concepción, Presidente Hayes) ........................A. phalerata
23. Abaxial and proximal part of pinnae mid-vein with dense patches of brown hairs; margins of staminate petals minutely denticu-
late ...................................................................................................................................................................................................24.

- Abaxial and proximal part of pinnae mid-vein without dense patches of brown hairs; margins of staminate petals not as above .....
.........................................................................................................................................................................................................29.
24. Stems acaulescent, 1.1(1.0–1.2) m long ..........................................................................................................................................25.
- Stems caulescent, 6.9(1.5–20.0) m long ..........................................................................................................................................26.
25. Stamens 8(7–9); staminodial rings with entire margins; Cerrado region of eastern Brazil (Bahia, Goiás, Mato Grosso, Minas
Gerais, São Paulo) and Paraguay (Amambay, Cordillera) ....................................................................................................A. exigua
- Stamens 6; staminodial rings with lobed margins; Atlantic Coastal Forest of Brazil (Bahia, Espírito Santo, Rio de Janeiro) ...........
............................................................................................................................................................................................. A. humilis
26. Petioles well-developed; Cerrado region of Brazil (Bahia) .......................................................................................... A. pindobassu
- Petioles short or absent ....................................................................................................................................................................27.
27. Staminate rachillae without or with few raphides; Atlantic Coastal Forest of Brazil (Alagoas, Bahia, Espírito Santo, Paraíba, Per-
nambuco, Sergipe) ...............................................................................................................................................................A. oleifera
- Staminate rachillae with raphides ....................................................................................................................................................28.
28. Pinnae irregularly arranged, usually spreading in different planes; Atlantic Coastal Forest of Brazil (Espírito Santo, Paraná, Rio de
Janeiro, Santa Catarina, São Paulo) ........................................................................................................................................ A. dubia
- Pinnae regularly arranged; Cerrado region of central Brazil (Goiás, Minas Gerais, Paraná, São Paulo) ........................... A. compta
29. Stems acaulescent; abaxial mid-vein with a continuous cover of brown hairs; Central Panama (Colón, San Blas) and northwestern
Colombia ..................................................................................................................................................................... A. iguadummat
- Stems caulescent; abaxial mid-vein without a continuous cover of brown hairs; Peru (Loreto, Madre de Dios) and Brazil (Acre) ..
........................................................................................................................................................................................ A. tessmannii
1. Attalea allenii Moore (1949: 191). Type:—PANAMA. Colón: Río Viejo, vicinity of Puerto Pilón, 10 m, 2 February
1947, P. Allen 4103 (holotype MO n.v., MO image!, isotype BH!)
Plants 4.2(3.3–5.0) m tall. Stems to 1 m long, diameter not recorded, acaulescent. Leaves 9(4–14) per stem; sheaths
not recorded; petioles 193.1(60.0–323.0) cm long; rachises 370.1(210.0–547.0) cm long; pinnae 104(55–152) per
side of rachis, arranged in distant clusters and spreading in different planes, usually without an auricle at the base;
middle pinnae 62.0(53.0–80.0) cm long, 3.4(2.2–4.7) cm wide; distalmost few pinnae remaining joined at their apices,
sometimes with ‘windows’ between them. Inflorescence peduncular bracts 54.0(48.0–60.0) cm long; peduncles
30.0(14.0–50.0) cm long; staminate rachillae number not recorded, 3.7(2.5–5.7) cm long, arranged all around rachis,
without raphides; staminate monad/dyad bracteoles not ribbed, crustaceous, different in texture from the rachilla and
appearing separate from it by an abaxial, proximal margin; staminate flowers 11.6(9.8–14.4) mm long, mostly in
monads, rarely dyads, superficial on rachillae, alternately arranged along abaxial surface of rachillae; staminate petals
lanceolate, acuminate, flattened, free, not or only slightly curved at the apices, flat in cross-section, the surfaces
ribbed, not coriaceous, granular tomentose, the margins minutely serrate; stamens 6, neither exserted from nor much
shorter than petals; anthers straight; pistillate rachillae number not recorded, 0.4(0.3–0.5) cm long, arranged all around
rachis; pistillate petals cuspidate, with elaborate, lobed margins; staminodial rings well-developed, cupular with entire
margins; fruits 74.8(59.4–91.6) mm long, 37.9(31.9–51.4) mm diameter, brown, light brown, brown-yellow, yellow,
or orange-yellow, not elongate and angled, borne on short rachillae with 1 fruit per rachilla; endocarp surfaces uneven,
longitudinally ridged, with stout, rounded, adherent fibers, without small pits proximally, the bases regularly lobed;
endocarps in cross-section with abundant, scattered fibers, 1-several-seeded, the abortive locules, when present, visible
in endocarp cross-section but not appressed to fertile locules; opercula sunken in endocarps and obscured by persistent
fibers; seeds 2(1–3), cross-section not recorded.
Distribution and habitat:—Central and eastern Panama and western Colombia at 139(15–350) m in lowland
rainforest (Fig. 4). The specimen in the Magdalena valley appears to be an outlier, but Galeano & Bernal (2010) show
a continuous distribution in Colombia.
Taxonomic notes:—Specimens determined as preliminary species Attalea allenii are polymorphic for auricles
and this variable is treated as a trait. Otherwise, specimens share a unique combination of qualitative variable states
and are therefore recognized as a phylogenetic species.
Morphologically, A. allenii appears most similar to A. nucifera, based on its granular, tomentose staminate petals
with minutely serrate margins, and pistillate petals with elaborate, lobed margins. Such pistillate petals are also found
in the similar A. barreirensis. It differs from both these species by its regularly lobed endocarp bases.
Glassman (1999) placed Attalea allenii in Attalea s.s. and considered it to be most closely related to A. exigua,
but the two do not seem to be related. Attalea allenii was polyphyletic in Freitas et al. (2016). They considered that
specimens from Colombia were sister to A. nucifera, whereas those from Panama were sister to A. guianensis (as
A. attaleoides). Based on morphology, this latter relationship seems unlikely. There are no quantitative differences

between the Colombian and Panamanian populations. Attalea allenii was not included by Pinheiro (1997) nor by
Meerow et al. (2009).
Subspecific variation:—Pistillate rachillae are scored as being arranged all around the rachis, although there is a
tendency in some specimens for them to be absent from the proximalmost, adaxial surface.
FIGURE 4. Distribution maps of Attalea allenii, A. amygdalina, A. barreirensis (closed circles = A. barreirensis, open circles, open
squares = unusual specimens, see text for details), and A. butyracea.
2. Attalea amygdalina Kunth in Humboldt, Bonpland & Kunth (1816: 310). Lectotype (designated by Glassman
1977a):—COLOMBIA. No locality, no date, F. Humboldt & A. Bonpland s.n. (holotype P!)
Attalea victoriana Dugand (1953: 9). Type:—COLOMBIA. Valle: Correg. Galicia y Ceilan, east of Bugalagrande, sitio El Almendronal,
June 1945, V. Patiño s.n. (holotype COL n.v., isotype F!).
Attalea uberrima Dugand (1953: 4). Type:—COLOMBIA. Caldas: Mun. Palestina, January 1942, R. Jaramillo Mejía 199 (holotype COL
n.v.).
Attalea rhynchocarpa Burret (1935: 617). Type:—COLOMBIA. Valle: Rio Frio bei Salonique, no date, E. Dryander s.n. (holotype B,
destroyed).

Plants height not recorded. Stems to 3.0 m long, 22.0 cm diameter, acaulescent or sometimes caulescent. Leaves
10(5–15) per stem; sheaths 75 cm long; petioles 88.4(3.0–224.0) cm long; rachises 739.0(615.0–910.0) cm long;
pinnae 146(126–172) per side of rachis, regularly arranged, usually with an auricle at the base; abaxial mid-vein with
a continuous cover of brown hairs; middle pinnae 88.8(85.5–92.0) cm long, 5.5(4.7–5.9) cm wide; distalmost few
pinnae remaining joined at their apices, sometimes with ‘windows’ between them, or pinnae free from one another.
Inflorescence peduncular bracts 195.3(125.0–327.0) cm long; peduncles 113.7(60.0–215.0) cm long; staminate
rachillae 251(250–252), 16.0(12.0–19.0) cm long, absent from adaxial surface of rachis, with abundant raphides;
staminate monad/dyad bracteoles usually ribbed, the same texture as the rachilla and continuous with it abaxially;
staminate flowers 15.0(11.5–16.9) mm long, mostly in monads, superficial on rachillae, loosely and singly arranged
along abaxial surface of rachillae; staminate petals lanceolate, acuminate, flattened, free, not or only slightly curved at
the apices, flat in cross-section, the surfaces ribbed, not coriaceous, not granular tomentose, the margins smooth; stamens
13(7–16), neither exserted from nor much shorter than petals; anthers straight; pistillate rachillae 50, 16.8(11.0–28.0)
cm long, absent from adaxial surface of rachis; pistillate petals cuspidate, with irregular margins; staminodial rings
well-developed, cupular with entire margins; fruits 88.1(70.9–105.6) mm long, 47.4(40.4–55.0) mm diameter, brown
or ferruginous, not elongate and angled, borne on long rachillae with 1 fruit per rachilla; endocarp surfaces uneven,
longitudinally ridged, with stout, rounded, adherent fibers, without small pits proximally, the bases irregularly pointed,
the endocarp becoming thinner and flanged; endocarps in cross-section with abundant fibers, 1-several-seeded, the
in endocarps and obscured by persistent fibers; seeds 2(1–3), solid in cross-section, rarely with an irregular central
cavity.
Distribution and habitat:—Cauca valley in Colombia (Antioquia, Valle del Cauca) at 1,354(1,200–1,470) m in
montane rainforest and persisting in disturbed areas (Fig. 4).
Taxonomic notes:—Specimens determined as preliminary species Attalea amygdalina are polymorphic for
stems, auricles, and distalmost pinnae and these variables are treated as a traits. Otherwise, specimens share a unique
combination of qualitative variable states and are therefore recognized as a phylogenetic species. Type specimens of
Attalea victoriana and A. uberrima share the same combination and are therefore included as synonyms. Glassman
(1999) considered Attalea rhynchocarpa to be an uncertain species based on the lack of a type specimen and incomplete
protologue. However, Galeano & Bernal (2010) placed it as a synonym of A. amygdalina and this is followed here.
Humboldt et al. (1816) wrote of Attalea amygdalina “Crescit in Provincia Chocoensi propre Zitara. Colitur in
hortis propre Carthago et Guaduas” (It grows in Chocó Province near Zitara. Cultivated in gardens near Cartago
and Guaduas). Dugand (1953), citing a paper by Karsten (1856) considered that Humboldt and Bonpland may have
confused two different species in Colombia. Guadas is in Cundinamarca, and is the type locality of another Attalea
species, A. nucifera, and this may have been the species that Humboldt and Bonpland saw cultivated there. Cartago is
in the Cauca valley and is within the range of A. amygdalina, as understood here. Apparently, Humboldt and Bonpland
were told that the cultivated Cartago plant came from Zitara (modern day Citará, an area that Humboldt and Bonpland
did not visit). Humboldt and Bonpland collected a specimen from Cartago, the type of A. amygdalina. This specimen
may have served in part for the illustration of this species in Humboldt et al. (1816, plate 96, fig. 1) and was lectotypified
by Glassman (1977a). However, Glassman cited this specimen as “Colombia, prov. Choco near Zitará, Humboldt &
Bonpland s.n.”, although there is no indication of its origin on the label, and it is probably not from Citará.
Morphologically, Attalea amygdalina appears most similar to A. racemosa and A. septuagenata. All three species
have staminate flowers loosely and singly arranged along the abaxial surface of rachillae, and pistillate rachillae
absent from the adaxial surface of the rachis. Glassman (1999) placed A. amygdalina in Attalea s.s., and considered it
to be most closely related to A. racemosa. Freitas et al. (2016) placed A. amygdalina in their Scheelea-like clade, and
there within a small clade with the polyphyletic A. butyracea, to which it bears little morphological resemblance. This
species was not included by Pinheiro (1997) nor by Meerow et al. (2009).
Subspecific variation:—Specimens occur in two separate areas but there are too few specimens to test for
differences, and the two occurrences may be an artifact of insufficient collecting. Galeano & Bernal (2010) showed a
continuous distribution. Although stems are usually recorded as acaulescent, one specimen is recorded as having a 3.0
m long stem, and Galeano & Bernal (2010) recorded stems up to 10 m long. Petioles are here scored as being present,
and the range is 3.0–224.0 cm long. However, cultivated plants were observed to lack petioles, but to have elongate
sheaths, with fibrous margins, and these appeared ‘petiolar’. This might account for the wide range in petiole length,
and a true petiole may be absent.

3. Attalea barreirensis Glassman (1999: 25). Type:—BRAZIL. Bahia: Mun. Barreiras, 28 km west of Barreiras,
August 1976, S. Glassman 13048 (holotype F!, isotype SP n.v.)
Plants height not recorded. Stems length and diameter not recorded, acaulescent. Leaves 6(5–8) per stem; sheaths
not recorded; petioles 22.9(3.1–36.0) cm long; rachises 147.0(115.0–174.0) cm long; pinnae irregularly arranged and
spreading in different planes, 85(67–104) per side of rachis, without an auricle at the base; abaxial and proximal
part of mid-vein with or without dense patches of brown hairs; middle pinnae 32.6(25.0–40.0) cm long, 3.6(2.4–5.0)
cm wide; distalmost few pinnae free from one another. Inflorescence peduncular bracts 32.6(31.0–35.0) cm long;
peduncles 11.4(6.0–17.0) cm long; staminate rachillae 47(41–57), 5.0(3.5–8.7) cm long, arranged all around rachis,
with abundant raphides; staminate monad/dyad bracteoles not ribbed, crustaceous, different in texture from the rachilla
and appearing separate from it by an abaxial, proximal margin; staminate flowers 11.6(9.9–13.1) mm long, mostly in
lanceolate, acuminate, flattened, free, not or only slightly curved at the apices, flat in cross-section, the surfaces ribbed,
not coriaceous, not granular tomentose, the margins smooth; stamens 6(5–6), neither exserted from nor much shorter
than petals; anthers straight; pistillate rachillae 21(18–24), 0.8(0.5–1.0) cm long, arranged all around rachis; pistillate
petals cuspidate, with elaborate, lobed margins; staminodial rings well-developed, cupular with entire margins; fruits
50.3(42.9–56.2) mm long, 34.4(30.6–37.4) mm diameter, gray-brown or reddish-orange, not elongate and angled,
borne on short rachillae with 1 fruit per rachilla; endocarp surfaces uneven, longitudinally ridged, with stout, rounded,
adherent fibers, proximally with clear outlines of three locule divisions, these marked by irregular, small pits, the
bases irregularly pointed, the endocarp becoming thinner and flanged; endocarps in cross-section without or with few
fibers, 1-seeded, the abortive locules visible in endocarp cross-section appressed to fertile locule; opercula sunken in
endocarps and obscured by persistent fibers; seeds 1, solid in cross-section, rarely with an irregular central cavity.
Distribution and habitat:—Brazil (Bahia, Goiás) at 695(510–840) m in cerrado vegetation (Fig. 4).
Taxonomic notes:—Specimens determined as preliminary species Attalea barreirensis are polymorphic for
abaxial mid-vein, and this variable is treated as a trait. Otherwise specimens share a unique combination of qualitative
variable states and are therefore recognized as a phylogenetic species, A. barreirensis. This is one of the few species
of Attalea having 1-seeded endocarps with appressed abortive locules and few or no fibers. Morphologically, A.
barreirensis appears most similar to A. nucifera and A. allenii (see notes under that species).
Glassman (1999) placed Attalea barreirensis in Attalea s.s. and considered it to be most closely related to A.
allenii and A. exigua. Freitas et al. (2016) placed it in their Orbignya-like clade, and there in a small clade with A.
moorei, and this clade as sister to Attalea maripa. However, A. barreirensis bears little morphological resemblance to
either A. moorei or A. maripa. Attalea barreirensis was not included by Pinheiro (1997) nor by Meerow et al. (2009).
Subspecific variation:—There are two outlying populations that bear a strong resemblance to Attalea
barreirensis, but are not included in the above description. Three specimens (Noblick 4960, 4961, Pereira-Silva 3432)
from Maranhão and Piauí (Fig. 4, open circles) are similar to A. barreirensis vegetatively and in habitat. However, the
only specimen with fruits (Noblick 4960) has scattered endocarp fibers and 2-seeded fruits with the abortive locules
not appressed to the fertile locules (unlike A. barreirensis without endocarp fibers and 1-seeded fruits with the abortive
locules appressed to the fertile locule).
Four specimens (Almeida 1201, Capucho 525, Lobo 109, Pardini 70) from further north from the eastern Amazon
region of Pará (Fig. 4, open squares) are unusual in several respects, and differ in their habitat. They resemble A.
barreirensis in their acaulescent habit, staminate and pistillate inflorescences, and flowers and fruits. One specimen
(Almeida 1201) has pinnae arranged in distant clusters and the distalmost few pinnae remain joined at their apices,
with ‘windows’ between them (unlike A. barreirensis), and two others have regularly arranged, much longer pinnae
(unlike A. barreirensis) and the distalmost ones are free. They are possibly hybrids with A. sagotii (see notes under that
species). More specimens are needed to understand this variation.
4. Attalea butyracea (Mutis ex Linnaeus f.) Wessels Boer (1988: 312). Cocos butyracea Mutis ex Linnaeus (1782:
454). Scheelea butyracea (Mutis ex Linnaeus) Karsten ex Wendland in Kerchove de Denterghem (1878: 256). Neotype
(here designated):—COLOMBIA. Cundinamarca: Municipio de Nilo, margen derecha del río Sumapaz, 7.5 km al
NE de Melgar en la carretera de Fusagasugá, 5°14’N, 74°37’W, 320 m, 2 September 2003, R. Bernal, G. Galeano, C.
Gamba, N. Pulgarín & N. Villalba 3448 (neotype COL!)
Attalea maracaibensis Martius (1844: 124). Scheelea maracaibensis (Martius) Burret (1929a: 541). Type:—VENEZUELA. Maracaibo,
no date, A. Plée s.n. (holotype P, destroyed). Neotype (designated by Glassman 1999):—Martius 1845: t. 167 fig. 3.

Attalea blepharopus Martius (1844: 116). Scheelea blepharopus (Martius) Burret (1929a: 541). Lectotype (designated by Moraes &
Pintaud 2016a):—Martius 1844, t. 5, fig. 2. Epitype (designated by Moraes & Pintaud 2016a):—BOLIVIA. Cochabamba: Prov.
Villa Tunari, Isinota, 1644’S 6538W, 203 m, 10 April 2015, M. Moraes & J.-C. Pintaud 2518 (epitype LPB n.v., isoepitypes BOLV
n.v., G n.v., NY!, P n.v.).
Scheelea regia Karsten (1856: 266). Lectotype (designated by Glassman 1977c):—Karsten 1866: t. 176, figs. 1–6.
Scheelea macrocarpa Karsten (1856: 268). Attalea macrocarpa (Karsten) Linden (1881: 15). Lectoype (designated by Wessels Boer
1988):—VENEZUELA. Without locality, without date, H. Karsten s.n. (lectotype LE n.v.).
Attalea rostrata Oersted (1858: 50). Scheelea rostrata (Oersted) Burret (1929a: 541). Type not designated.
Attalea humboldtiana Spruce (1869: 163). Scheelea humboldtiana (Spruce) Burret (1929a: 541). Type:—VENEZUELA. Amazonas:
lower angle(?) of Casiquiare & Orinoco, January 1854, R. Spruce 43 (holotype K!, isotypes F!, P!).
Scheelea leandroana Barbosa Rodrigues (1891a: 19). Attalea leandroana (Barbosa Rodrigues) Zona (2002: 133). Lectotype (here
designated):—Barbosa Rodrigues 1891a: t. 7, 8B.
Scheelea osmantha Barbosa Rodrigues (1894: 24). Attalea osmantha (Barbosa Rodrigues) Wessels Boer (1988: 318). Lectotype (here
designated):—Barbosa Rodrigues 1894: t. 4B.
Scheelea kewensis Hooker (1897: ts. 7552–7553). Attalea kewensis (Hooker) Zona (2002: 133). Type:—UNITED KINGDOM. Cultivated
plant, October 1985, no collector, no number (holotype K!).
Attalea wallisii Huber (1906: 267). Scheelea walllisii (Huber) Burret (1929a: 657). Neotype (designated by Glassman 1977b):—BRAZIL.
Amazonas or Acre: Rio Purus or Rio Acre, no date, G. Huebner 163 (holotype B n.v., B image!).
Scheelea liebmannii Beccari (1916: 113). Cocos regia Liebmann in Martius (1853: 323). Attalea liebmannii (Beccari) Zona (2002: 133).
Lectotype (designated by Glassman 1977b):—MEXICO. Veracruz: Xicaltepec, Río Nautla, 1841, F. Liebmann 6560 (lectotype C
n.v.).
Scheelea bassleriana Burret (1929a: 655). Attalea bassleriana (Burret) Zona (2002: 132). Type:—PERU. Loreto: Yarina Cocha, 155 m,
10 November 1925, G. Tessmann 5490 (holotype B n.v., B image!, isotypes NY!, US!).
Scheelea passargei Burret (1929a: 671). Type:—VENEZUELA. Guayana, no date, S. Passarge s.n. (holotype B n.v., B image!).
Scheelea urbaniana Burret (1929a: 672). Lectotype (here designated):—TRINIDAD AND TOBAGO. Easterfield, 11 April 1911, W.
Broadway 4015 (lectotype F!, isolectotypes BM!, G n.v., G image!, K!) (the holotype at B was destroyed).
Scheelea stenorhyncha Burret (1929a: 675). Lectotype (designated by Glassman 1977b):—PERU. Loreto: Río Itaya, Soledad, 110 m, 1
July 1925, G. Tessmann 5256 (lectotype F!, isolectotype G n.v., G image!) (the holotype at B was destroyed).
Scheelea preussii Burret (1929a: 678). Type:—GUATEMALA. Pacific side, 1926, H. Preuss s.n. (holotype B n.v., B image!).
Scheelea brachyclada Burret (1929a: 680). Lectotype (designated by Glassman 1977b):—PERU. Loreto: Soledad, 110 m, 26 June 1925,
G. Tessmann 5237 (lectotype F!, isolectotype NY!) (the holotype at B was destroyed).
Scheelea tessmannii Burret (1929a: 682). Attalea peruviana Zona (2002: 133). Type:—PERU. Loreto: Iquitos, 100 m, 23 April 1925, G.
Tessmann 5085 (holotype B n.v., B image!, isotype NY!).
Scheelea costaricensis Burret (1929a: 684). Type:—COSTA RICA. Without locality, no date, C. Hoffmann s.n. (holotype B, destroyed).
Scheelea macrolepis Burret (1929a: 688). Attalea macrolepis (Burret) Wessels Boer (1988: 311). Type:—VENEZUELA. Bolívar: Yopal,
14 February 1902, S. Passarge 774 (holotype B, destroyed). Neotype (designated by Stauffer & Fariñas 2006):—VENEZUELA.
Bolívar: El Tigre cerca del Río Cuchivero, 100 m, 15 June 1940, L. Williams 13315 (neotype VEN n.v., isoneotypes F!, US!).
Scheelea zonensis Bailey (1933: 37). Type:—PANAMA. Barro Colorado Island, June 1931, L. Bailey & Z. Bailey 1 (holotype BH!).
Scheelea dryanderae Burret (1934: 1049). Type:—COLOMBIA. Valle: near Cali, no date, E. Dryander 12 (holotype B n.v.).
Scheelea lundellii Bartlett (1935: 46). Attalea lundellii (Bartlett) Zona (2002: 133). Type:—GUATEMALA. Petén: Monte Polol, 11 June
1933, C. Lundell 3752 (holotype MICH n.v., isotypes BH!, BM!, GH n.v., K!, US!).
Scheelea curvifrons Bailey (1947: 443). Type:—TRINIDAD AND TOBAGO. St. Patrick: Leasehold Reservation, 30 January 1946, L.
Bailey 124 (holotype BH!).
Scheelea magdalenica Dugand (1959: 1). Attalea magdalenica (Dugand) Zona (2002: 133). Type:—COLOMBIA. Magdalena: Santa
Marta, near Tucurinca, 40 m, no date, A. Schultze 707 (holotype B, destroyed). Lectotype (designated by Glassman 1977b):—
COLOMBIA. Magdalena: Santa Marta, 21 October 1898, H. Smith 2639 (lectotype GH n.v., isolectotypes F!, K!, MO n.v., NY!, P!,
US!).
Scheelea wesselsboerii Glassman (1999: 170). Attalea wesselsboerii (Glassman) Zona (2002: 133). Type:—VENEZUELA. Barinas: near
Barrancas, 17 August 1967, J. Wessels Boer 1990 (holotype, F!, isotypes NY!, U n.v.).
Scheelea salazarii Glassman (1999: 146). Attalea salazarii (Glassman) Zona (2002: 133). Type:—PERU. Loreto: Maynas, Río Itaya
below Munichi, 17 May 1960, H. Moore, A. Salazar & E. Smith 8478 (holotype USM n.v., isotype BH!).
Scheelea fairchildensis Glassman (1999: 163). Attalea fairchildensis (Glassman) Zona (2002: 132). Type:—UNITED STATES. Florida:
cultivated, July 1963, R. Read 900 (holotype BH!, isotype FTG n.v., FTG image!).

Plants 13.9(5.0–30.0) m tall. Stems 8.8(1.5–30.0) m long, 43.7(19.0–75.0) cm diameter. Leaves 22(12–40) per stem;
sheaths 156.0(87.0–317.0) cm long; petioles short or absent, 1.0(0.0–20.0) cm long; rachises 785.0(555.0–1110.0) cm
long; pinnae 190(89–273) per side of rachis, regularly or sometimes irregularly arranged, with an auricles at the base;
abaxial and proximal part of mid-vein without or rarely with dense patches of brown hairs; middle pinnae 122.5(96.5–
140.0) cm long, 4.5(2.0–7.6) cm wide; distalmost few pinnae free from one another. Inflorescence peduncular bracts
216.6(107.0–320.0) cm long; peduncles 130.5(46.0–200.0) cm long; staminate rachillae 161(50–250), 26.4(10.5–53.0)
cm long, arranged all around rachis, with abundant raphides; staminate monad/dyad bracteoles usually ribbed, the same
texture as the rachilla and continuous with it abaxially; staminate flowers 13.1(7.9–18.0) mm long, in monads or dyads,
superficial on rachillae, loosely and spirally or somewhat irregularly arranged all around rachillae (sometimes fewer or
absent from adaxial, proximal surface of rachillae); staminate petals linear, free, not curved at the apices, flat in cross-
section, the surfaces ribbed, not coriaceous, not granular tomentose, the margins smooth; stamens 6, much shorter
than petals; anthers straight; pistillate rachillae 189(86–300), 18.1(3.5–37.5) cm long, arranged all around rachis;
pistillate petals cuspidate, with irregular margins; staminodial rings well-developed, cupular with entire margins; fruits
63.6(42.7–115.5) mm long, 34.3(25.5–67.0) mm diameter, brown, brownish-orangeish-yellowish, yellow, yellowish,
dull yellow, orange, orangeish, brownish-orange, orange-brown, orange-red, or pale purple, not elongate and angled,
borne on long rachillae with several fruits per rachilla; endocarp surfaces uneven, longitudinally ridged, with stout,
rounded, adherent fibers, without small pits proximally, the bases irregularly pointed, the endocarp becoming thinner
and flanged; endocarps in cross-section with abundant, scattered fibers, 1-several-seeded, the abortive locules, when
present, visible in endocarp cross-section but not appressed to fertile locules; opercula sunken in endocarps and
obscured by persistent fibers; seeds 2(1–3), solid in cross-section, rarely with an irregular central cavity.
Distribution and habitat:—Mexico, Guatemala, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Trinidad
and Tobago, Brazil, Ecuador, Peru, and Bolivia at 198(15–950) m in lowland rainforest (especially along river margins),
seasonal forest, or savannas, and persisting in disturbed areas (Fig. 4).
FIGURE 5. Attalea butyracea, left, caulescent habit; right infructescence (Colombia, images by Gloria Galeano).
Taxonomic notes:—Specimens determined as preliminary species Scheelea butyracea are polymorphic for
pinnae arrangement and abaxial mid-vein, and these variables are treated as traits. Otherwise specimens share a unique
combination of qualitative variable states and are therefore recognized as a phylogenetic species, Attalea butyracea

(Fig. 5). Specimens of 15 other species of Scheelea (S. bassleriana, S. fairchildensis, S. kewensis, S. leandroana, S.
liebmannii, S. lundellii, S. macrocarpa, S. macrolepis, S. magdalenica, S. maracaibensis, S. osmantha, S. rostrata,
S. salazarii, S. tessmannii, S. wesselsboerii) recognized by Glassman (1999) share the same combination and are
therefore included as synonyms of A. butyracea (but see below for pinnae arrangement). Type specimens of Scheelea
blepharopus, S. brachyclada, S. costaricensis, S. curvifrons, S. dryanderae, S. humboldtiana, S. passargei, S. preussii,
S. regia, S. stenorhyncha, S. urbaniana, S. wallisii, and S. zonensis also share the same combination and are therefore
included as synonyms. A type specimen of Attalea butyracea was never designated and there is no original material at
MA, and a neotype is therefore designated.
Glassman (1999) distinguished the 16 species he recognized based on two qualitative variables (petiole, pinna
arrangement) and five quantitative variables (staminate rachilla length, staminate flower length, pistillate rachilla
length, fruit length, fruit diameter). There are many inconsistencies in Glassman’s treatment. For example, Glassman
considered the Central American S. rostrata to have petioles 0.6–1.2 m long, but Henderson 1805 from Costa Rica is
recorded as having no petiole. Similarly, a specimen from Guatemala (Steyermark 37869), available to but not cited
by Glassman, has no petiole, but is otherwise identical to S. lundellii which is said to have a petiole 50–65 cm long.
Scheelea osmantha is described in the key as having petioles to 44 cm long, but the label of one specimen (Wessels
Boer 1837) states “petiole almost lacking”. Hammel et al. (2003) distinguished A. rostrata in Costa Rica from A.
butyracea based on its petioles. However, these are described as having fibrous margins. The confusion here is over
the problem of distinguishing sheaths from petioles, as discussed in the Morphology section, and the distal part of the
sheath is often recorded on labels as the petiole. A true petiole in A. butyracea, at least in adult plants, appears to be
either very short or absent.
Similar confusion is found in pinna arrangement. For example, Glassman considered Attalea wesselsboerii from
Apure in Venezuela to have clustered pinnae, but Wessels Boer (1988) wrote: “las poblaciones…en los Edos. Apure
y Barinas, normalmente consisten de plantas con hojas algo irregularmente pinnadas… Sin embargo, se encuentra
alguna variación a saber, plantas con pinnas a intervalos casi regulares que crecen lado a lado de plantas con hojas mas
irregularment pinnadas.…”. (“the populations… in the states of Apure and Barinas, normally consist of plants with
somewhat irregularly pinnate leaves…However, one finds some variation, plants with pinnae at almost regular intervals
growing side by side with more irregularly pinnate leaves…”). Among specimens determined by Glassman as Scheelea
magdalenica, supposedly with regularly arranged pinnae, there are some specimens with irregularly arranged pinnae.
Furthermore, it is commonly recorded on specimen labels that pinnae are irregularly arranged along the proximal part
of the rachis and regularly arranged distally. In summary, pinna arrangement is not useful taxonomically in this species.
It appears more related to habitat; almost all specimens from lowland forest areas have regularly arranged pinnae,
and specimens from more seasonal areas tend to have more clustered pinnae. For quantitative variables, there are few
significant differences between any of the species recognized by Glassman.
Stauffer & Fariñas (2006) considered Attalea macrolepis to be distinct, based on an unique combination of
variables. However, these variables are all quantitative, and specimens of A. macrolepis are smaller in size than other
specimens, notably in their short pistillate rachillae. This is exactly what one would expect from plants growing in a
dryer, savanna habitat, and apart from size these specimens are the same as others of A. butyracea. However, Stauffer
& Fariñas reported staminate flowers with 6–9 stamens. The specimens with stamens have not been seen in the present
study. Moraes & Pintaud (2016a) recognized Attalea blepharopus as distinct from other species, but the variables they
listed as distinguishing features are the same as those of A. butyracea.
Morphologically, Attalea butyracea appears most similar to four other species (A. cephalotus, A. insignis, A.
phalerata, A. plowmanii) all previously included in Scheelea, having pinnae with auricles, linear staminate petals, and
stamens much shorter than the petals.
Glassman (1999) placed Attalea butyracea in Scheelea s.s. Pinheiro (1997) considered that, of the three specimens
he examined of A. butyracea (as Scheelea butyracea), two exhibited anatomical patterns of a Scheelea and Orbignya
mixture, suggesting they were of hybrid origin. Meerow et al. (2009) and Freitas et al. (2016) placed A. butyracea
in a clade with other species formerly included in Scheelea. However, A. butyracea was polyphyletic in Freitas et al.
(2016).
Subspecific variation:—Attalea butyracea is almost continuously distributed from Mexico to Bolivia, and
through northern South America to Trinidad and Tobago, and there are few geographic disjunctions. Those that do
occur may be artifacts of insufficient collecting. Nevertheless, A. butyracea is not particularly variable, apart from the
variation in pinna arrangement discussed above. Some specimens from Amazonian Ecuador have larger fruits than
others (>100 mm long), and some other specimens from adjacent areas also tend to have large fruits. On the other
hand, specimens from Central America tend to have smaller fruits (mean 51.7 mm long). This variation in fruit size
may, like pinnae arrangement, be based on habitat, with plants from wetter, forested areas having larger fruits. Fruit
color is also variable, most commonly brown, yellow, or orange. Some specimens from the western Amazon basin
have exceptionally long staminate rachillae (e.g., Tessmann 5088, Galeano 2068), and others have exceptionally thick
staminate rachillae (e.g., Bernal 1404, 2081).
5. Attalea cephalotus Poeppig ex Martius (1844: 119). Scheelea cephalotes (Poeppig ex Martius) Karsten (1856:
269). Lectotype (designated by Glassman 1977b):—PERU. San Martín: Maynas alto, Tocache, July 1830, E. Poeppig
2000 (lectotype M n.v., isolectotype BR n.v., BR image!) (the holotype at W was destroyed)
Scheelea moorei Glassman (1999: 127). Attalea moorei (Glassman) Zona (2002: 133). Type:—PERU. Huánuco: Prov. Leoncio Prado,
near Naranjilla, May 1960, H. Moore 8392 (holotype BH!, isotype USM n.v.).
Attalea phalerata Martius ex Sprengel var. concinna Moreno Suárez & Moreno Suárez (2003: 68). Type:—BOLIVIA. Pando: Prov.
Nicolás Suarez, Chiquipaya, 11°08’S, 69°04’W, 260 m, 24 June 2002, L. Moreno 252 (holotype JBSC n.v., isotype NY!).
FIGURE 6. Distribution maps of Attalea cephalotus, A. cohune, A. colenda, and A. compta.
Plants 10.6(5.0–32.5) m tall. Stems 5.5(2.0–10.0) m long, 29.0(25.0–35.0) cm diameter, caulescent or rarely
acaulescent. Leaves 18(13–26) per stem; sheaths 119.3(66.0–200.0) cm long; petioles absent; rachises 723.0(520.0–

900.0) cm long; pinnae 182(105–218) per side of rachis, regularly or rarely irregularly arranged, with auricles at the
bases; middle pinnae 99.3(89.0–113.0) cm long, 4.2(2.6–5.2) cm wide; distalmost few pinnae free from one another.
Inflorescence peduncular bracts 144.5(125.0–164.0) cm long; peduncles 77.0(54.0–100.0) cm long; staminate rachillae
430, 10.0(6.5–13.5) cm long, arranged all around rachis, with abundant raphides; staminate monad/dyad bracteoles
usually ribbed, the same texture as the rachilla and continuous with it abaxially; staminate flowers 9.3(7.8–13.6) mm
long, mostly in monads, superficial on rachillae, loosely and irregularly arranged and tending to be absent from adaxial
surface of rachillae; staminate petals linear, free, not curved at the apices, flat in cross-section, the surfaces ribbed,
not coriaceous, not granular tomentose, the margins smooth; stamens 6, much shorter than petals; anthers straight;
pistillate rachillae number not recorded, 1.2(0.5–2.0) cm long, arranged all around rachis; pistillate petals cuspidate,
with irregular margins; staminodial rings well-developed, cupular with entire margins; fruits 87.4(63.6–110.6) mm
long, 36.8(30.1–53.0) mm diameter, brown, elongate and angled by mutual pressure, borne on very short rachillae,
with only 1 fruit per rachilla; endocarp surfaces uneven, longitudinally ridged, with stout, rounded, adherent fibers,
without small pits proximally, the bases irregularly pointed, the endocarp becoming thinner and flanged; endocarps in
cross-section with abundant fibers in regular clusters, 1-several-seeded, the abortive locules, when present, visible in
endocarp cross-section but not appressed to fertile locules; opercula sunken in endocarps and obscured by persistent
fibers; seeds 3(1–4), solid in cross-section, rarely with an irregular central cavity.
Distribution and habitat:—Colombia (Amazonas), Brazil (Acre), Peru (Huánuco, Loreto, Madre de Dios, San
Martín), and Bolivia (Pando) at 291(100–1,100) m in lowland rainforest or more open, disturbed areas (Fig. 6).
Taxonomic notes:—Specimens determined as preliminary species Scheelea cephalotes are polymorphic for stems
and pinnae arrangement and these variables are treated as traits. Otherwise, specimens share a unique combination of
qualitative variable states and are therefore recognized as a phylogenetic species, Attalea cephalotus (Fig. 7). Specimens
determined as preliminary species S. moorei, and the type specimen of A. phalerata var. concinna, also share the same
combination and the names are included as synonyms (but see below for pinnae arrangement). Moreno Suárez and
Moreno Suárez (2003) gave a complete account of this taxon in Bolivia (as Attalea phalerata var. concinna).
FIGURE 7. Attalea cephalotus, left, caulescent habit; right, infructescence, with elongate fruits angled by mutual pressure, borne on very
short rachillae, with only one fruit per rachilla (Madre de Dios, Peru).
The original description of Attalea cephalotus by Martius (1844) described the pinnae as being irregularly

arranged. However, no specimens were cited nor illustrations given. The isolectotype specimen at BR, Poeppig 2000,
has staminate flowers but lacks leaves and fruits (the lectotype at M has not been seen). Martius (1845) illustrated part
of an infructescence and fruits of A. cephalotus (t. 169 II) . The short pistillate rachillae and long, angled fruits with
clustered endocarp fibers are so distinctive that there is little doubt about application of this name. However, almost
all specimens examined have regularly arranged pinnae (but see below), and the original description of the pinna
arrangement remains a problem.
Glassman (1999) described the staminate flowers of Attalea cephalotus (as Scheelea cephalotes) as being spirally
arranged around the rachillae. It is not clear where this information came from, since the only specimen he cited is
sterile. All the specimens examined here with staminate rachillae have the staminate flowers irregularly arranged and
tending to be absent from adaxial surface of rachillae. Even the original description of Martius (1844) described the
staminate flower arrangement as “in facie rachin communem spectante floribus denudati” (lacking flowers on the face
[of the rachilla] looking toward the rachis).
Morphologically, Attalea cephalotus appears most similar to four other species (A. butyracea, A. insignis, A.
phalerata, A. plowmanii) all previously included in Scheelea, having pinnae with auricles, linear staminate petals, and
stamens much shorter than the petals. Of these four species it appears most similar to A. phalerata, based on staminate
flower arrangement. Glassman (1999) included Attalea cephalotus in Scheelea. It was not included by Pinheiro (1997)
nor by Meerow et al. (2009). Freitas et al. (2016) included a single specimen that was placed in the Scheelea-like clade,
within which it was sister to A. barreirensis. Based on morphology this seems an unlikely relationship.
Subspecific variation:—One specimen (Alexiades 1292) appears to have slightly irregularly arranged pinnae.
6. Attalea cohune Martius (1844: 121). Orbignya cohune (Martius) Dahlgren ex Standley (1932: 3). Lectotype (here
designated):—Martius 1845: t. 167 IV
Cocos guacuyule Liebmann ex Martius (1853: 323). Orbignya guacuyule (Liebmann ex Martius) Hernández Xolocotzi (1949: 17). Attalea
guacuyule (Liebmann ex Martius) Zona (2002: 133). Type:—MEXICO. Oaxaca: pr. Guatulco, no date, F. Liebmann 6559 (holotype
C n.v., C image!).
Cocos cocoyule Martius (1853: 324). Lectotype (designated by Glassman 1977c):—MEXICO. Acapulco, no date, W. Karwinski s. n.
(holotype M n.v., M image!).
Orbignya dammeriana Barbosa Rodrigues (1903a: 62). Type:—BRAZIL. Rio de Janeiro, cultivated plant, no date, Anon 260 (holotype
destroyed). Neotype (designated by Glassman 1977c):—BRAZIL. Rio de Janeiro: cultivated plant, no date, A. Glaziou 16468
(holotype B n.v., B image!, isotypes BH!, C n.v., C image!, MO!).
Plants 11.6(8.0–20.0) m tall. Stems 3.9(2.0–7.0) m long, 36.5(30.0–47.0) cm diameter. Leaves 18(15–24) per stem;
sheaths 141.3(131.0–150.0) cm long; petioles absent; rachises 731.7(614.0–881.0) cm long; pinnae 187(177–216) per
side of rachis, regularly arranged, with or without auricles at the bases; abaxial and proximal part of mid-vein with or
without dense patches of brown hairs; middle pinnae 83.9(69.0–114.0) cm long, 3.8(3.2–4.8) cm wide; distalmost few
pinnae free from one another. Inflorescence peduncular bracts 282.0(210.0–354.0) cm long; peduncles 124.3(50.0–
203.0) cm long; staminate rachillae 128, 19.4(9.5–24.0) cm long, arranged all around rachis, with abundant raphides;
staminate monad/dyad bracteoles not ribbed, crustaceous, different in texture from the rachilla and appearing separate
from it by an abaxial, proximal margin; staminate flowers 8.9(5.2–13.0) mm long, in dyads, slightly sunken in rachillae,
spirally arranged all around rachillae; staminate petals spathulate, free, curved over at the apices, flat in cross-section,
the surfaces smooth, coriaceous, not granular tomentose, the margins smooth; stamens 27(19–34), neither exserted
from nor much shorter than petals; anthers coiled and twisted; pistillate rachillae 94(60–127), 10.9(6.0–25.0) cm
long, arranged all around rachis; pistillate petals cuspidate, with irregular margins; staminodial rings well-developed,
cupular, with entire margins; fruits 66.0(51.9–83.4) mm long, 41.6(36.3–51.0) mm diameter, brown, not elongate and
angled, borne on long rachillae with 1-several fruits per rachilla; endocarp surfaces uneven, longitudinally ridged, with
stout, rounded, adherent fibers, without small pits proximally, the bases irregularly pointed, the endocarp becoming
thinner and flanged; endocarps in cross-section with abundant, scattered fibers, 1-several-seeded, the abortive locules,
when present, visible in endocarp cross-section but not appressed to fertile locules; opercula sunken in endocarps and
obscured by persistent fibers; seeds 1(1–3), solid in cross-section, or with an irregular central cavity.
Distribution and habitat:—Along the Pacific coast of Mexico (Colima, Guerrero, Nayarit, Oaxaca), the
Caribbean slope of Mexico, Guatemala, Belize, and Honduras, and the Magdalena valley of Colombia (Boyacá,
Caldas) at 253(125–600) m in lowland rainforest or deciduous forest and persisting in disturbed areas (Fig. 6).
Taxonomic notes:—Specimens determined as preliminary species Orbignya cohune are polymorphic for auricles
and abaxial mid-veins and these variables are treated as traits. Otherwise, specimens share a unique combination of
qualitative variable states and are therefore recognized as a phylogenetic species, Attalea cohune (Fig. 8). Specimens
determined as preliminary species O. guacuyule, and the type specimens of Cocos cocoyule and Orbignya dammeriana,
also share the same combination and are included.
FIGURE 8. Attalea cohune, left, caulescent habit; right, staminate inflorescence (Honduras).
Specimens come from three separate areas: the Caribbean slope of Mexico, Guatemala, Belize, and Honduras; the
Pacific slope of Mexico; and the Magdalena valley of Colombia (Fig. 6). The Colombian specimens were unknown
to Glassman (1999), but he recognized the two Central American populations as separate species—Orbignya cohune
(Caribbean slope of Mexico, Guatemala, Belize, and Honduras) and O. guacuyule (Pacific slope of Mexico). He
considered that O. cohune differed from O. guacuyule in its longer staminate rachillae, longer staminate flowers, wider
staminate petals, longer pistillate rachillae, more stamens (24 versus 19–20), and inconspicuous endocarp fibers. There
are no significant differences between the two for staminate rachilla length, staminate flower length, and number of
stamens, and too few specimens to test for pistillate rachillae length. Staminate flower width and conspicuousness of
endocarp fibers have not been measured in the present study. Here, O. cohune has 29(23–34) stamens and O. guacuyule
has 23(19–27) stamens. Of the three other variables (pinnae width, fruit length, fruit width) with sufficient data for
testing, there are no significant differences between the Caribbean slope and Pacific slope populations. The single
Colombian specimen with staminate flowers has 28 stamens (24 reported in Galeano & Bernal 2010). Despite the
geographic disjunction (the Colombian population occurs more than 1,600 km from the nearest Honduran population,
and the two central American populations are more than 500 km apart), the three populations are recognized as one
species.
Attalea cohune appears similar to a group of species (A. cuatrecasana, A. eichleri, A. sagotii, A. speciosa)
having coiled and twisted anthers. Of these, it appears most similar to A. cuatrecasana based on its staminate flower
arrangement and spathulate staminate petals. Glassman placed Attalea cohune in Orbignya, and Meerow et al. (2009)
included it in their Orbignya group. Freitas et al. (2016) placed it in their Orbignya-like clade, as sister species to A.
cuatrecasana. Anatomically these two species are similar (Pinheiro 1997).

7. Attalea colenda (Cook) Balslev & Henderson (1987: 1). Ynesa colenda Cook (1942: 71). Type:—ECUADOR. Los
Rios: Hacienda Santa Lucia, Cantón Vinces, 50 m, 20 October 1934, Y. Mexia 6574 (holotype US!, isotype BH!)
Plants 28.6(25.0–30.0) m tall. Stems 15.4(10.0–30.0) m long, 43.0(27.0–63.0) cm diameter. Leaves 22(19–26) per
stem; sheaths 251.0(200.0–297.0) cm long; petioles absent; rachises 584.0(500.0–680.0) cm long; pinnae 145(131–168)
per side of rachis, regularly arranged, with or without auricles at the bases; middle pinnae 158.0(139.0–193.0) cm long,
6.5(4.3–8.5) cm wide; distalmost few pinnae free from one another. Inflorescence peduncular bracts 268.7(236.0–
290.0) cm long; peduncles 97.5(80.0–130.0) cm long; staminate rachillae 799(598–1,000), 22.6(15.5–27.0) cm long,
arranged all around rachis, with abundant raphides; staminate monad/dyad bracteoles usually ribbed, the same texture
as the rachilla and continuous with it abaxially; staminate flowers 11.8(10.8–12.4) mm long, mostly in monads, rarely
dyads, superficial on rachillae, alternately arranged along abaxial surface of rachillae; staminate petals lanceolate, free,
not curved at the apices, flat in cross-section, the surfaces smooth, coriaceous, not granular tomentose, the margins
smooth; stamens 10(9–12), neither exserted from nor much shorter than petals; anthers straight; pistillate rachillae
420(200–640), 26.8(20.0–40.0) cm long; pistillate petals cuspidate, with irregular margins; staminodial rings well-
developed, cupular with entire margins; fruits 48.7(44.9–51.8) mm long, 26.2(24.0–27.8) mm diameter, orange or
yellow, not elongate and angled, borne on long rachillae with 1-several fruits per rachilla; endocarp surfaces smooth
or with few, thin fibers, proximally with clear outlines of three locule divisions, these marked by irregular, small pits,
smooth, not ridged, the bases regularly rounded; endocarps in cross-section without or with few fibers, 1-seeded, the
abortive locules visible in endocarp cross-section appressed to fertile locule; opercula superficial or slightly sunken in
endocarps, not obscured by fibers; seeds 1, solid in cross-section or with an irregular central cavity.
Distribution and habitat:—Pacific coast of southern Colombia and Ecuador at 179(50–470) m in lowland
rainforest and persisting in disturbed areas (Fig. 6).
Taxonomic notes:—Specimens determined as preliminary species Ynesa colenda are polymorphic for auricles
and are therefore recognized as a phylogenetic species, Attalea colenda (Fig. 9).
FIGURE 9. Attalea colenda, left, caulescent habit; right, staminate inflorescence (Ecuador).

Attalea colenda has unique staminate petals (lanceolate, free, not curved at the apices, flat in cross-section).
Morphologically it does not appear to have any close relatives, although it is similar to similarly isolated species (A.
crassispatha, A. guianensis, A. luetzelburgii, A. maripa) in having superficial or slightly sunken opercula that are not
obscured by fibers.
Glassman (1999) placed Attalea colenda in a separate genus, Ynesa, and considered Ynesa colenda to be of
possible hybrid origin, although this was not supported by Pinheiro (1997). Freitas et al. (2016) placed it in their
Scheelea-like clade, and within this in a sub-clade with A. ferruginea and A. racemosa (the former here treated as a
synonym of the latter). Morphologically it has little in common with A. racemosa. Attalea colenda was not included
by Meerow et al. (2009).
8. Attalea compta Martius (1826: 137). Lectotype (designated by Glassman 1977a):—BRAZIL. Unknown state: no
locality, no date, M. Wied-Neuwied s.n. (holotype M n.v., M image!). Epitype (here designated):—BRAZIL. Minas
Gerais: Municipio de Vazante, 2 km SE of the city, Fazenda Jaburu, 18°01’S 46°54’W, 650-700 m, 29 March 1992, L.
Noblick 4855 (epitype BHCB n.v., isoepitype NY!)
Attalea monogyna Burret (1929a: 534). Lectotype (designated by Glassman 1977a):—BRAZIL. Goiás: Serra dos Pyreneos, 18 July 1894,
A. Glaziou 22269 (lectotype BR!, isolectotypes BH n.v., C n.v., F!, K!, MO!, P!, US!).
Attalea camposportoana Burret (1938: 257). Type:—BRAZIL. Minas Gerais: Serra de Mantiqueira, on road between Juiz de Fora and
Barbacena, by Palmyra, December 1937, M. Burret 17 (holotype B n.v., B image!, isotype RB n.v.).
Attalea brasiliensis Glassman (1999: 65). Type:—BRAZIL. Goiás: 23.5 km north of central Brasília, Fer Cal area, 24 August 1976, S.
Glassman & G. Eiten 13057 (holotype F!, isotype SP n.v.).
stem; sheaths 63.3(40.0–100.0) cm long; petioles 10.0(0.0–20.0) cm long; rachises 594.6(412.0–786.0) cm long;
pinnae 168(133–205) per side of rachis, regularly arranged, without auricles at the base, abaxial and proximal part
of mid-vein with dense patches of brown hairs; middle pinnae 94.0(78.5–119.0) cm long, 3.1(2.8–3.5) cm wide;
distalmost few pinnae free from one another. Inflorescence peduncular bracts 195.0(170.0–220.0) cm long; peduncles
43.2(11.0–88.0) cm long; staminate rachillae 173(150–195), 10.8(7.0–16.0) cm long, arranged all around rachis, with
abundant raphides; staminate monad/dyad bracteoles usually ribbed, the same texture as the rachilla and continuous
with it abaxially, or staminate monad/dyad bracteoles not ribbed, crustaceous, different in texture from the rachilla
and appearing separate from it by an abaxial, proximal margin; staminate flowers 13.1(9.0–15.8) mm long, mostly in
lanceolate, acuminate, flattened, free, not or only slightly curved at the apices, flat in cross-section, the surfaces ribbed,
not coriaceous, not granular tomentose, the margins minutely denticulate; stamens 8(7–10), neither exserted from nor
much shorter than petals; anthers straight; pistillate rachillae 235(200–275), 7.5 cm long; pistillate petals cuspidate,
long, 46.4(33.1–59.3) mm diameter, brown, not elongate and angled, borne on short rachillae with 1-several fruits
per rachilla; endocarp surfaces uneven, longitudinally ridged, with stout, rounded, adherent fibers, without small pits
proximally, the bases irregularly pointed, the endocarp becoming thinner and flanged; endocarps in cross-section with
abundant, scattered fibers, 1-several-seeded, the abortive locules, when present, visible in endocarp cross-section but
not appressed to fertile locules; opercula sunken in endocarps and obscured by persistent fibers; seeds 3(1–5), solid in
cross-section, rarely with an irregular central cavity.
Distribution and habitat:—Cerrado region of central Brazil (Goiás, Minas Gerais, Paraná, São Paulo) at
874(570–1,400) m in seasonal forest or more open areas (Fig. 6).
Taxonomic notes:—Specimens determined as preliminary species Attalea compta are polymorphic for staminate
monad/dyad bracteoles and this variable is treated as a trait (see discussion below). Otherwise specimens share a
unique combination of qualitative variable states and are recognized as a phylogenetic species, A. compta. Specimens
determined as preliminary species A. camposportoana and A. brasiliensis also share the same combination and are
included.
Glassman (1999) included A. camposportoana as a synonym of A. apoda. However, the type represents a tall-
stemmed palm and the name is here included as a synonym of A. compta. Glassman (1999) tentatively included A.
monogyna as a synonym of the acaulescent A. exigua (as A. geraensis). However, the type specimen of A. monogyna
clearly comes from a caulescent rather than an acaulescent palm, and it is here treated as a synonym of A. compta. The
lectotype of Attalea compta consists of several pinnae only, without precise locality. Given the confusion over the type,
as discussed by Glassman (1999) and Noblick (1991), a fertile epitype is here designated.

Attalea compta is one of six, morphologically similar species (A. compta, A. dubia, A. exigua, A. humilis, A.
oleifera, A. pindobassu), sharing pinnae with the abaxial and proximal part of mid-vein with dense patches of brown
hairs, and staminate petals with minutely denticulate margins. Of these, it appears most similar to A. oleifera (differing
in its staminate rachillae with raphides) and to A. dubia (differing in its regularly arranged pinnae). In fact, the only
character separating A. compta from A. dubia is pinnae arrangement and this is one of the few cases where pinnae
arrangement is treated as a character. Habitat of the two species also differs with A. compta occurring more inland in
the Cerrado region and A. dubia occurring in coastal regions in the Atlantic Coastal Forest.
Glassman (1999) placed Attalea compta in Attalea s.s., and considered it closely related to A. oleifera. Meerow et
al. (2009) placed it in the Attalea clade and Freitas et al. (2016) (as A. brasiliensis) in the Attalea-like clade with other
Attalea s.s. although it was polyphyletic in Freitas et al. This species was not included by Pinheiro (1997).
Subspecific variation:—Attalea compta remains a poorly known species. Several specimens (Balick 1691,
Glassman 13057, Noblick 4855) from the most northern part of the range are scored as having the staminate dyad
bracteoles not ribbed, crustaceous, and different in texture from the rachilla and appearing separate from it by an
abaxial, proximal margin. This is similar to the sympatric A. speciosa, and there may be introgression between these
two species here. Balick et al. (1987a) described a hybrid between A. compta and A. speciosa (as Orbignya oleifera)
from this area: × Attabignya minarum Balick, Anderson & Medeiros-Costa (1987a: 27). Attalea × minarum (Balick,
Anderson & Medeiros-Costa) Zona (2002: 133). Type:—BRAZIL. Minas Gerais: Mun. de Santa Fé, 11 km from town
of Santa Fé, near stream called Logro do Rio, on Fazenda Santa Maria, 30 November-1 December 1984, M. Balick, J.
Medeiros-Costa, J. Frazão & J. Vieira 1694 (holotype CEN n.v., isotype NY!).
9. Attalea crassispatha (Martius) Burret (1929b: 23). Maximiliana crassispatha Martius (1844: 110). Orbignya
crassispatha (Martius) Glassman (1999: 94). Lectotype (designated by Glassman 1977c): Plumier 1703 t. 1. Epitype
(here designated):—HAITI. Department du Sud: Fond des Nègres, 18°22’N 73°14’W, 11 November 1988, A.
Henderson & M. Aubry 805 (epitype NY!)
Plants height not recorded. Stems 9.3(8.0–10.0) m long, 33.0 cm diameter. Leaves 17(15–19) per stem; sheaths
140.0(130.0–150.0) cm long; petioles absent; rachises 360.0(320.0–400.0) cm long; pinnae 157(127–178) per side
of rachis, regularly arranged, without auricles at the bases; middle pinnae 96.5(75.0–111.0) cm long, 4.4(3.6–5.0)
cm wide; distalmost few pinnae free from one another. Inflorescence peduncular bracts not recorded; peduncles not
recorded; staminate rachillae arranged all around rachis, with abundant raphides; staminate monad/dyad bracteoles
usually ribbed, the same texture as the rachilla and continuous with it abaxially; staminate flowers 7 mm long, mostly
in monads, rarely dyads, superficial on rachillae, alternately arranged along abaxial surface of rachillae; staminate
petals irregularly shaped, joined at the bases, sometimes fused, curved over and often split at the apices, flat in cross-
section, the surfaces ribbed, not coriaceous, the margins smooth; stamens 8, neither exserted from nor much shorter
than petals; anthers coiled and twisted; pistillate rachillae number not recorded, 16.0(15.0–17.0) cm long, arranged
all around rachis; pistillate petals cuspidate, with irregular margins; staminodial rings poorly developed, forming a
low ring; fruits 38.6(36.0–40.7) mm long, 24.2(22.2–25.3) mm diameter, red, not elongate and angled, borne on
long rachillae with 1-several fruits per rachilla; endocarp surfaces smooth or with few, thin fibers, proximally with
clear outlines of three locule divisions, these marked by irregular, small pits, smooth, not ridged, the bases regularly
rounded; endocarps in cross-section without or with few fibers, 1-seeded, the abortive locules visible in endocarp
cross-section appressed to fertile locule; opercula superficial or slightly sunken in endocarps, not obscured by fibers;
seeds 1, with a well-developed central cavity in cross-section.
Distribution and habitat:—Southwestern Haiti at 203(50–300) m in secondary forest or disturbed areas (Fig.
10).
Taxonomic notes:—Specimens determined as preliminary species Orbignya crassispatha share a unique
combination of qualitative variable states and are recognized as a phylogenetic species, Attalea crassispatha (Fig. 11).
It is the only Caribbean species of Attalea and is known only from southwestern Haiti (Henderson & Balick 1991;
Timyan & Reep 1994).
Morphologically Attalea crassispatha does not appear to have any close relatives, although it is similar to similarly
isolated species (A. colenda, A. guianensis, A. luetzelburgii, A. maripa) in having superficial or slightly sunken opercula
that are not obscured by fibers.
Glassman (1999) placed Attalea crassispatha in Orbignya. He described the pinnae as being irregularly arranged,
which is not correct. Pinheiro (1997) considered it more similar to Attalea s.s. species. Meerow et al. (2009) placed it
as sister species to their Orbignya clade. Freitas et al. (2016) placed it in their Orbignya-like clade, as sister species to
a clade containing Orbignya species.

FIGURE 10. Distribution maps of Attalea crassispatha, A. cuatrecasana, A. dubia and A. eichleri (closed circles = A. eichleri, open
circles = A. × teixeriana).
10. Attalea cuatrecasana (Dugand) Henderson, Galeano & Bernal (1995: 265). Orbignya cuatrecasana Dugand
(1943: 285). Type:—COLOMBIA. Valle: Río Naya, Puerto Merizalde, 20 February 1943, J. Cuatrecasas 13980
(holotype COL!, isotype F!)
Plants 7.8(4.0–12.0) m tall. Stems 1 m long, diameter not recorded, usually acaulescent. Leaves 8(5–12) per stem;
sheaths 80.5(70.0–97.0) cm long; petioles 171.9(94.0–250.0) cm long; rachises 574.4(440.0–655.0) cm long; pinnae
97(85–112) per side of rachis, regularly arranged, without auricles at the bases; abaxial mid-vein with a continuous cover
of brown hairs; middle pinnae 131.0(116.0–146.0) cm long, 6.2(5.0–6.9) cm wide; distalmost few pinnae remaining
joined at their apices, sometimes with ‘windows’ between them. Inflorescence peduncular bracts 113.0(76.0–150.0)
cm long; peduncles 51.7(30.0–75.0) cm long; staminate rachillae 34(21–56), 8.5(4.7–13.0) cm long, arranged all
around rachis, with abundant raphides; staminate monad/dyad bracteoles not ribbed, crustaceous, different in texture
from the rachilla and appearing separate from it by an abaxial, proximal margin; staminate flowers 10.7(7.3–15.5)
mm long, in dyads, slightly sunken in rachillae, spirally arranged all around rachillae; staminate petals spathulate,

free, curved over at the apices, flat in cross-section, the surfaces smooth, coriaceous, not granular tomentose, the
margins smooth; stamens 21(19–23), neither exserted from nor much shorter than petals; anthers coiled and twisted;
pistillate rachillae 40, 2.1(1.0–4.5) cm long, arranged all around rachis; pistillate petals cuspidate, with aculeate or
irregular margins; staminodial rings well-developed, cupular with entire margins; fruits 119.9(111.5–143.1) mm long,
77.0(58.6–88.0) mm diameter, brown or light brown, not elongate and angled, borne on short rachillae with 1 fruit
per rachilla; endocarp surfaces uneven, longitudinally ridged, with stout, rounded, adherent fibers, proximally with
clear outlines of three locule divisions, these marked by irregular, small pits, the bases regularly rounded; endocarps
in cross-section with abundant fibers, several-seeded, the abortive locules, when present, visible in endocarp cross-
section but not appressed to fertile locules; opercula sunken in endocarps and obscured by persistent fibers; seeds
3(2–3), cross-section not recorded.
FIGURE 11. Attalea crassispatha, left, caulescent habit; right, predominantly staminate inflorescence (Haiti).
Distribution and habitat:—Pacific coast of Colombia (Cauca, Chocó, Nariño, Valle del Cauca) at 82(9–500) m
in lowland rainforest (Fig. 10).
Taxonomic notes:—Specimens determined as preliminary species Orbignya cuatrecasana share a unique
combination of qualitative variable states and are recognized as a phylogenetic species, Attalea cuatrecasana (Fig.
12).
Attalea cuatrecasana appears similar to a group of species (A. cohune, A. eichleri, A. sagotii, A. speciosa) having
coiled and twisted anthers. It is a remarkable species, having the largest fruits in the genus and unusual endocarp bases.
The outer surface of the endocarp is strongly ridged. Endocarps bear a strong resemblance to those of A. iguadummat.
Attalea cuatrecasana occurs in scattered localities but this may be an artifact of insufficient collecting.
Glassman (1999) placed Attalea cuatrecasana in Orbignya, and considered that it had few close relatives. Freitas
et al. (2016) placed it in the Scheelea-like clade, as sister species to A. cohune, and Pinheiro (1997) gave similar results.
It was not included in the study of Meerow et al. (2009).

FIGURE 12. Attalea cuatrecasana, left, staminate inflorescence; right, acaulescent habit and infructescence (Valle del Cauca, Colombia,
images by Gloria Galeano).
11. Attalea dubia (Martius) Burret (1929a: 516). Orbignya dubia Martius (1845: 304). Pindarea dubia (Martius)
Hawkes (1952: 191). Lectotype (designated by Glassman 1977a):—Martius 1845: t. 169, fig. 6. Epitype (here
designated):—BRAZIL. São Paulo: São Miguel, rodovia SP 250, 16 July 1992, R. Pardini & A. Marcato 83 (epitype
NY!)
Attalea indaya Drude (1881: 437). Neotype (designated by Dahlgren 1959):—BRAZIL. Rio de Janeiro: Corcovado, cultivated, 15 June
1876, A. Glaziou 8070 (holotype C n.v., C image!, isotypes BR!, F!, P!).
Pindarea concinna Barbosa Rodrigues (1896: 17). Attalea concinna (Barbosa Rodrigues) Burret (1929a: 537). Type:—BRAZIL. Rio de
Janeiro: cultivated plant, no date, Anon 108 (holotype destroyed). Lectotype (here designated):—Barbosa Rodrigues 1896: t. 4C.
Pindarea fastuosa Barbosa Rodrigues (1896: 23). Type:—BRAZIL. Rio de Janeiro: cultivated plant, no date, Anon 2153 (holotype
destroyed). Lectotype (here designated):—Barbosa Rodrigues 1896: t. 5A.
Plants 15.0(10.0–20.0) m tall. Stems 6.7(5.0–10.0) m long, 38.0(30.0–50.0) cm diameter. Leaves 35 per stem;
sheaths 150 cm long; petioles absent; rachises 472.5(345.0–600.0) cm long; pinnae 183(167–213) per side of rachis,
irregularly arranged, usually spreading in different planes, with or without auricles at the bases, abaxial and proximal
part of mid-vein with dense patches of brown hairs; middle pinnae 79.0 cm long, 3.3 cm wide; distalmost few pinnae
free from one another. Inflorescence peduncular bracts 120 cm long; peduncles 44.3(38.5–50.0) cm long; staminate
rachillae number not recorded, 20.5 cm long, arrangement not recorded, with abundant raphides; staminate monad/
dyad bracteoles usually ribbed, the same texture as the rachilla and continuous with it abaxially; staminate flowers
21.1 mm long, mostly in monads, rarely dyads, superficial on rachillae, alternately arranged along abaxial surface
of rachillae; staminate petals lanceolate, acuminate, flattened, free, not or only slightly curved at the apices, flat in
cross-section, the surfaces ribbed, not coriaceous, the margins minutely denticulate; stamens number not recorded,
neither exserted from nor much shorter than petals; anthers straight; pistillate rachillae number not known, 16.5 cm

long; pistillate petals cuspidate, with irregular margins; staminodial rings well-developed, cupular with entire margins;
fruits 48.5 mm long, 22.9 mm diameter, color not recorded, not elongate and angled, borne on short to long rachillae
with 1-several fruits per rachilla; endocarp surfaces uneven, longitudinally ridged, with stout, rounded, adherent fibers,
the surfaces without small pits proximally, the bases irregularly pointed, the endocarp becoming thinner and flanged;
endocarps in cross-section with abundant, scattered fibers, 1-seeded, the abortive locules, when present, visible in
endocarp cross-section but not appressed to fertile locules; opercula sunken in endocarps and obscured by persistent
fibers; seeds 1, cross-section not recorded.
Distribution and habitat:—Atlantic Coastal Forest of Brazil (Paraná, Rio de Janeiro, Santa Catarina, São Paulo)
at 141(10–394) m in lowland rainforest or disturbed areas (Fig. 10). Glassman (1999) reported this species also occurs
in the Brazilian state of Espírito Santo, but no specimens from there have been seen in this study.
Taxonomic notes:—Specimens determined as preliminary species Attalea dubia are polymorphic for auricles
and are recognized as a phylogenetic species, A. dubia (Fig. 13). Type specimens of Attalea indaya, Pindarea concinna
and Pindarea fastuosa also have this combination and are included in synonymy. The lectotype of Attalea dubia is an
illustration of a single fruit and therefore an epitype is designated.
FIGURE 13. Attalea dubia, caulescent habit, with irregularly arranged pinnae (São Paulo, Brazil).
Attalea dubia appears similar to a group of morphologically similar species (A. compta, A. exigua, A. humilis, A.
oleifera, A. pindobassu), sharing pinnae with the abaxial and proximal part of mid-vein with dense patches of brown
hairs, and staminate petals with minutely denticulate margins. Attalea dubia shares most qualitative variable states
with another species, A. compta. It differs from A. compta in its clustered rather than regularly arranged pinnae.
Glassman (1999) placed Attalea dubia in Attalea s.s. He considered it to be related to A. funifera. Pinheiro (1997)
considered A. dubia to be a problematic species. He wrote: “one of the specimens shows clear signs of crossings
between species of Attalea and Orbignya in its leaf anatomy.” He considered A. oleifera to be one of the possible
parents. Attalea dubia was not included in either Meerow et al. (2009) or Freitas et al. (2016).

12. Attalea eichleri (Drude) Henderson (1995: 143). Orbignya eichleri Drude (1881: 449). Type:—BRAZIL. Brésil
central, Sertão d’Amaroleité, September–October 1844, H. Weddell 2705 (holotype P!)
Orbignya urbaniana Dammer (1901: 23). Type:—BRAZIL. Goiás: Goyas, 1 August 1895, A. Glaziou 22265 (holotype C n.v., C image!,
isotypes F!, G n.v., G image!, K!, P!, excluding leaves).
Plants 1.2(1.0–1.5) m tall. Stems length and diameter not recorded, acaulescent. Leaves 5(4–6) per stem; sheaths
not recorded; petioles 47.2(30.0–62.0) cm long; rachises 155.7(106.0–217.0) cm long; pinnae 83(48–127) per side
of rachis, irregularly arranged, spreading in different planes, without auricles at the bases; middle pinnae 43.6(30.0–
49.2(33.0–65.0) cm long; peduncles 10.8(4.5–17.0) cm long; staminate rachillae 27(19–34), 7.7(6.5–8.7) cm long,
absent from adaxial surface of rachis, with abundant raphides; staminate monad/dyad bracteoles not ribbed, crustaceous,
different in texture from the rachilla and appearing separate from it by an abaxial, proximal margin; staminate flowers
7.1(5.2–9.4) mm long, mostly in monads, rarely dyads, slightly sunken in rachillae, alternately arranged along abaxial
surface of rachillae; staminate petals irregularly shaped, joined at the bases, sometimes fused, curved over and often
split at the apices, flat in cross-section, the surfaces smooth, coriaceous, not granular tomentose, the margins smooth;
stamens 18(14–21), neither exserted from nor much shorter than petals; anthers coiled and twisted; pistillate rachillae
23(14–34), 2.1(0.2–4.0) cm long, absent from adaxial surface of rachis, at least proximally; pistillate petals cuspidate,
long, 38.9(32.3–41.1) mm diameter, brown or dark brown, not elongate and angled, borne on short rachillae with 1
fruit per rachilla; endocarp surfaces uneven, longitudinally ridged, with stout, rounded, adherent fibers, without small
pits proximally, the bases irregularly pointed, the endocarp becoming thinner and flanged; endocarps in cross-section
with abundant, scattered fibers, several-seeded, the abortive locules, when present, visible in endocarp cross-section
but not appressed to fertile locules; opercula sunken in endocarps and obscured by persistent fibers; seeds 4(2–6), solid
in cross-section, rarely with an irregular central cavity.
Distribution and habitat:—Brazil (Bahia, Goiás, Maranhão, Mato Grosso, Pará, Tocantins) and Bolivia (Beni)
at 403(200–750) m in cerrado vegetation (Fig. 10).
Taxonomic notes:—Specimens determined as preliminary species Orbignya eichleri share a unique combination
of qualitative variable states and are recognized as a phylogenetic species, Attalea. eichleri (Fig. 14). Glassman (1999)
tentatively included Orbignya urbaniana as a synonym, noting that the regularly arranged pinnae of the type were
discordant. The type may be a mixed collection, and here the leaves are excluded. Then, the specimen has the same
combination of states and is included in synonymy.
Attalea eichleri appears similar to a group of species (A. cohune, A. cuatrecasana, A. sagotii, A. speciosa) having
coiled and twisted anthers. Of these it appears most similar to A. speciosa, differing in its acaulescent habit and
irregularly arranged pinnae.
Glassman (1999) placed Attalea eichleri in Orbignya. He considered it to be close to A. speciosa (as O. phalerata)
based on their staminate rachillae and flowers. This was supported by Pinheiro (1997), Meerow et al. (2009), and
Freitas et al. (2016).
Attalea eichleri forms hybrids with A. speciosa, named as Orbignya × teixeriana by Bondar (1954)(Fig. 10, open
circles): Orbignya × teixeriana Bondar (1954: 58). Attalea × teixeriana (Bondar) Zona (2002: 133). Type:—BRAZIL.
Maranhão: Caixas, 10 January 1952, G. Bondar s.n. (holotype RB n.v., isotype F!).
Balick et al. (1987b) gave a complete description of this hybrid in Brazil, and Moreno Suárez (1999) gave
a description of it in Bolivia. However, there are some confusing specimens which are not included in the above
description. One specimen (Balick 1313) from Maranhão in Brazil, cited by Balick et al. (1987b) as Attalea eichleri
has unusual fruits with lobed endocarp bases, unlike other specimens of A. eichleri. In fact, these endocarps resemble
more those of the unusual specimens (Noblick 4960, 4961, Silva 3432) of A. barreirensis (see discussion under that
species) from nearby in Maranhão and Piauí. There are other specimens (Pinheiro 891, 892, 894, 898) from further
north in Maranhão with similar, unusual endocarps. One other specimen (Pinheiro 8910) from the same locality has
staminate flowers and these are typical of those of A. eichleri. It seems possible that there may be three species
hybridizing in this part of Maranhão, but the problem is unresolved.

FIGURE 14. Attalea eichleri, left, acaulescent habit with irregularly arranged pinnae; right, infructescences with pistillate rachillae
absent from adaxial surface of rachis, at least proximally (Mato Grosso, Brazil).
13. Attalea exigua Drude (1881: 439). Lectotype (designated by Glassman 1999):—BRAZIL. Entre Goyaz and
Cujaba, November-December 1844, H. Weddell 2965 (holotype P!, isotype F!)
Attalea geraensis (“ceraensis”) Barbosa Rodrigues (1898a: 22). Lectotype (here designated):—Barbosa Rodrigues 1898a: t. 7.
Attalea guaranitica Barbosa Rodrigues (1899: 27). Lectotype (here designated):—Barbosa Rodrigues 1903a t. 57 B.
Attalea apoda Burret (1933: 105). Type:—BRAZIL. Minas Gerais: between Porto de Rio Paracatu and Piquiero, 29 July 1893, A. Glaziou
22266 (holotype P!, isotypes BR!, C n.v., G n.v., K!, MO!).
Plants 1.5 m tall. Stems 1.2 m long, 15.0 cm diameter, acaulescent. Leaves 6(5–7) per stem; sheaths 15.0 cm long;
petioles 22.5(3.0–60.0) cm long; rachises 174.0(135.0–229.0) cm long; pinnae 82(61–102) per side of rachis, regularly
arranged, without auricles at the bases, abaxial and proximal part of mid-vein with dense patches of brown hairs; middle
pinnae 48.4(36.0–73.5) cm long, 2.2(1.5–3.2) cm wide; distalmost few pinnae free from one another. Inflorescence
peduncular bracts 47.3(41.0–50.0) cm long; peduncles 12.6(3.3–18.0) cm long; staminate rachillae 36(27–60), 5.5(2.8–
7.0) cm long, arranged all around rachis, without or with few raphides; staminate monad/dyad bracteoles usually ribbed,
the same texture as the rachilla and continuous with it abaxially; staminate flowers 14.3(11.0–17.0) mm long, mostly
in monads, rarely dyads, superficial on rachillae, alternately arranged along abaxial surface of rachillae; staminate
petals lanceolate, acuminate, flattened, free, not or only slightly curved at the apices, flat in cross-section, the surfaces
ribbed, not coriaceous, not granular tomentose, the margins minutely denticulate; stamens 8(7–9), neither exserted
from nor much shorter than petals; anthers straight; pistillate rachillae 21(14–39), 3.0(1.0–6.0) cm long, arranged all
around rachis; pistillate petals cuspidate, with irregular margins; staminodial rings well-developed, cupular with entire
margins; fruits 65.5(43.1–77.1) mm long, 45.4(33.6–56.5) mm diameter, brownish-orange, brown with orange base,
reddish-brown, dark chocolate, dark brown, brown and gray, or russet to chestnut, not elongate and angled, borne on
short rachillae with 1 fruit per rachilla; endocarp surfaces uneven, longitudinally ridged, with stout, rounded, adherent

fibers, without small pits proximally, the bases irregularly pointed, the endocarp becoming thinner and flanged;
endocarps in cross-section with abundant, scattered fibers, several-seeded, the abortive locules, when present, visible
fibers; seeds 3(2–4), solid in cross-section, rarely with an irregular central cavity.
Distribution and habitat:—Cerrado region of eastern Brazil (Bahia, Goiás, Mato Grosso, Minas Gerais, São
Paulo) and Paraguay (Amambay, Cordillera) at 672(270–960) m in open vegetation (Fig. 15).
FIGURE 15. Distribution maps of Attalea exigua, A. funifera, A. guianensis, and A. humilis.
Taxonomic notes:—Specimens determined as preliminary species Attalea geraensis share a unique combination
of qualitative variables and are therefore recognized as a phylogenetic species, A. exigua (Fig. 16). The holotype of
A. exigua consists of five sheets, representing a small, acaulescent palm with regularly arranged pinnae and a small
inflorescence with typical Attalea s.s. staminate flowers with 9 stamens. One sheet contains an Orbignya eichleri
inflorescence and flowers and is excluded. However, Glassman (1999) described A. exigua as having clustered pinnae,
whereas all specimens examined, including the type, have regularly arranged pinnae. This misinterpretation was
repeated by Lorenzi et al. (2010), and the plants they described and illustrated as A. exigua appear to be A. eichleri. The
type specimen of A. apoda also shares this combination and is included as a synonym. Glassman (1999) recognized
Attalea apoda as a distinct species and described it as having a tall stem and 12–14 cm long staminate rachillae.

However, the type of A. apoda at P clearly states ‘acaule’ on the label and has short, 6.5 cm long staminate rachillae
(and regularly arranged pinnae) typical of other specimens of A. exigua.
FIGURE 16. Attalea exigua, left, acaulescent habit; right, infructescence (São Paulo, Brazil).
Attalea exigua appears morphologically similar to other species (A. compta, A. dubia, A. humilis, A. oleifera,
A. pindobassu) having pinnae with the abaxial and proximal part of mid-vein with dense patches of brown hairs,
and staminate petals with minutely denticulate margins. Within this group it is similar to A. oleifera, differing in its
acaulescent (versus caulescent) habit and well-developed (versus short or absent) petioles. Glassman (1999) considered
A. exigua similar to A. allenii and A. barreirensis, but morphologically it does not appear close to these two species.
Freitas et al. (2016) placed it (as polyphyletic A. geraensis) in the Attalea-like clade, as did Pinheiro (1997). It was not
included by or Meerow et al. (2009).
Subspecific variation:—There are too few data from most variables to test, but the Paraguayan population has
significantly shorter petioles (t-test, P <0.05).
14. Attalea funifera Martius (1826: 136). Sarinia funifera (Martius) Cook (1942: 78). Lectotype (designated by
Glassman 1977a):—Martius 1826: t. 95–96. Epitype (here designated):—BRAZIL. Bahia: Ilhéus, Fazenda Barra do
Manguinho, ramal a direita com entrada no km 10 da rod. Ilhéus-Olivença (BA001), 14°54’S 39°12’W, 7 August 1985,
L. Noblick & L. Silva 4245 (epitype HUEFS n.v., isoepitype NY!)
Attalea acaulis Burret (1933: 103). Attalea funifera var. acaulis (Burret) Bondar (1942: fig. 8). Type:—BRAZIL. Bahia: Bolandeiras bei
San Salvador, April 1932, E. Werdermann 3182 (holotype B n.v., B image!).
Leaves 8(5–14) per stem; sheaths 228.3(60.0–550.0) cm long, the margins with long, rigid fibers; petioles 170.0(44.0–
300.0) cm long; rachises 432.9(328.0–536.0) cm long; pinnae 127(105–151) per side of rachis, irregularly arranged
and spreading in different planes, without auricles at the bases; middle pinnae 87.3(56.5–113.0) cm long, 4.0(2.6–
5.2) cm wide; distalmost few pinnae free from one another. Inflorescence peduncular bracts 152.4(120.0–170.0) cm
long; peduncles 68.5(37.0–110.0) cm long; staminate rachillae 120(80–145), 14.5(9.0–20.5) cm long, arranged all
around rachis, without or with few raphides; staminate monad/dyad bracteoles usually ribbed, the same texture as the
rachilla and continuous with it abaxially; staminate flowers 20.1(15.5–23.8) mm long, mostly in monads, superficial
on rachillae, loosely and singly arranged along abaxial surface of rachillae; staminate petals lanceolate, acuminate,
flattened, free, not or only slightly curved at the apices, flat in cross-section, the surfaces ribbed, not coriaceous, the
margins smooth; stamens 6, neither exserted from nor much shorter than petals; anthers straight; pistillate rachillae
113(75–150), 16.3(10.0–19.0) cm long, arranged all around rachis; pistillate petals cuspidate, with irregular margins;
staminodial rings well-developed, cupular with entire margins; fruits 85.4(72.1–115.6) mm long, 51.7(40.0–66.7)
mm diameter, color not recorded, not elongate and angled, borne on long rachillae with 1-several fruits per rachilla;
endocarp surfaces uneven, longitudinally ridged, with stout, rounded, adherent fibers, the surfaces without small pits
Distribution and habitat:—Atlantic Coastal Forest of Brazil (Bahia, Sergipe) at 51(1–149) m in coastal areas,
often on stabilized sand dunes where it can form pure stands (Fig. 15). This species also occurs in the Brazilian state
of Alagoas (Noblick 1991) but no specimens from there have been seen in the present study.
Taxonomic notes:—Specimens determined as preliminary species Attalea funifera are polymorphic for stems (see
below). This variable is treated as a trait. Otherwise, specimens share a unique combination of qualitative variable states
and are treated as a phylogenetic species, A. funifera (Fig. 17). The type of Attalea acaulis also has this combination
and is included as a synonym.
FIGURE 17. Attalea funifera, left, caulescent habit with irregularly arranged pinnae; left, staminate inflorescence (Bahia, Brazil).
Attalea funifera stands somewhat apart from other similar species (A. compta, A. dubia, A. exigua, A. humilis, A.
oleifera, A. pindobassu) in its long, rigid fibers along the leaf sheath margins and lack of dense patches of brown hairs
on the abaxial and proximal part of the mid-vein. Glassman (1999) placed A. funifera in Attalea s.s., and considered it
most closely related to A. dubia. Pinheiro (1997) placed it in a clade with other Attalea s.s., as did Meerow et al. (2009)
and Freitas et al. (2016).

Subspecific variation:—Some specimens from the northern part of the range are reported to be acaulescent and
there is a possibility of clinal variation in this species (see Noblick 1991).
Two hybrids have been named. Bondar (1942) named a species that Glassman (1999) considered a hybrid between
Attalea funifera and A. oleifera:
Attalea piassabossu Bondar (1942: 30). Lectotype (designated by Glassman 1977a):—BRAZIL. Bahia: Salvador,
1940, G. Bondar 6 (lectotype F!).
Glassman (1999) named a hybrid between Attalea funifera and A. humilis:
Attalea × voeksii Noblick ex Glassman (1999: 191). Type:—BRAZIL. Bahia: Municipio Ilhéus, Fazenda Barra
do Manguinho, ramal a direita, com entrada no km 10 do Rodovia Ilhéus-Olivença (BA 001), 7 July 1985, L. Noblick,
L. Silva, T. Santos & M. Lemos 4241 (holotype CEPEC n.v., isotypes F!, HUEFS n.v.).
15. Attalea guianensis (Glassman) Zona (2002: 133). Scheelea guianensis Glassman (1999: 137). Type:—FRENCH
GUIANA. Haut Camopi: au nord du Mont Belvédère, 180 m, December 1984, J.-J. de Granville 7087 (holotype CAY
n.v., isotype F!)
Scheelea maripensis Glassman (1999: 140). Attalea maripensis (Glassman) Zona (2002: 133). Type:—FRENCH GUIANA. Maripa Soula:
Montagne Bellvue de l’Inini, 750 m, August 1985, J.-J. de Granville 7823 (holotype CAY n.v., isotype F!).
Plants 2.0 m tall. Stems length and diameter not recorded, acaulescent. Leaves 11(8–16) per stem; sheaths 30.0 cm
long; petioles 83.3(40.0–100.0) cm long; rachises 446.6(330.0–600.0) cm long; pinnae 82(67–105) per side of rachis,
regularly arranged, without auricles at the bases; middle pinnae 85.4(58.0–105.0) cm long, 3.2(2.3–4.5) cm wide;
distalmost few pinnae free from one another. Inflorescence peduncular bracts 72.7(68.0–80.0) cm long; peduncles 100
cm long; staminate rachillae number not recorded, 6.2(3.0–8.7) cm long, arranged all around rachis, with abundant
raphides; staminate monad/dyad bracteoles usually ribbed, the same texture as the rachilla and continuous with it
abaxially; staminate flowers 10.2(8.8–11.7) mm long, in monads or dyads, superficial on rachillae, loosely and spirally
or somewhat irregularly arranged all around rachillae (sometimes fewer or absent from adaxial, proximal surface of
rachillae); staminate petals linear, free, not curved at the apices, concave in cross-section, the surfaces ribbed, not
coriaceous, not granular tomentose, the margins smooth; stamens 6, neither exserted from nor much shorter than
petals; anthers straight; pistillate rachillae number not recorded, 1.8(1.5–2.0) cm long, arranged all around rachis;
pistillate petals cuspidate, with irregular margins; staminodial rings well-developed, cupular, with entire margins;
fruits 49.9(41.6–61.4) mm long, 25.1(22.3–27.8) mm diameter, yellow-brown, not elongate and angled, borne on
short rachillae with 1-several fruits per rachilla; endocarp surfaces smooth or with few, thin fibers, surfaces proximally
with clear outlines of three locule divisions, these marked by irregular, small pits, smooth, not ridged, the bases
regularly rounded; endocarps in cross-section without or with few fibers, 1-several-seeded, the abortive locules, when
present, visible in endocarp cross-section but not appressed to fertile locules; opercula superficial or slightly sunken in
endocarps, not obscured by fibers; seeds 2(1–3), solid in cross-section, rarely with an irregular central cavity.
Distribution and habitat:—Northeastern Amazon region of Surinam, French Guiana, and Brazil (Amazonas), at
353(130–750) m in lowland rainforest (Fig. 15).
Taxonomic notes:—Specimens determined as preliminary species Scheelea guianensis and S. maripensis share a
unique combination of qualitative variable states and are therefore recognized as a single phylogenetic species, Attalea
guianensis (Fig. 18). The name Attalea attaleoides (Barbosa Rodrigues) Wessels Boer was used for this species by
Wessels Boer (1965) and Henderson (1995). However, following Glassman (1999) it is here treated as an Excluded
Name.
Morphologically A. guianensis does not appear to have any close relatives, although it is similar to similarly
isolated species (A. colenda, A. crassispatha, A. luetzelburgii, A. maripa) in having superficial or slightly sunken
opercula that are not obscured by fibers. Specimens occur in two separate areas, central Brazil and the Guianas. Only
one specimen (de Granville 11604) from the Guianas has fruits and these are considerably larger than others.
Pinheiro (1999) placed Attalea guianensis (as S. attaleoides) in a Scheelea clade, close to A. insignis (as S.
insignis) and considered it represented a ‘true Scheelea’. However, Freitas et al. (2016) placed A. guianensis (as A.
attaleoides) in a small clade along with the polyphyletic A. allenii, this clade being embedded in the middle of their
Orbignya-like clade. Attalea guianensis bears little morphological resemblance to A. allenii or to species included in
Orbignya. Attalea guianensis was not included by Meerow et al. (2009).

FIGURE 18. Attalea guianensis, left, acaulescent habit; right, infructescence (Amazonas, Brazil).
Subspecific variation:—There appear to be several hybrids. Glassman (1999) described two species, the types of
which are here considered to be hybrids between Attalea guianensis and A. maripa:
Scheelea camopiensis Glassman (1999: 138). Attalea camopiensis (Glassman) Zona (2002: 132). Type:—FRENCH
GUIANA. Rivière Camopi S. Yanie, March 1974, J.-J. de Granville 2087 (holotype CAY n.v. CAY image!).
Scheelea degranvillei Glassman (1999: 139). Attalea degranvillei (Glassman) Zona (2002: 132). Type:—FRENCH
GUIANA. S de la Crique Martin, RN2 (route de l’Est) PK22, April 1983, J.-J. de Granville 5566 (holotype CAY n.v.,
CAY image!, isotype F!).
Other specimens appear to represent this hybrid. One (de Granville 9094) has similar staminate flowers to those
of the type of Scheelea degranvillei. Another (de Granville 11606) is described on the label as having a stem 1–10 m
tall and proximal pinnae irregularly arranged.
Henderson (1995) considered that A. maripa and A. guianensis (as A. attaleoides) produced hybrids in Reserva
Ducke, Brazil. Glassman (1999) considered that two specimens (Wessels Boer 1330, 1371) from Surinam, identified
by Wessels Boer (1965) as A. macropetala, were hybrids between A. maripa and a Scheelea (possibly A. guianensis).
16. Attalea humilis Martius ex Sprengel (1825: 624). Lectotype (designated here):—BRAZIL. State unknown: no
locality, no date, M. Wied-Neuwied s.n. (holotype M n.v., M image!)
Attalea compta var. acaulis Martius (1826: t. 75). Lectotype (here designated):—Martius 1826: t. 75.
Attalea borgesiana Bondar (1939: 16). Neotype (designated by Glassman 1977a):—BRAZIL. Bahia: Reconcavo de Bahia, Fazenda
Carmo, Mun. São Sebastião, no date, G. Bondar s.n. (holotype F n.v.).
Attalea borgesiana Hawkes (1952: 176), superfluous name.
Plants 3.5 m tall. Stems 1.0 m long, 37.0 cm diameter, acaulescent. Leaves 8(6–12) per stem; sheaths not recorded;
petioles 76.2(2.0–125.0) cm long; rachises 311.2(275.0–345.0) cm long; pinnae 107(97–123) per side of rachis,
regularly arranged, without auricles at the bases, abaxial and proximal part of mid-vein with dense patches of brown

hairs; middle pinnae 60.3(42.5–80.0) cm long, 3.0(2.1–3.9) cm wide; distalmost few pinnae free from one another.
83(74–96), 5.5(3.2–9.0) cm long, arranged all around rachis, without or with few raphides; staminate monad/dyad
bracteoles usually ribbed, the same texture as the rachilla and continuous with it abaxially; staminate flowers 15.6(11.9–
18.8) mm long, mostly in monads, rarely dyads, superficial on rachillae, alternately arranged along abaxial surface
cross-section, the surfaces ribbed, not coriaceous, not granular tomentose, the margins minutely denticulate; stamens
6, neither exserted from nor much shorter than petals; anthers straight; pistillate rachillae 75, 4.0 cm long, arranged
all around rachis; pistillate petals cuspidate, with irregular margins; staminodial rings well-developed, cupular, with
lobed margins; fruits 66.7(58.6–75.3) mm long, 45.1(39.3–55.2) mm diameter, brown or light brown, not elongate
and angled, borne on short rachillae with 1 fruit per rachilla; endocarp surfaces uneven, longitudinally ridged, with
thinner and flanged; endocarps in cross-section with abundant, scattered fibers, several-seeded, the abortive locules,
obscured by persistent fibers; seeds 3(2–3), solid in cross-section.
Distribution and habitat:—Atlantic Coastal Forest of Brazil (Bahia, Espírito Santo, Rio de Janeiro) at 6(2–10)
m in lowland rainforest or restinga forest (Fig. 15).
Taxonomic notes:—Specimens determined as preliminary species Attalea humilis share a unique combination
of qualitative variable states and are therefore recognized as a phylogenetic species, A. humilis (Fig. 20). The type
specimens of Attalea compta var. acaulis and Attalea borgesiana also share this combination and are included.
Morphologically A. humilis appears to be a member of a group of species (A. compta, A. dubia, A. exigua, A. oleifera,
A. pindobassu) having pinnae with the abaxial and proximal part of the mid-vein with dense patches of brown hairs and
staminate petals with minutely denticulate margins. It is distinguished from these by its staminodial ring with lobed
margins.
Attalea humilis was placed by Glassman (1999) in Attalea s.s., and was considered to be related to A exigua (as
A. geraensis). This position was supported by Pinheiro (1999), Meerow et al. (2009), and Freitas et al. (2016).
Glassman (1999) named a species and considered it could be a hybrid between Attalea humilis and A. oleifera:
Attalea salvadorensis Glassman (1999: 63). Type:—BRAZIL. Bahia: 78 km N of Salvador, near turnoff for
Itapetingui, along highway BR324, 19 August 1976, S. Glassman 13016 (holotype F!, isotypes K!, SP n.v.).
17. Attalea iguadummat de Nevers (1987: 506). Type:—PANAMA. Colón: Santa Rita Ridge 13.8 km from the
Transisthmica Highway, 9°20’N, 79°45’W, 350 m, 24 February 1986, G. de Nevers 7197 (holotype CAS n.v., isotypes
K!, MO!, PMA n.v.)
Plants height not recorded. Stems length and diameter not recorded, acaulescent. Leaves 14(13–15) per stem; sheaths
not recorded; petioles 62.5(60.0–65.0) cm long; rachises 738.3(689.0–815.0) cm long; pinnae 108(100–117) per side
of rachis, regularly arranged, without auricles at the bases; abaxial mid-vein with a continuous cover of brown hairs;
middle pinnae 116.3(108.0–123.0) cm long, 5.4(4.4–6.6) cm wide; distalmost few pinnae free from one another.
number not recorded, 11.5(9.5–13.5) cm long, arranged all around rachis, with abundant raphides; staminate monad/
15.4(13.7–16.6) mm long, in monads or dyads, superficial on rachillae, loosely and spirally or somewhat irregularly
arranged all around rachillae (sometimes fewer or absent from adaxial, proximal surface of rachillae); staminate petals
linear, free, not curved at the apices, flat in cross-section, the surfaces ribbed, not coriaceous, not granular tomentose,
the margins smooth; stamens 9(8–9), neither exserted from nor much shorter than petals; anthers straight; pistillate
rachillae number not recorded, 1.3 cm long, arranged all around rachis; pistillate petals cuspidate, with aculeate or
irregular margins; staminodial rings well-developed, cupular, with entire margins; fruits 92.3(92.0–92.6) mm long,
54.6(49.2–63.6) mm diameter, color not recorded, not elongate and angled, borne on short rachillae with 1 fruit per
rachilla; endocarp surfaces uneven, longitudinally ridged, with stout, rounded, adherent fibers, proximally with clear
outlines of three locule divisions, these marked by irregular, small pits, the bases regularly rounded; endocarps in
cross-section with abundant, scattered fibers, several-seeded, the abortive locules, when present, visible in endocarp
cross-section but not appressed to fertile locules; opercula sunken in endocarps and obscured by persistent fibers; seeds
3(2–3), cross-section not recorded.

FIGURE 19. Distribution maps of Attalea iguadummat, A. insignis, A. luetzelburgii, and A. maripa (closed circles = A. maripa, open
circles = Markleya dahlgreniana).
Distribution and habitat:—Central Panama (Colón, San Blas) at 339(65–500) m in lowland rainforest (Fig. 19).
Cano et al. (2017) recorded Attalea iguadummat from northwestern Colombia, but no specimens from there have been
seen in the present study.
Taxonomic notes:—Specimens determined as preliminary species Attalea iguadummat share a unique combination
of qualitative variable states and are therefore recognized as a phylogenetic species, A. iguadummat. It appears to be
a taxonomically isolated species without any obvious relatives. Specimens occur in two separate areas in Panama, but
there are too few to test for differences.
Glassman (1999) considered that Attalea iguadummat had staminate petals typical of other species of Attalea s.s.
However, staminate petals are here scored as linear, free, not curved at the apices, and flat in cross-section, i.e., typical
of Scheelea. Attalea iguadummat is also similar to some species described under Scheelea in its staminate flower
arrangement (loosely and spirally or somewhat irregularly arranged all around rachillae but tending to be absent from
adaxial, proximal surface of rachillae). Apart from its unusual staminate flowers and their arrangement, it has endocarp
surfaces proximally with clear outlines of three locule divisions, these marked by irregular, small pits. In fact, the
endocarps bear a remarkable resemblance to those of A. cuatrecasana. Although pistillate flowers with intact petals

are lacking in the material examined, old pistillate flowers appear to have petals with elaborate, lobed margins, like
those of the sympatric A. allenii. Attalea iguadummat was not included in the studies of Pinheiro (1999), Meerow et
al. (2009), or Freitas et al. (2016).
FIGURE 20. Attalea humilis, left, acaulescent habit; right, staminate inflorescence (Espírito Santo, Brazil).
18. Attalea insignis (Martius) Drude in Engler & Prantl (1897: 56). Maximiliana insignis Martius (1826: 133).
Scheelea insignis (Martius) Karsten (1856: 269). Englerophoenix insignis (Martius) Kuntze (1898: 322). Type:—
COLOMBIA. Caquetá: in sylvis Japuruensibus, no date, C. Martius s.n. (holotype M n.v., M image!)
Scheelea attaleoides Karsten (1856: 265). Lectotype (here designated):—Karsten 1856 t. 67.
Attalea goeldiana Huber (1906: 268). Scheelea goeldiana (Huber) Burret (1929a: 541). Type not designated.
48.0 cm long; petioles 280.6(164.0–370.0) cm long; rachises 572.6(430.0–750.0) cm long; pinnae 134(110–161) per
side of rachis, arranged in distant clusters and spreading in different planes, with auricles at the bases; middle pinnae
75.5(64.0–93.0) cm long, 3.8(2.8–4.6) cm wide; distalmost few pinnae remaining joined at their apices, sometimes with
‘windows’ between them. Inflorescence peduncular bracts 105.0 cm long; peduncles 40.0 cm long; staminate rachillae
46, 16.0(9.7–25.5) cm long, arranged all around rachis, with abundant raphides; staminate monad/dyad bracteoles
usually ribbed, the same texture as the rachilla and continuous with it abaxially; staminate flowers 11.8(10.0–15.8)
mm long, in monads or dyads, superficial on rachillae, loosely and spirally or somewhat irregularly arranged all around
rachillae (sometimes fewer or absent from adaxial, proximal surface of rachillae); staminate petals linear, free, not
curved at the apices, flat in cross-section, the surfaces ribbed, not coriaceous, not granular tomentose, the margins
smooth; stamens 6, much shorter than petals; anthers straight; pistillate rachillae number not recorded, 1.0(0.5–1.5) cm
long, absent from adaxial surface of rachis, at least proximally; pistillate petals not cuspidate, with irregular margins;
staminodial rings well-developed, cupular, with entire margins; fruits 66.7(55.0–79.3) mm long, 38.3(29.4–46.0) mm
diameter, cream, brown, or brown-red, not elongate and angled, borne on short rachillae with 1 fruit per rachilla;

endocarp surfaces uneven, longitudinally ridged, with stout, rounded, adherent fibers, the surfaces without small pits
abundant, scattered fibers, several-seeded, the abortive locules, when present, visible in endocarp cross-section but
Distribution and habitat:—Western Amazon region of Colombia (Amazonas, Caquetá, Casanare, Guaviare,
Meta), Brazil (Acre, Amazonas), Ecuador (Napo), and Peru (Loreto) at 184(100–300) m in lowland forest or gallery
forest in more open areas (Fig. 19).
Taxonomic notes:—Specimens determined as preliminary species Scheelea insignis share a unique combination
of qualitative variable states and are therefore recognized as a phylogenetic species, Attalea insignis (Fig. 21). Type
specimens of Scheelea attaleoides and Attalea goeldiana share the same combination and are included. Glassman
(1999) treated Attalea goeldiana as an uncertain species, based on the lack of staminate flowers. However, from the
locality and brief protologue, it appears to belong here.
FIGURE 21. Attalea insignis, staminate inflorescence, infructescence (Amazonas, Brazil).
Morphologically, Attalea insignis appears most similar to four other species (A. butyracea A. cephalotus, A.
phalerata, A. plowmanii), all previously included in Scheelea, having pinnae with auricles, linear staminate petals, and
stamens much shorter than the petals. However, only the proximal pinna of a cluster of pinnae has an auricle at the
base.
Glassman (1999) considered Scheelea insignis to be related to two other species, S. lauromuelleriana (here placed
as a synonym of A. phalerata) and S. leandroana (here placed as a synonym of A. butyracea). Pinheiro (1999) placed
it as sister to A. guianensis (as A. attaleoides). It was placed in the Scheelea-like clade of Freitas et al. (2016) but
is polyphyletic there. According to Freitas et al. the two separate groups of specimens are from separate geological
formations. This species was not included in the study of Meerow et al. (2009).
Subspecific variation:—Pistillate rachillae are scored as being arranged all round the rachis, although there is a
tendency in some specimens for them to be absent from the adaxial surface proximally. One specimen (Pardini 20) is
reported to have clustered stems. The label of one specimen (Bernal 2044) reports the plants to be invasive.

19. Attalea luetzelburgii (Burret) Wessels Boer (1988: 303). Orbignya luetzelburgii Burret (1930: 1019). Parascheelea
luetzelburgii (Burret) Dugand (1941: 24). Type:—BRAZIL. Amazonas: Jutica, Varadouro, Rio Ayari, Rio Uaupés, 16
November 1928, P. von Luetzelburg 21969 (holotype B n.v., B image!, isotypes M n.v., M image!, R n.v.)
Parascheelea anchistropetala Dugand (1940: 12). Type:—COLOMBIA. Vaupés: Cerro de Circasia, 300–500 m, 10 October 1939, J.
Cuatrecasas 7172 (holotype COL!).
Plants 2.5 m tall. Stems length and diameter not recorded, acaulescent. Leaves 6(5–6) per stem; sheaths 50 cm
long; petioles 96.7(50.0–150.0) cm long; rachises 475.0(460.0–490.0) cm long; pinnae 80(72–89) per side of rachis,
regularly arranged, without auricles at the bases; middle pinnae 68.3(44.0–84.5) cm long, 3.8(2.9–4.5) cm wide;
distalmost few pinnae remaining joined at their apices, sometimes with ‘windows’ between them or free from one
another. Inflorescence peduncular bracts 68.3(55.0–80.0) cm long; peduncles length not recorded; staminate rachillae
100, 6.3(3.5–8.0) cm long, arranged all around rachis, without or with few raphides; staminate monad/dyad bracteoles
usually ribbed, the same texture as the rachilla and continuous with it abaxially; staminate flowers 6.6(5.5–8.0) mm
long, in dyads, slightly sunken in rachillae, irregularly arranged on abaxial surface of rachillae; staminate petals linear,
joined at the bases, curved over at the apices, flat in cross-section, the surfaces smooth, coriaceous, not granular
tomentose, the margins smooth; stamens 6, neither exserted from nor much shorter than petals; anthers coiled and
twisted; pistillate rachillae 105, 2.8(2.0–3.8) cm long, arranged all around rachis; pistillate petals cuspidate, with
21.5(19.9–23.1) mm diameter, color not recorded, not elongate and angled, borne on short rachillae with 1-several
fruits per rachilla; endocarp surfaces smooth or with few, thin fibers, proximally with clear outlines of three locule
divisions, these marked by irregular, small pits, smooth, not ridged, the bases regularly rounded; endocarps in cross-
section without or with few fibers, 1-seeded, the abortive locules, when present, visible in endocarp cross-section but
not appressed to fertile locules; opercula superficial or slightly sunken in endocarps, not obscured by fibers; seeds 1,
cross-section not recorded.
Distribution and habitat:—Colombia (Vaupés), Venezuela (Amazonas), and Brazil (Amazonas) at 200(120–
400) m in open, sandy areas (Fig. 19). Galeano & Bernal (2010) reported that this species also occurs in the Colombian
department of Guainía but no specimens from there have been seen in the present study.
Taxonomic notes:—Specimens determined as preliminary species Orbignya luetzelburgii are polymorphic for
distalmost few pinnae and this is treated as a trait. Otherwise, specimens share a unique combination of qualitative
variable states and are therefore recognized as a phylogenetic species, A. luetzelburgii. The type specimen of
Parascheelea anchistropetala also shares this combination and is here included.
Attalea luetzelburgii is a taxonomically isolated species, formerly included in its own genus, Parascheelea.
Morphologically it does not appear to have any close relatives, although it is similar to similarly isolated species
(A. colenda, A. crassispatha, A. guianensis, A. maripa) in having superficial or slightly sunken opercula that are not
obscured by fibers. Specimens come from two areas but this may be an artifact of insufficient collecting.
Glassman (1999) placed Attalea luetzelburgii in Orbignya and Pinheiro (1997) also placed it with species formerly
included in Orbignya. Freitas et al. (2016) placed A. luetzelburgii it in their Orbignya-like clade, along with A. speciosa.
This species was not included in the study of Meerow et al. (2009).
20. Attalea maripa (Aublet) Martius (1844: 123). Palma maripa Aublet (1775: 974). Scheelea maripa (Aublet)
Wendland in Kerchove de Denterghem (1878: 256). Maximiliana maripa (Aublet) Drude (1881: 452). Englerophoenix
maripa (Aublet) Kuntze (1891: 728). Type:—FRENCH GUIANA. Without locality, no date, J. Aublet s.n. (holotype
P n.v.)
Maximiliana regia Martius (1826: 132). Maximiliana martiana Karsten (1856: 273). Englerophoenix regia (Martius) Kuntze (1891: 728).
Attalea regia (Martius) Wessels Boer (1965: 150). Lectotype (designated by Glassman 1978):—BRAZIL. Provinciae Paraensis et
Maragnaniensis: without locality, no date, C. Martius s.n. (holotype M n.v., M image!).
Maximiliana elegans Karsten (1856: 271). Neotype (designated by Glassman 1999):—COLOMBIA. Nova Grenada, Villavicencio, prov.
Bogotá, 1851–1857, J. Triana 734 (neotype P!).
Maximiliana caribaea Grisebach & Wendland ex Grisebach (1864: 522). Englerophoenix caribaeum (Grisebach & Wendland) Kuntze
(1891: 728). Type:—TRINIDAD AND TOBAGO. No locality, no date, H. Crueger s.n. (holotype K, destroyed).
Maximiliana longirostrata Barbosa Rodrigues (1891b: 112). Englerophoenix longirostrata (Barbosa Rodrigues) Barbosa Rodrigues
(1903a: 77). Lectotype (here designated):—Barbosa Rodrigues 1903a: t. 60B.

Maximiliana macrogyne Burret (1929a: 692). Type:—BRAZIL. Maranhão: Tury-assu, 19 October 1923, E. Snethlage 279 (holotype B,
destroyed).
Maximiliana stenocarpa Burret (1929a: 696). Lectotype (here designated):—PERU. Loreto: Iquitos, no date, G. Tessmann 5081 (lectotype
NY!) (the holotype at B was destroyed).
Maximiliana macropetala Burret (1929a: 699). Attalea macropetala (Burret) Wessels Boer (1965: 155). Type:—VENEZUELA. State
unknown: Rosalia, no date, S. Passarge 63 (holotype B, destroyed).
Plants 15.7(5.0–25.0) m tall. Stems 11.8(1.0–25.0) m long, 28.2(20.0–45.0) cm diameter, caulescent. Leaves 16(8–
25) per stem, arranged in five vertical ranks; sheaths 91.8(29.0–160.0) cm long; petioles 229.2(120.0–400.0) cm long;
rachises 604.2(448.0–960.0) cm long; pinnae 217(57–318) per side of rachis, arranged in distant clusters, without
auricles at the bases; middle pinnae 119.3(92.0–160.0) cm long, 4.5(3.3–6.6) cm wide; distalmost few pinnae free
from one another. Inflorescence peduncular bracts 155.5(90.0–250.0) cm long; peduncles 71.0(17.0–100.0) cm long;
staminate rachillae 635(245–1,000), 15.8(9.0–37.0) cm long, arranged all around rachis, with abundant raphides;
from it by an abaxial, proximal margin; staminate flowers 9.5(6.0–13.3) mm long, in monads or dyads, superficial
on rachillae, loosely and spirally or somewhat irregularly arranged all around rachillae (sometimes fewer or absent
from adaxial, proximal surface of rachillae); staminate petals linear, free, not curved at the apices, concave in cross-
section, the surfaces ribbed, not coriaceous, the margins smooth; stamens 6, exserted from petals; anthers straight;
pistillate rachillae 480(476–483), 15.2(6.2–21.0) cm long, arranged all around rachis; pistillate petals cuspidate, with
24.0(18.5–30.7) mm diameter, brown, pale brown, chestnut, not elongate and angled, borne on short to long rachillae
with 1-several fruits per rachilla; endocarp surfaces smooth or with few, thin fibers, the surfaces proximally with
clear outlines of three locule divisions, these marked by irregular, small pits, smooth, not ridged, the bases regularly
rounded, several-seeded, the abortive locules, when present, visible in endocarp cross-section but not appressed
to fertile locules; endocarps in cross-section without or with few fibers; opercula superficial or slightly sunken in
endocarps, not obscured by fibers; seeds 2, solid in cross-section, rarely with an irregular central cavity.
Distribution and habitat:—Widespread in the Amazon region and adjacent areas in Colombia, Venezuela,
Guyana, Surinam, French Guiana, Trinidad, Brazil, Ecuador, Peru, and Bolivia at 181(5–350) m in lowland rainforest
and persisting in disturbed areas (Fig. 19).
Taxonomic notes:—Specimens determined as preliminary species Maximiliana maripa share a unique combination
of qualitative variable states and are therefore recognized as a phylogenetic species, Attalea maripa (Fig. 23). The type
specimens of M. caribaea, M. elegans, M. longirostrata, M. macrogyne, M. macropetala, M. regia, and M. stenocarpa
also share this combination and are included as synonyms.
Attalea maripa is unique in the genus in its leaves that are arranged in five vertical ranks. Morphologically, it does
not appear to have any close relatives, although it is similar to similarly isolated species (A. colenda, A. crassispatha,
A. guianensis, A. luetzelburgii) in having superficial or slightly sunken opercula that are not obscured by fibers.
Pinheiro (1997) placed Attalea maripa as sister to the rest of the species of Attalea. Freitas et al. (2016) placed it
in their Orbignya-like clade, in a sub-clade with A. barreirensis and A. moorei, although morphologically it has little
in common with these two species. It was not included in the study of Meerow et al. (2009).
Subspecific variation:—Attalea maripa is a variable species, especially in terms of staminate rachillae size,
flower arrangement, and staminate flowers (Wessels Boer 1965). Inflorescences are not always unisexual and are
commonly observed with both staminate and pistillate flowers.
Bondar (1957) described a genus, Markleya, which he and Glassman (1999) considered to be an intergeneric
hybrid between Attalea maripa (as Maximiliana maripa) and A. speciosa (as Orbignya martiana). Wessels Boer
(1965) did not agree with this, although Pinheiro (1997) supported its hybrid status. It occurs in Surinam and Brazil
(Maranhão) (Fig. 19, open circles):
Markleya dahlgreniana Bondar (1957: 50). Attalea dahlgreniana (Bondar) Wessels Boer (1965: 158). ×
Maximbignya dahlgreniana (Bondar) Glassman (1999: 199). Type:—BRAZIL. Pará: Mun. Bragança, Tracuateua, 24
October 1956, G. Bondar 95829 (holotype RB n.v., RB image!).
Wessels Boer (1988) described a species, the type of which Glassman (1999) considered to be a hybrid between
Attalea maripa (as Maximiliana maripa) and an unknown species:
Attalea cryptanthera Wessels Boer (1988: 310). Type:—VENEZUELA. Amazonas: near San Carlos de Río
Negro, approx. 1°55’N, 67°W, 125 m, 22 January 1968, J. Wessels Boer 2323 (holotype, U n.v., isotypes F!, NY!).

FIGURE 22. Distribution maps of Attalea nucifera, A. oleifera, A. phalerata, and A. pindobassu.
21. Attalea nucifera Karsten (1856: 255). Lectotype (designated by Glassman 1977a):—Karsten 1861: t. 68. Epitype
(designated here):—COLOMBIA. Santander: carretera entre Barrancabermeja y Bucaramanga, ca. 1 hora antes de
Bucaramanga, 200 m, 26 February 1987, G. Galeano & A. Barfod 1276 (epitype COL!, isoepitypes AAU!, NY!)
Plants 3.0 m tall. Stems 0.3 m long, 31.8 cm diameter, acaulescent. Leaves 12(11–13) per stem; sheaths length not
recorded; petioles 95.0(60.0–130.0) cm long; rachises 360.3(341.0–390.0) cm long; pinnae 100(94–106) per side of
rachis, regularly arranged, without auricles at the bases; middle pinnae 70.8(65.0–76.5) cm long, 3.3(2.7–4.3) cm
wide; distalmost few pinnae remaining joined at their apices, sometimes with ‘windows’ between them or free from
one another. Inflorescence peduncular bracts not recorded; peduncles 25.0 cm long; staminate rachillae number not
recorded, 5.5(4.5–6.5) cm long, absent from adaxial surface of rachis, without or with few raphides; staminate monad/
dyad bracteoles not ribbed, crustaceous, different in texture from the rachilla and appearing separate from it by an
abaxial, proximal margin; staminate flowers 13.6(12.0–15.2) mm long, mostly in monads, rarely dyads, superficial on
rachillae, alternately arranged along abaxial surface of rachillae; staminate petals lanceolate, acuminate, flattened, free,
not or only slightly curved at the apices, flat in cross-section, the surfaces smooth, coriaceous, the margins minutely

serrate; stamens 6, neither exserted from nor much shorter than petals; anthers straight; pistillate rachillae number not
recorded, 0.4(0.2–0.5) cm long, arranged all around rachis; pistillate petals cuspidate, with elaborate, lobed margins;
staminodial rings well-developed, cupular with entire margins; fruits 67.7(66.1–71.1) mm long, 45.0(40.3–50.4) mm
diameter, brown or light brown, not elongate and angled, borne on short rachillae with 1 fruit per rachilla; endocarp
surfaces uneven, longitudinally ridged, with stout, rounded, adherent fibers, without small pits proximally, the bases
irregularly pointed, the endocarp becoming thinner and flanged; endocarps in cross-section with abundant, scattered
fibers, endocarps several-seeded, the abortive locules, when present, visible in endocarp cross-section but not appressed
to fertile locules; opercula sunken in endocarps and obscured by persistent fibers; seeds 2–3, solid in cross-section,
rarely with an irregular central cavity.
FIGURE 23. Attalea maripa, left, caulescent habit; right staminate inflorescence, with leaves arranged in a few, distinctive spirals (Acre,
Brazil).
Distribution and habitat:—Colombia (Santander) in the Magdalena valley at 227(200–300) m in lowland

rainforest and persisting in disturbed vegetation (Fig. 22). It also occurs in the Colombian departments of Antioquia,
Bolívar, and Cundinamarca (Galeano & Bernal 2010, Prada-Ríos & García 2019) but no specimens from there have
been seen in the present study.
Taxonomic notes:—Specimens determined as preliminary species Attalea nucifera are polymorphic for distalmost
few pinnae and this is treated as a trait. Otherwise, specimens share a unique combination of qualitative variable states
and are therefore recognized as a phylogenetic species, A. nucifera. Morphologically, it is similar to both A. allenii and
A. barreirensis, all three having cuspidate pistillate petals with elaborate, lobed margins.
Pinheiro (1997) considered that A. nucifera exhibited unusual anatomical characters, more similar to those of
Scheelea. He called this species “the most intriguing species included in this study” and considered that, anatomically,
it was a Scheelea. It is of interest to note that Galeano & Bernal (2010) considered that hybrids existed in Colombia
between A. nucifera and A. butyracea, and this may explain Pinheiro’s results. However, these hybrids were said to
occur in El Banco, Magdalena, further to the north of the known range of A. nucifera. Freitas et al. (2016) placed it as
sister species to A. allenii, in their Scheelea-like clade. It was not included in the study of Meerow et al. (2009).

22. Attalea oleifera Barbosa Rodrigues (1881: 123). Lectotype (here designated):—Barbosa Rodrigues 1903a: t. 58
Attalea concentrista Bondar (1942: 30). Lectotype (designated by Glassman 1977a):—BRAZIL. Bahia: Mun. S. Antonia de Jesus, Castro
Alves, Amargosa, Aeria, S. Ines, 200–400 m, no date, G. Bondar s.n. (holotype F!).
Attalea burretiana Bondar (1942: 30). Neotype (designated by Glassman 1977a):—BRAZIL. Bahia: Aratú, no date, G. Bondar 24
(holotype F!).
Plants 10.0 m tall. Stems 7.7(1.5–15.0) m long, 39.4(30.0–50.0) cm diameter, caulescent. Leaves 16(10–22) per
stem; sheaths length not recorded; petioles 3.1(0.0–4.4) cm long, short or absent; rachises 660.0(503.0–880.0) cm
long; pinnae 169(137–191) per side of rachis, regularly arranged, without auricles at the bases, abaxial and proximal
part of mid-vein with or without dense patches of brown hairs; middle pinnae 98.9(66.0–117.0) cm long, 3.9(3.0–
4.8) cm wide; distalmost few pinnae free from one another. Inflorescence peduncular bracts 135.5(59.0–187.0) cm
long; peduncles 47.5(20.0–80.0) cm long; staminate rachillae 256(142–300), 14.7(11.0–20.0) cm long, arranged all
around rachis, without or with few raphides; staminate monad/dyad bracteoles usually ribbed, the same texture as
the rachilla and continuous with it abaxially; staminate flowers 16.2(13.4–18.9) mm long, mostly in monads, rarely
dyads, superficial on rachillae, alternately arranged along abaxial surface of rachillae; staminate petals lanceolate,
acuminate, flattened, free, not or only slightly curved at the apices, flat in cross-section, the surfaces ribbed, not
coriaceous, not granular tomentose, the margins minutely denticulate; stamens 7(6–8), neither exserted from nor much
shorter than petals; anthers straight; pistillate rachillae number not recorded, 13.4(9.0–21.5) cm long, arranged all
around rachis; pistillate petals cuspidate, with irregular margins; staminodial rings well-developed, cupular, with entire
margins; fruits 79.9(66.3–97.7) mm long, 48.3(40.0–65.6) mm diameter, orange-brown, not elongate and angled,
borne on short to long rachillae with 1-several fruits per rachilla; endocarp surfaces uneven, longitudinally ridged, with
thinner and flanged; endocarps in cross-section with abundant, scattered fibers, 1-several-seeded, the abortive locules,
FIGURE 24. Attalea oleifera, left, caulescent habit; right, infructescence (Espírito Santo, Brazil).

Distribution and habitat:—Atlantic Coastal Forest of Brazil (Bahia) with an outlier in Paraíba, at 343(75–775)
m in lowland rainforest or transitional forest between coastal rainforest and dryer, inland areas (Fig. 22). Noblick
(1991) reported that Attalea oleifera (as A. burretiana) also occurred in Sergipe, just to the north of Bahia, and Espírito
Santo, just to the south, although no specimens from there have been seen. He also noted that A. oleifera occurred
further north in Alagoas, Paraíba, and Pernambuco. However, only one specimen (Lima 165) from one of these three
states (Paraíba) has been seen.
Taxonomic notes:—Specimens determined as preliminary species Attalea burretiana are polymorphic for abaxial
mid-vein surfaces and stamen number. Two specimens (Bondar s.n., Noblick 3733) have staminate flowers with 6
stamens, all others have 7–8 stamens. Abaxial mid-vein surfaces and stamen number are treated as traits. Otherwise,
specimens share a unique combination of qualitative variable states and are recognized as a phylogenetic species, A.
oleifera (Fig. 24). The type specimens of Attalea concentrista and A. burretiana also shares this combination and
are included (see also Noblick 1991). Attalea oleifera is similar to other species (A. compta, A. dubia, A. exigua, A.
humilis, A. pindobassu) having pinnae with the abaxial and proximal part of mid-vein with dense patches of brown
hairs, and staminate petals with minutely denticulate margins.
Pinheiro (1997) placed Attalea oleifera in Attalea s.s. Meerow et al. (2009) placed it in their Attalea group, and
Freitas et al. (2016) included it in their Attalea-like clade.
23. Attalea phalerata Martius ex Sprengel (1825: 624). Scheelea phalerata (Martius ex Sprengel) Burret (1929a:
541). Neotype (designated by Glassman 1977b):—Martius 1845: t. 169. Epitype (here designated):—BRAZIL. Goiás:
Municipio de Goiânia, 14 km NE of the city on BR-060/BR153 to Anapolis, 16°40’S 49°16’W, 700–800 m, 7 May
1992, L. Noblick & H. Ferreira 4864 (epitype UFG n.v., isoepitypes FTG n.v., NY!)
Attalea princeps Martius (1844: 113). Scheelea princeps (Martius) Karsten (1856: 269). Type:—BOLIVIA. Santa Cruz: Moxos and
Chiquitos, no date, A. d’Orbigny 16 (holotype P, destroyed). Lectotype (designated by Glassman 1977b):—Martius 1844: t. 31B.
Attalea gomphococca Martius (1845: 301). Scheelea gomphococca (Martius) Burret (1929a: 541). Lectotype (here designated):—Martius
1845, t. 167, fig. 6.
Maximiliana tetrasticha Drude (1881: 455). Englerophoenix tetrasticha (Drude) Barbosa Rodrigues (1903a: 76). Scheelea tetrasticha
(Drude) Burret (1929a: 667). Type:—BRAZIL. Pará: Rio Tocantins and Araguaia, August–September 1844, M. Weddell 2331
(holotype P n.v., isotype F!).
Scheelea amylacea Barbosa Rodrigues (1891a: 17). Attalea amylacea (Barbosa Rodrigues) Zona (2002: 132). Lectotype (here
designated):—Barbosa Rodrigues 1891a: ts. 5A, 6.
Scheelea anizitziana Barbosa Rodrigues (1898b: 63). Attalea anisitsiana (Barbosa Rodrigues) Zona (2002: 132). Type:—BRAZIL. Matto
Grosso: no locality, no date, Anon 223 (holotype destroyed). Lectotype (here designated):—Barbosa Rodrigues 1898b: t. 20.
Scheelea princeps var. corumbaensis Barbosa Rodrigues (1898b: 66). Scheelea corumbaensis (Barbosa Rodrigues) Barbosa Rodrigues
(1903a: 54). Type:—BRAZIL. Matto Grosso: Corumbá, no date, J. Barbosa Rodrigues 218 (holotype destroyed). Lectotype (here
designated):—Barbosa Rodrigues 1898b: t. 21A.
Scheelea quadrisperma Barbosa Rodrigues (1899: 23). Lectotype (designated by Glassman, 1972):—Barbosa Rodrigues 1899: t. 6.
Attalea parviflora Barbosa Rodrigues (1903b: 625). Scheelea parviflora (Barbosa Rodrigues) Barbosa Rodrigues (1903a: 53). Type:—
PARAGUAY. Concepción: prope Concepción, August 1901, E. Hassler 7165 (holotype G n.v., G image!).
Scheelea quadrisulcata Barbosa Rodrigues (1907: 107). Lectotype (designated by Glassman 1977b):—Barbosa Rodrigues 1907: t. 22B.
Scheelea lauromuelleriana Barbosa Rodrigues (1907: 108). Attalea lauromuelleriana (Barbosa Rodrigues) Zona (2002: 133). Lectotype
(here designated):—Barbosa Rodrigues 1907: t. 25.
Attalea hoehnei Burret (1929a: 522). Type:—BRAZIL. Mato Grosso or Acre: Agua Limpa, no date, F. Hoehne 2196 (holotype B
destroyed).
Scheelea huebneri Burret (1929a: 663). Type:—BRAZIL. Amazonas: Rio Purus, no date, G. Huebner 23a (holotype B n.v., B image!).
Scheelea weberbaueri Burret (1929a: 659). Attalea weberbaueri (Burret) Zona (2002: 133). Type:—PERU. Junín: Tarma, La Merced,
Chanchamayo, 1,000 m, December 1902, A. Weberbauer 1848 (holotype B n.v., B image!).
Scheelea microspadix Burret (1940: 104). Type:—BRAZIL. Mato Grosso: without locality, no date, W. Hopp 3010 (holotype B n.v., B
image!).
Attalea pacensis Moraes & Pintaud (2016b: 164). Type:—BOLIVIA.. La Paz: Prov. Abel Iturralde, San Buenaventura, 19 November
2015, M. Moraes & T. Cartagena MMR2531 (holotype LPB n.v., isotype K n.v.).
Leaves 19(11–27) per stem; sheaths 60.0(50.0–70.0) cm long; petioles 26.4(0.0–90.0) cm long, short or absent or
well-developed; rachises 469.7(340.0–619.0) cm long; pinnae 169(104–205) per side of rachis, irregularly arranged,
usually spreading in different planes, with auricles at the bases; middle pinnae 92.1(65.5–112.0) cm long, 3.1(2.1–4.9)
cm wide; distalmost few pinnae free from one another. Inflorescence peduncular bracts 109.7(95.0–135.0) cm long;
peduncles 56.3(15.0–120.0) cm long; staminate rachillae 246(110–348), 9.0(6.0–15.0) cm long, arranged all around
rachis, with abundant raphides; staminate monad/dyad bracteoles usually ribbed, the same texture as the rachilla and
continuous with it abaxially; staminate flowers 7.4(6.0–9.4) mm long, in monads, superficial on rachillae, loosely and
irregularly arranged and tending to be absent from adaxial surface of rachillae; staminate petals linear, free, not curved
at the apices, flat in cross-section, the surfaces ribbed, not coriaceous, not granular tomentose, the margins smooth;
stamens 6, much shorter than petals; anthers straight; pistillate rachillae 63(50–75), 4.8(2.0–7.7) cm long, arranged all
around rachis; pistillate petals cuspidate, with irregular margins; staminodial rings well-developed, cupular, with entire
margins; fruits 61.2(48.4–79.1) mm long, 32.2(24.2–47.4) mm diameter, brown, light brown, yellow-green, or red, not
elongate and angled, borne on short to long rachillae with 1-several fruits per rachilla; endocarp surfaces with stout,
rounded, adherent fibers, without small pits proximally, the bases irregularly pointed, the endocarp becoming thinner
and flanged; endocarps in cross-section with abundant fibers in regular clusters, several-seeded, the abortive locules,
Distribution and habitat:—Brazil (Acre, Goiás, Mato Grosso, Mato Grosso do Sul, Maranhão, Pará, São
Paulo, Tocantins), Peru (Junín, Madre de Dios), Bolivia (Beni, La Paz, Pando, Santa Cruz), and Paraguay (Amambay,
Concepción, Presidente Hayes) at 305(6–750) m in seasonal forest or, less often, in lowland rainforest, and persisting
in disturbed areas (Fig. 22).
FIGURE 25. Attalea phalerata, left, acaulescent habit with irregularly arranged pinnae; right, infructescence (Rondônia, Brazil).
Taxonomic notes:—Specimens determined as preliminary species Scheelea princeps are polymorphic for stems
and petioles and these variables are treated as traits. Petioles are scored as either short or absent or well-developed.
However, it is not clear if a true petiole is present or not; the evidence suggests it is probably absent. One specimen
(Noblick 5018) label states “on adult trees the true leaf petiole is practically none”. Otherwise, specimens share a
unique combination of qualitative variable states and are recognized as a phylogenetic species, Attalea phalerata (Fig.

25). Specimens determined as preliminary species S. phalerata, and the type specimens of S. amylacea, S. anizitziana,
S. gomphococca, A. hoehnei, S. huebneri, S. lauromuelleriana, S. microspadix, S. parviflora, S. princeps, S. princeps
var. corumbaensis, A. pacensis, S. quadrisperma, S. quadrisulcata, S. tetrasticha, and S. weberbaueri also share this
combination and are included. Because the neotype of Attalea phalerata is a plate, an epitype is here designated.
Glassman (1999) recognized two groups of species: Scheelea anisitsiana, S. weberbaueri, S. phalerata, and S.
amylacea with “staminate flowers arranged on one side of the rachillae”; and S. princeps and S. cephalotes with
“staminate flowers spirally arranged around staminate rachillae”. Scheelea anisitsiana and S. amylacea are treated as
synonyms, and S. cephalotes as a separate species (Attalea cephalotus). As for Scheelea princeps, Glassman (1999)
considered it to have spirally arranged staminate flowers although two of the specimens (Balick 1358, Hopkins 173)
he cited have the staminate flowers absent from adaxial surface of rachillae, as in all other specimens here included
in A. phalerata. Pintaud et al. (2016) also considered S. princeps to have spirally arranged flowers (although in their
key it was said to have unilaterally arranged flowers). Rodríguez del Castillo et al. (2016) considered it to have “a
mostly spiral arrangement”. In fact, as stated by Martius (1844), the staminate flowers of S. princeps are absent from
the adaxial surface of the rachillae (“Postice hinc florigeris inde nudis, antice undique florigeris”), and S. princeps is
clearly a synonym of A.phalerata.
Scheelea lauromuelleriana was placed by Pintaud (2008) in the “Attalea macrolepis complex” along with names
here considered to be synonyms of A. butyracea. However, by its fruits with clustered endocarp fibers, it appears to
represent A. phalerata.
Scheelea huebneri was mistakenly included as a synonym of A. butyracea by Henderson (1995) and Henderson
et al. (1995). Although the type is sterile it apparently belongs under A. phalerata. Pintaud (2008) considered it had
spirally arranged staminate flowers, whereas Pintaud et al. (2016) considered it had the staminate flowers arranged on
one side of the rachillae. Rodríguez del Castillo et al. (2016) illustrated an endocarp with clustered fibers.
The type of Scheelea weberbaueri is tentatively included here; the endocarps in cross-section have scarcely
clustered fibers, unlike other specimens. Pintaud et al. (2016) distinguished it by its pinnae being more than 5 cm wide.
The type is reported to have 3 cm wide pinnae and the only specimen (Killip 25141) cited by Glassman (1999) has
pinnae slightly less than 5 cm wide.
The type of Scheelea microspadix has a remarkably small infructescence, not unlike another specimen (Prance
59372) from Mato Grosso.
Attalea phalerata appears most similar to four other species (A. butyracea A. cephalotus, A. insignis, A. plowmanii)
all previously included in Scheelea, having pinnae with auricles, linear staminate petals, and stamens much shorter
than the petals. Specimens occur in a wide arc around the western, southern, and eastern margins of the Amazon
region, but there appears to be no geographic disjunction.
Pinheiro (1997), based on leaf anatomy, considered that Attalea phalerata could be a hybrid between unspecified
species of Attalea s.s. and Scheelea. However, Pinheiro’s specimens came from two different species—A. phalerata
and A. cephalotus. Meerow et al. (2009) placed it in their Scheelea group and Freitas et al. (2016) included it in their
Scheelea-like clade.
24. Attalea pindobassu Bondar (1942: 462). Neotype (designated by Glassman 1977a):—BRAZIL. Bahia: Serra de
Ouro, 1940, G. Bondar s.n. (holotype F n.v.)
Attalea seabrensis Glassman (1999: 50). Type:—BRAZIL. Bahia: Mun. Seabra, 291 km W of Salvador, Veredinha, 21 August 1976, S.
Glassman 13034 (holotype F!, isotypes SP n.v., US!).
Plants 20.0 m tall. Stems 9.1(3.0–20.0) m long, 38.0(30.0–50.0) cm diameter, caulescent. Leaves 18(8–22) per stem;
sheaths 150 cm long; petioles 106.3(10.0–175.0) cm long; rachises 560.5(417.0–679.0) cm long; pinnae 173(149–183)
per side of rachis, regularly or irregularly arranged, without auricles at the bases, abaxial and proximal part of mid-vein
with dense patches of brown hairs; middle pinnae 91.5(77.0–106.0) cm long, 3.7(3.0–4.8) cm wide; distalmost few
pinnae free from one another. Inflorescence peduncular bracts 143.2(105.0–187.0) cm long; peduncles 51.9(30.0–
90.0) cm long; staminate rachillae 312(261–401), 13.3(9.0–17.0) cm long, arranged all around rachis, without or with
few raphides; staminate monad/dyad bracteoles usually ribbed, the same texture as the rachilla and continuous with
it abaxially; staminate flowers 15.0(11.2–19.0) mm long, mostly in monads, rarely dyads, superficial on rachillae,
alternately arranged along abaxial surface of rachillae; staminate petals lanceolate, acuminate, flattened, free, not or
only slightly curved at the apices, flat in cross-section, the surfaces ribbed, not coriaceous, not granular tomentose,
the margins minutely denticulate; stamens 10(8–12), neither exserted from nor much shorter than petals; anthers
straight; pistillate rachillae number not recorded, 11.9(8.0–14.0) cm long, arranged all around rachis; pistillate petals
cuspidate, with irregular margins; staminodial rings well-developed, cupular, with entire margins; fruits 78.3(64.3–
91.8) mm long, 50.9(36.8–62.7) mm diameter, dark brown, not elongate and angled, borne on short to long rachillae
cross-section with abundant, scattered fibers, 1-several-seeded, the abortive locules, when present, visible in endocarp
3(1–5), solid in cross-section, rarely with an irregular central cavity.
Distribution and habitat:—Cerrado region of Brazil (Bahia) on the western and eastern margins of the Serra do
Tombador and in the Serra do Sincorá at 738(375–1,060) m in wet areas along slopes (Fig. 22).
Taxonomic notes:—Specimens determined as preliminary species Attalea pindobassu share a unique combination
of qualitative variable states and are recognized as a phylogenetic species, A. pindobassu. Specimens determined as
preliminary species A. seabrensis share the same combination, except for pinnae arrangement. Noblick (1991) separated
A. seabrensis from A. pindobassu based on pinnae arrangement. He considered A. seabrensis to have “clustered leaflets
on the lower one third to one-half of the leaf”, whereas A. pindobassu was said to have pinnae “almost regularly inserted
along the rachis (except at the very base)”. Based on this, pinnae arrangement is treated as a trait, and specimens of A.
seabrensis are included in A. pindobassu. The population of Attalea seabrensis is somewhat isolated from that of A.
pindobassu, on the western side of the Serra do Tombador, and specimens differ significantly from A. pindobassu in
three variables (rachis length, staminate rachillae length, staminate flower length)(t-test, P <0.05) with A. seabrensis
having lower values for these three variables.
Attalea pindobassu is distinguished from A. oleifera by its well-developed petioles. Although petioles have been
shown to be problematic in Attalea taxonomy, as discussed in the Morphology section, in this particular case we have
reliable data on petioles due to the intensive field work and numerous specimens of Larry Noblick in Bahia (Noblick
1991).
Pinheiro (1997) placed Attalea pindobassu with other Attalea species (see notes under A. compta). Meerow et al.
(2009) placed it in the Attalea group and Freitas et al. (2016) placed it in their Attalea-like clade.
25. Attalea plowmanii (Glassman) Zona (2002: 133). Scheelea plowmanii Glassman (1999: 144). Type:—PERU.
Loreto: Maynas, Río Yaguasyacu, Brillo Nuevo, 18 April 1977, T. Plowman 6778 (holotype BH!, isotypes GH n.v.,
USM n.v.)
Plants 4.7(4.0–6.0) m tall. Stems 1.0 m long, diameter not recorded, caulescent or acaulescent. Leaves 12(5–16) per
stem; sheaths 54.0(34.0–74.0) cm long; petioles 131.6(70.0–185.0) cm long; rachises 404.8(298.0–600.0) cm long;
pinnae 104(86–136) per side of rachis, regularly arranged, with auricles at the bases; abaxial and proximal part of
mid-vein with or without dense patches of brown hairs; middle pinnae 90.2(74.5–129.0) cm long, 4.0(2.7–5.5) cm
wide; distalmost few pinnae remaining joined at their apices, sometimes with ‘windows’ between them. Inflorescence
peduncular bracts 114.7(100.0–123.0) cm long; peduncles 35.0 cm long; staminate rachillae 20, 13.3(10.0–16.5) cm
long, arranged all around rachis, with abundant raphides; staminate monad/dyad bracteoles usually ribbed, the same
texture as the rachilla and continuous with it abaxially; staminate flowers 12.6(9.6–16.0) mm long, in monads or
dyads, superficial on rachillae, loosely and spirally or somewhat irregularly arranged all around rachillae (sometimes
fewer or absent from adaxial, proximal surface of rachillae); staminate petals linear, free, not curved at the apices, flat
in cross-section, the surfaces ribbed, not coriaceous, not granular tomentose, the margins smooth; stamens 6, much
shorter than petals; anthers straight; pistillate rachillae 20, 1.4(0.7–3.0) cm long, arranged all around rachis; pistillate
petals not cuspidate, with irregular margins; staminodial rings well-developed, cupular, with entire margins; fruits
79.5(71.1–96.7) mm long, 47.0(37.9–52.4) mm diameter, brown, not elongate and angled, borne on short rachillae
with 1 fruit per rachilla; endocarp surfaces uneven, longitudinally ridged, with stout, rounded, adherent fibers, the
surfaces without small pits proximally, the bases irregularly pointed, the endocarp becoming thinner and flanged;
endocarps in cross-section with abundant, scattered fibers, several-seeded, the abortive locules, when present, visible
fibers; seeds 3, solid in cross-section, rarely with an irregular central cavity.
Distribution and habitat:—Colombia (Amazonas) and Peru (Loreto) at 148(109–250) m in lowland rainforest
(Fig. 27).

FIGURE 26. Attalea plowmanii, infructescence (Amazonas, Colombia).
Taxonomic notes:—Specimens determined as preliminary species Scheelea plowmanii are polymorphic for
abaxial mid-vein surfaces and this variable is treated as a trait. Otherwise specimens share a unique combination of
qualitative variable states and are recognized as a phylogenetic species, A. plowmanii (Fig. 26). This appears most
similar to four other species (A. butyracea A. cephalotus, A. insignis, A. phalerata) all previously included in Scheelea,
having pinnae with auricles, linear staminate petals, and stamens much shorter than the petals.
Glassman (1999) considered Attalea plowmanii closely related to Scheelea salazarii, here treated as a synonym
of A. butyracea. Freitas et al. (2016) placed it in their Scheelea-like clade where it was sister to A. insignis. It was not
included by Pinheiro (1997) nor by Meerow et al. (2009).
Subspecific variation:—Specimens are scored as having the pistillate rachillae arranged all around rachis,
although there is a tendency to one-sidedness in some specimens. Fruits tend to be longitudinally fissured. One
specimen (Moore 10214) has endocarps in cross-section with dense fibers arranged almost in a ring around the seeds,
unlike other specimens and more like the endocarps of the sympatric Attalea racemosa. It is here scored as having
scattered fibers.
26. Attalea racemosa Spruce (1869: 166). Orbignya racemosa (Spruce) Drude (1881: 448). Type:—VENEZUELA.
Amazonas: in sylvis inter fluvios Negro et Guasíe, October 1854, R. Spruce 54 (holotype K!, isotype P!)
Attalea ferruginea Burret (1934: 1044). Type:—VENEZUELA. Amazonas: San Carlos de Río Negro, March 1933, C. Lakó/G. Huebner
166 (holotype B n.v., B image!).
Plants 3.5 m high. Stems length and diameter not recorded, acaulescent. Leaves 12(6–35) per stem; sheaths 61.8(40.0–
75.0) cm long; petioles 152.6(82.0–240.0) cm long; rachises 386.3(210.0–525.0) cm long; pinnae 103(68–130) per side
of rachis, regularly arranged, with or without auricles at the bases; middle pinnae 71.2(51.0–97.0) cm long, 3.4(2.5–
4.5) cm wide; distalmost few pinnae remaining joined at their apices, sometimes with ‘windows’ between them or free
from one another. Inflorescence peduncular bracts 95.0(65.0–140.0) cm long; peduncles 121.0(74.0–160.0) cm long;

staminate rachillae number not recorded, 8.7(6.0–13.5) cm long, absent from adaxial surface of rachis, without or with
few raphides; staminate monad/dyad bracteoles usually ribbed, the same texture as the rachilla and continuous with it
abaxially; staminate flowers 16.3(12.7–20.8) mm long, mostly in monads, superficial on rachillae, loosely and singly
arranged along abaxial surface of rachillae; staminate petals lanceolate, acuminate, flattened, free, not or only slightly
curved at the apices, flat in cross-section, the surfaces ribbed, not coriaceous, not granular tomentose, the margins
tomentose; stamens 32(21–40), neither exserted from nor much shorter than petals; anthers straight; pistillate rachillae
20, 2.6(1.5–4.0) cm long, absent from adaxial surface of rachis, at least proximally; pistillate petals cuspidate, with
43.0(29.6–51.1) mm diameter, brown or light brown, not elongate and angled, borne on short rachillae with 1 fruit
FIGURE 27. Distribution maps of Attalea plowmanii, A. racemosa, A. sagotii (closed circles = A. sagotii, open circles = unusual
specimens, see text for details), and A. septuagenata.

Distribution and habitat:—Amazon region in Colombia (Amazonas, Caquetá, Guainía, Guaviare), Venezuela
(Amazonas), Peru (Loreto), and Brazil (Amazonas) at 165(85–425) m in lowland forest or more open, white sand areas
(Fig. 27).
Taxonomic notes:—Specimens determined as preliminary species Attalea ferruginea are polymorphic for auricles
and distalmost pinnae and these variables are treated as traits. Otherwise, specimens share a unique combination of
qualitative variable states and are recognized as a phylogenetic species, using the earlier name A. racemosa (Fig. 28).
Glassman (1999) considered A. racemosa to be a doubtful species of Orbignya, based on a misinterpretation of Drude
(1881). However, the type clearly belong to Attalea s.s. Glassman (1999) tentatively included two specimens; Davidse
17025 (here placed in A. sagotii) and Renteria 1499 (here placed in A. nucifera).
FIGURE 28. Attalea racemosa, left, acaulescent habit; right, pistillate inflorescence (Amazonas, Venezuela, images by Hans Beck).
Attalea racemosa is similar to A. septuagenata, differing in its acaulescent habit and fruits that are not grayish-
white tomentose at the apex.
Pinheiro (1997), based on leaf anatomical data, considered that specimens of A. racemosa showed “strong conflict
in terms of their character variation and correlation” and were of possible hybrid origin. This may explain the confusing
variation seen in this species. Freitas et al. (2016) included A. racemosa (as A. ferruginea) in their Scheelea-like clade.
Morphologically this relationship seems unlikely. It was not included by Meerow et al. (2009).
Subspecific variation:—Attalea racemosa is a variable and poorly understood species. Two specimens (Schultes
18040, Wessels Boer 2322) from Venezuela have larger fruits than other Venezuelan ones, and have somewhat clustered
fibers. These fruits are similar to those of specimens from Peru.
27. Attalea sagotii (Trail ex Im Thurn) Wessels Boer (1965: 162). Orbignya sagotii Trail ex Im Thurn (1884: 276).
Lectotype (designated by Wessels Boer 1965):—FRENCH GUIANA. Karouany, 1855–1856, P. Sagot 831 (lectotype
K!, isolectotype P!)
Orbignya sabulosa Barbosa Rodrigues (1888: 54). Lectotype (here designated):—Barbosa Rodrigues 1903a: t. 48.
Attalea polysticha (Burret) Wessels Boer (1988: 301). Orbignya polysticha Burret (1932: 324). Type:—PERU. Loreto: Mishuyacu, near

Iquitos, October–November 1929, G. Klug 205 (holotype US!, isotypes F!, NY!).
Attalea pycnocarpa Wessels Boer (1988: 299). Type:—VENEZUELA. Amazonas: Puerto Ayacucho, 5°44’N 67°38’W, 28 July 1967, J.
Wessels Boer 1910 (holotype U n.v., U image!).
43.5(37.0–50.0) cm long; petioles 143.4(70.0–300.0) cm long; rachises 477.5(270.0–700.0) cm long; pinnae 93(67–
130) per side of rachis, regularly arranged, with or without auricles at the bases; middle pinnae 93.7(45.0–143.0)
cm long, 4.1(2.0–6.1) cm wide; distalmost few pinnae remaining joined at their apices, sometimes with ‘windows’
between them or free from one another. Inflorescence peduncular bracts 100.0 cm long; peduncles 43.3(20.0–70.0)
cm long; staminate rachillae 90(73–103), 8.2(4.0–11.5) cm long, arranged all around rachis, with abundant raphides;
from it by an abaxial, proximal margin; staminate flowers 7.2(5.2–9.8) mm long, mostly in monads, rarely dyads,
slightly sunken in rachillae, spirally arranged but tending to be absent from adaxial surface of rachillae, especially
proximally; staminate petals spathulate, free, curved over at the apices, flat in cross-section, the surfaces smooth,
coriaceous, not granular tomentose, the margins smooth; stamens 11(9–16), neither exserted from nor much shorter
than petals; anthers coiled and twisted; pistillate rachillae number not recorded, 5.6(3.5–8.0) cm long, arranged all
around rachis; pistillate petals cuspidate, with irregular margins; staminodial rings well-developed, cupular, with
entire margins; fruits 37.9(29.9–56.9) mm long, 23.5(20.1–31.2) mm diameter, brown, tawny brown, reddish-brown,
brownish-yellow, light yellow, or pale yellow, not elongate and angled, borne on short to long rachillae with 1-several
fruits per rachilla; endocarp surfaces uneven, longitudinally ridged, with stout, rounded, adherent fibers, without small
pits proximally, the bases regularly rounded; endocarps in cross-section without or with few fibers, 1-seeded, the
in endocarps and obscured by persistent fibers; seeds 1, solid in cross-section, or with an irregular central cavity.
Distribution and habitat:—Northern Amazon region of Colombia, Venezuela, the Guianas, Peru, and Brazil at
147(5–350) m in lowland rainforest or sometimes in more open, white sand areas (Fig. 27).
FIGURE 29. Attalea sagotii, left, acaulescent habit; right, infructescence (Amazonas, Brazil).

Taxonomic notes:—Specimens determined as preliminary species Orbignya sagotii are polymorphic for
petioles, auricles, and distalmost pinnae and these variables are treated as traits. Otherwise, specimens share a unique
combination of qualitative variable states and are recognized as a phylogenetic species, Attalea sagotii (Fig. 29).
Specimens determined as preliminary species Orbignya polysticha, and the type of Orbignya sabulosa, share the same
combination and are included. Attalea pycnocarpa is tentatively included here. The image of the type shows rachillae
and an endocarp and these resemble those of A. sagotii. However, the label reports the stem to be 12 m tall. The type
may represent a mixed collection. Attalea sagotii is similar to a group of species (A. cohune, A. cuatrecasana, A.
eichleri, A. speciosa) having coiled and twisted anthers.
Henderson (1995) and Henderson et al. (1995) used the name Attalea microcarpa Martius for this species.
Following Glassman (1999) this name is here treated as a Doubtful Name.
Freitas et al. (2016) included Attalea sagotii (as A. polysticha) in their Orbignya-like clade. It was not included by
Meerow et al. (2009).
Subspecific variation:—Glassman (1999) recognized two species here: Orbignya sagotii with staminate flowers
absent from the abaxial surface of the rachillae and O. polysticha with staminate flowers arranged spirally all round the
rachillae. This distinction is not as clear as he supposed, and here all specimens are scored as having staminate flowers
spirally arranged but tending to be absent from adaxial surface of rachillae, especially proximally. Most specimens
from the western part of the range tend to have less of the proximal, adaxial surface of the rachillae without flowers,
and most specimens from the eastern part of the range tend to have more of the proximal, adaxial surface of the
rachillae without flowers. However, some western specimens (e.g., Vásquez 7756) from Peru have the abaxial surface
of the rachillae almost completely without flowers, and a similar situation occurs in eastern specimens in French
Guiana (Pintaud 2008).
In eastern Brazil, north of the Amazon in Pará and Amapá, there is an area of more open, savanna type vegetation,
approximately from the Rio Trombetas in the west to the Rio Jari in the east. The specimens from here are confusing
(Fig. 27, open circles). Two specimens (Henderson 1741, 1742) from Monte Alegre are acaulescent with no petiole,
long, stiff leaf rachis, staminate flowers with three spathulate petals and 9–12 stamens, and rather small fruits. Another
specimen from the Rio Jari (Pires-O’Brien 1765) is similar (see Pires-O’Brien 1993, who described these Jari plants
as invasive weeds), and similar specimens have been observed on the Rio Nhamunda, slightly more to the west
(Henderson, pers. obs.). These differ from another specimen (Henderson 1736), also from Monte Alegre, which is
acaulescent or caulescent (Henderson, pers. obs.) with no petiole, shorter leaves with the rachis curved over near the
apex, staminate flowers with two, non-spathulate petals and 18 stamens, and larger fruits. All these specimens appear
similar to A. sagotii but differ in several respects, particularly the habitat, absent petiole, and staminate flowers.
Henderson (1995) and Henderson et al. (1995) used the name Attalea spectabilis Martius for these eastern Brazilian
specimens. This name is here treated as an Excluded Name because the type is destroyed (Glassman 1999), although
the type locality, Serra de Barú in Pará, is also within this savanna region. In the present study these specimens are not
recognized as a distinct species because they have the same character state combinations as A. sagotii. As noted by
Henderson (1995), some of these specimens may be hybrids.
Pinheiro (1997), using anatomical data, considered that specimens of three similar species, which he called O.
polysticha, O. sagotii, and O. spectabilis, were the most confusing of all species he studied, and exhibited characters
similar to those of the Scheelea group and of the Attalea group. However, specimens studied by Pinheiro of two of
these species represent a mixture of species; for O. spectabilis, Pardini 70 is here referred to as an unusual specimen
near A. barreirensis; Mori 17421 to A. sagotii; and Wessels Boer 1503 to Markleya dahlgreniana; and for O. sagotii,
Lobo 109 is also an unusual specimen near A. barreirensis (see notes under that species).
Two other specimens (de Granville 11186, 13027), not included in the above description, are tentatively included
here although the fruits are larger than usual.
28. Attalea septuagenata Dugand (1953: 3). Type:—COLOMBIA. Amazonas: Río Miriti-Paraná, Caño Guacayá, 2–8
March 1952, R. Schultes & Cabrera 15796 (holotype COL n.v., isotype BH!)
Plants height not recorded. Stems 9.5 m long, 27.5 cm diameter, caulescent. Leaves 19 per stem; sheaths length not
recorded; petioles length not recorded, well-developed; rachises 900 cm long; pinnae 183 per side of rachis, regularly
arranged, without auricles at the bases; middle pinnae 152.5 cm long, 4.5 cm wide; distalmost few pinnae free from
one another. Inflorescence peduncular bracts 109 cm long; peduncles 100 cm long; staminate rachillae number
not recorded, 16.5 cm long, absent from adaxial surface of rachis, without or with few raphides; staminate monad/
16.1(15.3–16.8) mm long, mostly in monads, superficial on rachillae, loosely and singly arranged along abaxial surface
cross-section, the surfaces ribbed, not coriaceous, not granular tomentose, the margins tomentose; stamens 55(39–70),
neither exserted from nor much shorter than petals; anthers straight; pistillate rachillae 51, 4.5 cm long, absent from
adaxial surface of rachis, at least proximally; pistillate petals cuspidate, with irregular margins; staminodial rings
well-developed, cupular, with entire margins; fruits 105.2 mm long, 55.1(54.9–55.3) mm diameter, grayish-brown,
grayish-white tomentose at the apices, not elongate and angled, borne on short to long rachillae with 1-several fruits
not appressed to fertile locules; opercula sunken in endocarps and obscured by persistent fibers; seeds 2, solid in cross-
section, rarely with an irregular central cavity.
Distribution and habitat:—Colombia (Amazonas) at 250 m in lowland rainforest (Fig. 27).
Taxonomic notes:—Specimens determined as preliminary species Attalea septuagenata share a unique
combination of qualitative variable states and are recognized as a phylogenetic species, A. septuagenata. This shares
all qualitative variable states with A. racemosa except for the caulescent habit and distinctive grayish-white tomentum
at the fruit apices (Fig. 30).
FIGURE 30. Attalea septuagenata, infructescence with fruits grayish-white tomentose at the apices (Amazonas, Colombia).
Attalea septuagenata was not included by Pinheiro (1997), Meerow et al. (2009), or by Freitas et al. (2016).
Subspecific variation:—There is a large difference in stamen number from the only two specimens examined.
Dugand (1953) reported 60–75 stamens from the type and here 64 and 70 stamens have been counted from two flowers
from the type. However, ten flowers from Galeano 2078 had 34(32–39) stamens (G. Galeano, pers. comm.). This latter
range is much nearer to that of A. racemosa and emphasizes the similarity between the two species.

29. Attalea speciosa Martius (1826: 138). Orbignya martiana Barbosa Rodrigues (1898b: 68). Orbignya barbosiana
Burret (1932: 690). Lectotype (designated by Henderson 1995):—Martius 1826: t. 96, figs. 3–6. Epitype (here
designated):—BRAZIL. Pará: Mun. Itupiranga, on Rio Tocantins, ca. 20 km downstream from Itupiranga at Caja-
zeirinha, 5°05’S 49°20’W, 23 November 1981, M. Balick, A. Anderson, J. Jangoux, J. Frazão, N. Rosa & L. Marinho
1304 (epitype NY!)
Orbignya phalerata Martius (1844: 126). Attalea glassmanii Zona (2002: 132). Type:—BOLIVIA. Chiquitos, no date, A. d’Orbigny 20
(holotype P!, isotypes F!, M n.v., M image!).
Orbignya lydiae Drude (1881: 448). Attalea lydiae (Drude) Barbosa Rodrigues (1903a: 65). Type:—BRAZIL. Rio de Janeiro: cultivated,
no date, A. Glaziou 9006 (holotype C n.v., C image!, isotypes K!, NY!, P!).
Orbignya oleifera Burret (1938: 240). Attalea vitrivir Zona (2002: 133). Type:—BRAZIL. Minas Gerais: Pirapora, 18 December 1937, M.
Burret & A. Brade 19 (holotype RB n.v., RB image!, isotype B n.v., B image!).
Orbignya brejinhoensis Glassman (1999: 84). Attalea brejinhoensis (Glassman) Zona (2002: 132). Type:—BRAZIL. Bahia: Mun. Oliveira
dos Brejinhos, 40 km E of Ibotarama, Fazenda de Brejinho da Serra Negra, 21 August 1976, S. Glassman 13041 (holotype F!, isotype
SP n.v.).
stem; sheaths 101.6(40.0–200.0) cm long; petioles 35.5(0.0–160.0) cm long, absent or well-developed; rachises
738.7(396.0–1250.0) cm long; pinnae 180(111–255) per side of rachis, regularly arranged, without auricles at the
bases; abaxial and proximal part of mid-vein with or without dense patches of brown hairs; middle pinnae 108.4(73.0–
167.5(142.0–218.0) cm long; peduncles 89.3(45.0–175.0) cm long; staminate rachillae 282(215–360), 20.8(9.5–30.0)
cm long, arranged all around rachis, with abundant raphides; staminate monad/dyad bracteoles not ribbed, crustaceous,
different in texture from the rachilla and appearing separate from it by an abaxial, proximal margin; staminate flowers
9.6(7.0–13.0) mm long, mostly in monads, rarely dyads, slightly sunken in rachillae, alternately arranged along abaxial
surface of rachillae; staminate petals irregularly shaped, joined at the bases, sometimes fused, curved over and often
split at the apices, flat in cross-section, the surfaces smooth, coriaceous, not granular tomentose, the margins smooth;
stamens 28(21–36), neither exserted from nor much shorter than petals; anthers coiled and twisted; pistillate rachillae
350(225–475), 7.3(4.5–10.5) cm long, arranged all around rachis; pistillate petals cuspidate, with irregular margins;
staminodial rings well-developed, cupular with entire margins; fruits 96.8(74.6–130.4) mm long, 56.8(42.4–99.3) mm
diameter, brown, light brown, yellow-brown, yellowish, or reddish-brown, not elongate and angled, borne on short to
long rachillae with 1-several fruits per rachilla; endocarp surfaces uneven, longitudinally ridged, with stout, rounded,
adherent fibers, without small pits proximally, the bases irregularly pointed, the endocarp becoming thinner and
flanged; endocarps in cross-section with abundant, scattered fibers, several-seeded, the abortive locules, when present,
visible in endocarp cross-section but not appressed to fertile locules; opercula sunken in endocarps and obscured by
persistent fibers; seeds 4(2–7), solid in cross-section, rarely with an irregular central cavity.
Distribution and habitat:—Surinam, Brazil (Amazonas, Bahia, Ceará, Maranhão, Mato Grosso, Minas Gerais,
Pará, Piauí, Rondônia), and Bolivia (Beni, Pando, Santa Cruz) at 404(35–850) m in deciduous forest or secondary
forest, or often forming large stands, persisting in disturbed areas (Fig. 32).
Taxonomic notes:—Specimens determined as preliminary species Orbignya brejinhoensis, O. oleifera, and O.
phalerata are polymorphic for petioles (see below for discussion of petioles) and abaxial mid-vein surfaces, and these
variables are treated as a traits. Otherwise specimens share a unique combination of qualitative variable states and are
recognized as a phylogenetic species, A. speciosa (Fig. 31). Type specimens of Orbignya lydiae and O. phalerata also
share this combination and are included.
Martius (1826) described Attalea speciosa on page 138, and gave a short description but did not refer to an
illustration nor type specimen. However, in the index to this work on page 151 he referred to illustrations (“t. 96 f. III.
3–6”). These illustrations show a fruit and seed, although the caption of the plate does not include the name A. speciosa.
The fruits from which these drawings were made are apparently lost. Henderson (1995) therefore lectotypified the
name with the illustration, t. 96. Martius (1844) gave a fuller description of A. speciosa. Subsequent authors have
confused this species (e.g., Barbosa Rodrigues 1898b, Burret 1932, Glassman 1999). Because of their confusion, and
given the type is lost and the lectotype is an illustration of fruits only, an epitype with leaves, staminate flowers, and
fruits is here designated. Cavallari & Toledo (2016) have given a discussion of the nomenclature of this species.

FIGURE 31. Attalea speciosa, left, staminate inflorescence; right, infructescence (Amazonas, Brazil).
FIGURE 32. Distribution maps of Attalea speciosa and A. tessmannii.
Glassman (1999) distinguished Orbignya brejinhoensis, O. oleifera, and O. phalerata based on several quantitative
variables, all of them overlapping. He distinguished O. oleifera from O. brejinhoensis by the 0–10 cm long petioles
and 2–3 cm long pistillate rachillae of the former, and the 80–100 cm petioles and 1 cm long pistillate rachillae of
the latter. However, the elongate petioles on specimens of O. brejinhoensis are recorded on labels as having fibrous

margins, and appear to refer to the sheaths rather than the petioles, which appears short or absent, as in O. oleifera. The
problem of petiole measurements has been discussed above in the Morphology section. The two are regarded here as
geographically separate populations, with specimens identified as O. oleifera from west of the Rio San Francisco in
Bahia and Minas Gerais and those of O. brejinhoensis from the east. Quantitatively, O. oleifera and O. brejinhoensis
are generally smaller in size than other populations. Of the other two populations from Bolivia and adjacent areas and
northeastern Brazil and adjacent areas, number of pinnae per side of the rachis differs significantly between the two
(t-test, P <0.05).
Attalea speciosa is similar to a group of species (A. cohune, A. cuatrecasana, A. eichleri, A. sagotii) having coiled
and twisted anthers. Of these it appears most similar to A. eichleri based on its staminate petal shape. Pinheiro (1997)
included this species (as Orbignya phalerata) and included it in his Brazilian Orbignya group. Freitas et al. (2016)
included A. speciosa in their Orbignya-like clade, where it was polyphyletic. Meerow et al. (2009) included it in their
Orbignya group.
Attalea speciosa forms hybrids with A. maripa, named as Markleya dahlgreniana (see notes under A. maripa). It
also forms hybrids with A. eichleri, as discussed under that species.
30. Attalea tessmannii Burret (1929a: 538). Type:—PERU. Loreto: Ost-Peru: Stromgebiet des Maranon von Iquitos
aufwärts bis zur Santiago-Mündung am Pongo de Manseriche, ca. 77°30’W, 1924, G. Tessmann 5167 (holotype B n.v.,
B image!, isotypes F!, G n.v., G image! NY!)
stem; sheaths 192.5(140.0–290.0) cm long; petioles 186.3(130.0–265.0) cm long; rachises 808.0(650.0–932.0) cm
long; pinnae 220(190–295) per side of rachis, irregularly arranged, spreading in different planes, with auricles at the
bases; middle pinnae 140.0 cm long, 5.5(4.5–6.8) cm wide; distalmost few pinnae free from one another. Inflorescence
peduncular bracts 246.7(180.0–330.0) cm long; peduncles 161.0(110.0–210.0) cm long; staminate rachillae 200,
22.9(16.5–31.0) cm long, arranged all around rachis, with abundant raphides; staminate monad/dyad bracteoles usually
ribbed, the same texture as the rachilla and continuous with it abaxially; staminate flowers 12.6(8.4–14.3) mm long,
in monads or dyads, superficial on rachillae, loosely and spirally or somewhat irregularly arranged all around rachillae
(sometimes fewer or absent from adaxial, proximal surface of rachillae); staminate petals lanceolate, acuminate,
flattened, free, not or only slightly curved at the apices, flat in cross-section, the surfaces ribbed, not coriaceous, not
granular tomentose, the margins tomentose; stamens 10(7–12), neither exserted from nor much shorter than petals;
anthers straight; pistillate rachillae 124(85–163), 11.3(9.0–13.5) cm long, arranged all around rachis; pistillate petals
cuspidate, with irregular margins; staminodial rings well-developed, cupular with entire margins; fruits 122.3(102.1–
142.1) mm long, 51.7(39.1–63.0) mm diameter, brown or brownish, not elongate and angled, borne on long rachillae
cross-section with abundant, scattered fibers, 1-several-seeded, the abortive locules, when present, visible in endocarp
2(1–3), solid in cross-section, rarely with an irregular central cavity.
Distribution and habitat:—Peru (Loreto, Madre de Dios) and Brazil (Acre) at 215(120–450) m in lowland
rainforest (Fig. 32).
Taxonomic notes:—Specimens determined as preliminary species Attalea tessmannii share a unique combination
of qualitative variable states and are recognized as a phylogenetic species, A. tessmannii (Fig. 33).
Burret (1929a) gave the type locality as Soledad on the Río Itaya (Soledad also appears on the NY isotype label
but not, apparently, on the holotype label).
Glassman (1999) considered Attalea tessmannii closely related to A. iguadummat. Both are unusual, taxonomically
isolated species but may not be closely related. Although staminate flowers of A. tessmannii are scored as having the
stamens as neither exserted from nor much shorter than petals, and having lanceolate, acuminate, flattened, petals, they
bear a remarkable resemblance to staminate flowers of Scheelea s.s. Pinnae of A. tessmannii are slightly gray abaxially
and have auricles at the bases, similar to those of A. butyracea (formerly included in Scheelea). Pinheiro (1997), based
on leaf anatomical data, considered this species to be of hybrid origin with anatomical characteristics intermediate
between those of Attalea s.s. and Scheelea. Freitas et al. (2016) placed it in their Scheelea-like clade, in a sub-clade
with A. butyracea. Attalea tessmannii was not included by Meerow et al. (2009).

FIGURE 33. Attalea tessmannii, left, habit with irregularly arranged pinnae; right, infructescence (Acre, Brazil).
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Appendix I. Quantitative variables
Abbreviations in parentheses at the end of each variable are the column labels in the Data Matrix (https://www.nybg.
org/content/uploads/2020/03/Attalea.xlsx).
1. Plant height (m); data taken from specimen labels (plheight)

2. Stem length (m); data taken from specimen labels. (stemlength)
3. Stem diameter (cm); data taken from specimen labels. (stemdiameter)
4. Number of leaves per stem; data taken from specimen labels. (leafnumber)
5. Sheath length (cm); data taken from specimen labels. (sheath)
6. Petiole length (cm); data taken from specimen labels. (petiole)
7. Rachis length (cm); data taken from specimen labels. (rachislen)
8. Number of pinnae per side of rachis; data taken from specimen labels (nodivisions)
9. Middle pinna length (cm); data taken from specimens or specimen labels. (pinlen)
10. Middle pinna width (cm); data taken from specimens. (pinwid)
11. Peduncular bract length (cm) data taken from specimen labels. (pedbract)
12. Peduncle length (cm); data taken from specimen labels. (pedunclelen)
13. Staminate rachilla length (cm); data taken from specimens or from specimen labels. (straclen)
14. Number of staminate rachillae; data taken from specimen labels. (stracnum)
15. Staminate flower length (mm); data taken from specimens. (stflowlen)
16. Number of stamens; data taken from specimens. (nostamen)
17. Pistillate rachilla length (cm); data taken from specimens or from specimen labels. (psraclen)
18. Number of pistillate rachillae; data taken from specimen labels. (psracnum)
19. Fruit length (mm); data taken from specimens. (fruitlen)
20. Fruit diameter (mm); data taken from specimens. (fruitdiam)
21. Number of seeds per fruit; data taken from specimens. (seeds)
Appendix II. Qualitative Variables
Abbreviations in parentheses at the end of each character are the column labels in the Data Matrix. The states of the
characters here are scored as ‘(1)’ or ‘(2)’ etc., and these correspond with the states in the Data Matrix (https://www.
nybg.org/content/uploads/2020/03/Attalea.xlsx).
1. Stems caulescent (1); stems acaulescent (2). (acaule)

2. Leaves arranged in five vertical ranks (1); leaves not arranged in vertical ranks (2). (vertic)
3. Petioles short or absent (1); petioles well-developed (2). (petiol)
4. Sheath margins with long, rigid fibers (1); sheath margins without long fibers (2). (shefib)
5. Pinnae regularly arranged (1); pinnae irregularly arranged, usually spreading in different planes (2); pinnae
arranged in distant clusters and usually spreading in different planes (3). (pinnae)
6. Abaxial and proximal part of mid-vein with dense patches of brown hairs (1); abaxial mid-vein with a continuous
cover of brown hairs (2); abaxial mid-vein without patches of hairs (3). (ahairs)
7. Pinnae with auricles at the bases (1); pinnae without auricles at the bases (2). (auricl)
8. Distalmost few pinnae remaining joined at their apices, sometimes with ‘windows’ between them (1); distalmost
few pinnae free from one another (2). (flabel)
9. Staminate rachillae arranged all around rachis (1); staminate rachillae absent from adaxial surface of rachis (2).
(starac)
10. Staminate rachillae with abundant raphides (1); staminate rachillae without or with few raphides (2). (toment)
11. Staminate monad/dyad bracteoles usually ribbed, the same texture as the rachilla and continuous with it abaxially
(1); staminate monad/dyad bracteoles not ribbed, crustaceous, different in texture from the rachilla and appearing
separate from it by an abaxial, proximal margin (2). (bracte)
12. Staminate flowers mostly in monads, rarely dyads, superficial on rachillae, alternately arranged along abaxial
surface of rachillae (1); staminate flowers in dyads, slightly sunken in rachillae, spirally arranged all around

rachillae (2); staminate flowers in dyads, slightly sunken in rachillae, irregularly arranged on abaxial surface of
rachillae (3); staminate flowers mostly in monads, rarely dyads, slightly sunken in rachillae, alternately arranged
along abaxial surface of rachillae (4); staminate flowers mostly in monads, rarely dyads, slightly sunken in rachillae,
spirally arranged but tending to be absent from adaxial surface of rachillae, especially proximally (5); staminate
flowers in monads or dyads, superficial on rachillae, loosely and spirally or somewhat irregularly arranged all
around rachillae (sometimes fewer or absent from adaxial, proximal surface of rachillae) (6); staminate flowers in
monads, superficial on rachillae, loosely and irregularly arranged and tending to be absent from adaxial surface
of rachillae (7); staminate flowers mostly in monads, superficial on rachillae, loosely and singly arranged along
abaxial surface of rachillae (8). (arrang)
13. Staminate petals lanceolate, acuminate, flattened, free, not or only slightly curved at the apices, flat in cross-section
(1); staminate petals linear, free, not curved at the apices, flat in cross-section (2); staminate petals spathulate, free,
curved over at the apices, flat in cross-section (3); staminate petals linear, joined at the bases, curved over at the
apices, flat in cross-section (4); staminate petals lanceolate, joined at the bases, curved outward at the apices, flat
in cross-section (5); staminate petals irregularly shaped, joined at the bases, sometimes fused, curved over and
often split at the apices, flat in cross-section (6); staminate petals lanceolate, free, not curved at the apices, flat in
cross-section (7); staminate petals linear, free, not curved at the apices, concave in cross-section (8). (stampe)
14. Surfaces of staminate petals smooth, coriaceous (1); surfaces of staminate petals ribbed, not coriaceous (2).
(surfac)
15. Surfaces of staminate petals granular tomentose (1); surfaces of staminate petals not granular tomentose (2).
(granul)
16. Margins of staminate petals minutely denticulate (1); margins of staminate petals smooth (2); margins of staminate
petals minutely serrate (3); margins of staminate petals tomentose (4). (margin)
17. Stamens 6, very rarely fewer (1); stamens more than 6 (2). (stamen)
18. Stamens exserted from petals (1); stamens much shorter than petals (2); stamens neither exserted from nor much
shorter than petals (3). (stalen)
19. Anthers straight (1); anthers coiled and twisted (2). (anther)
20. Pistillate rachillae arranged all around rachis (1); pistillate rachillae absent from adaxial surface of rachis, at least
proximally (2). (pisrac)
21. Pistillate petals cuspidate (1); pistillate petals not cuspidate (2) (cuspid)
22. Pistillate petals with elaborate, lobed margins (1); pistillate petals with irregular margins (2). (pismar)
23. Staminodial rings well-developed, cupular with entire margins (1); staminodial rings well-developed, cupular
with lobed margins (2); staminodial rings poorly developed, forming a low ring (3). (string)
24. Fruits elongate and angled by mutual pressure, borne on very short rachillae, with only 1 fruit per rachilla (1);
fruits not elongate and angled, borne on short to long rachillae with 1-several fruits per rachilla (2). (angled)
25. Fruits grayish-white tomentose at the apices (1); fruits not grayish-white tomentose at the apices (2). (frgray)
26. Endocarp surfaces uneven, longitudinally ridged, with stout, rounded, adherent fibers (1); endocarp surfaces
smooth or with few, thin fibers (2). (fibers)
27. Endocarp surfaces proximally with clear outlines of three locule divisions, these marked by irregular, small pits
(1); endocarp surfaces without small pits proximally (2). (locpit)
28. Endocarp bases irregularly pointed, the endocarp becoming thinner and flanged (1); endocarp bases regularly
rounded (2); endocarp bases regularly lobed (3). (endbas)
29. Endocarps in cross-section with abundant fibers (1); endocarps in cross-section without or with few fibers (2).
(xfiber)
30. Endocarp fibers, when present, in regular clusters (1); endocarp fibers, when present, scattered or rarely in a few,
irregular clusters (2). (endfib)
31. Endocarps 1-seeded, the abortive locules visible in endocarp cross-section appressed to fertile locule (1); endocarps
1-several-seeded, the abortive locules, when present, visible in endocarp cross-section but not appressed to fertile
locules (2). (seeded)
32. Opercula sunken in endocarps and obscured by persistent fibers (1); opercula superficial or slightly sunken in
endocarps, not obscured by fibers (2). (opercu)
33. Seeds with a well-developed central cavity in cross-section (1); seeds solid in cross-section, rarely with an irregular
central cavity (2). (hollow)
34. Fruit color is taken directly from the specimen labels. Because so many different colors are given on labels, it is
not possible to score it as in other characters. (fruitcolor)
Appendix III. Excluded and Doubtful Names
This list of name comprises published names that have been included in Attalea or related genera but for which no
type specimen is available (mostly destroyed at B); or the type specimen is insufficient for identification (doubtful
names, following Glassman 1999 or Burret 1929a); or the names are invalidly published; or the names are illegitimate;
or nomina nuda (names of new taxa published without a description or diagnosis or reference to a description or
diagnosis); or unplaced names (following Govaerts & Dransfield 2005).
Attalea acaulis Wendland (1854: 4), nom. nud.

Attalea agrestis Barbosa Rodrigues (1875: 42). Orbignya agrestis (Barbosa Rodrigues) Burret (1929a: 511). Type:—
BRAZIL. Pará: “ad fluvium Uauinchá, confluentem amnis Jamundá”, no date, J. Barbosa Rodrigues 324 (holotype
destroyed). Glassman (1999) treated this as an uncertain species because staminate flowers were not described or
illustrated in the protologue. This is followed here.
Attalea boehmii Drude (1897: 59). This name is not included by Glassman (1999) and is listed under ‘Unplaced names’
by Govaerts & Dransfield (2005).
Attalea butyrosa Loddiges ex Wendland in Kerchove de Denterghem (1878: 232), nom. nud.
Attalea coronata Loddiges ex Wendland in Kerchove de Denterghem (1878: 233). This name is not included by
Glassman (1999) and is listed under ‘Unplaced names’ by Govaerts & Dransfield (2005).
Attalea excelsa Martius (1826: 138). Lectotype (designated by Henderson 1995):—Martius 1826: t. 96. Henderson
(1995) treated this as a synonym of Attalea phalerata. However, Glassman (1999) treated the name as an uncertain
one, based on the lack of staminate flowers, and this is followed here.
Attalea grandis Wendland in Kerchove de Denterghem (1878: 233). This name is not included by Glassman (1999) and
is listed under ‘Unplaced names’ by Govaerts & Dransfield (2005).
Attalea lapidea (Gaertner) Burret (1929a: 533). Cocos lapidea Gaertner (1788: 16). Type:—Gaertner 1788: t. 6, fig. 1.
Glassman (1999) considered this to be an uncertain species based on the incomplete protologue, where only fruits
were described. This is followed here.
Attalea limbata Seemann ex Wendland in Kerchove de Denterghem (1878: 233). This name is not included by Glassman
(1999) and is listed under ‘Unplaced names’ by Govaerts & Dransfield (2005).
Attalea macoupi Sagot ex Drude (1881: 441), nom. nud.
Attalea magdalenae Linden (1881: 15). This name is not included by Glassman (1999) and is listed under ‘Unplaced
names’ by Govaerts & Dransfield (2005).
Attalea manaca Linden (1881: 15). This name is not included by Glassman (1999) and is listed under ‘Unplaced
Attalea microcarpa Martius (1844: 125). Orbignya microcarpa (Martius) Burret (1929a: 507). Neotype (designated
by Henderson 1995):—Martius 1845: t. 168. Glassman (1999) considered this to be an uncertain species based on
the lack of a type specimen and incomplete protologue. This is followed here.
Attalea microclada Burret (1935: 520), nom. nud.
Attalea monosperma Barbosa Rodrigues (1875: 42). Attalea spectabilis var. monosperma (Barbosa Rodrigues) Drude
(1881: 440). Glassman (1999) considered this to be an uncertain species based on the lack of a type specimen and
incomplete protologue. This is followed here.
Attalea pixuna Barbosa Rodrigues (1875: 43). Orbignya pixuna (Barbosa Rodrigues) Barbosa Rodrigues (1879: 49).
Glassman (1999) considered this to be an uncertain species based on the lack of a type specimen and incomplete
protologue. This is followed here.
Attalea purpurea Linden (1881: 15). This name is not included by Glassman (1999) and is listed under ‘Unplaced
Attalea puruensis Linden (1881: 15). This name is not included by Glassman (1999) and is listed under ‘Unplaced
Attalea rosii Loddiges ex Loudon (1830: 387). This name is not included by Glassman (1999) and is listed under
‘Unplaced names’ by Govaerts & Dransfield (2005).
Attalea spectabilis Martius (1826: 136). Orbignya spectabilis (Martius) Burret (1929a: 508). Type:—BRAZIL. Pará:
Serra de Barú, near Pará and Rio Negro, no date, C. Martius s.n. (holotype M, destroyed). Glassman (1999)
considered this to be an uncertain species based on the lack of a type specimen and incomplete protologue. This
is followed here.

Attalea spectabilis var. polyandra Drude (1881: 440). Type:—BRAZIL. Rio Purus, no date, Wallis s.n. (holotype
K, destroyed). This was given as a doubtful species by Glassman (1999) based on the lack of a type, and this is
followed here.
Attalea spectabilis var. typica Drude (1881: 440), nom. inval.
Attalea spinosa Meyen (1843: 469). This name is not included by Glassman (1999) and is listed under ‘Unplaced
Attalea tiasse Linden (1881: 15). This name is not included by Glassman (1999) and is listed under ‘Unplaced names’
by Govaerts & Dransfield (2005).
Attalea transitiva Barbosa Rodrigues (1879: 49). This name was treated by Glassman (1999) as a confused name.
Attalea venatorum Martius (1853: 325). Unplaced name - Govaerts & Dransfield 2005. This name is not included by
Glassman (1999) and is listed under ‘Unplaced names’ by Govaerts & Dransfield (2005).
Bornoa crassispatha (Martius) Cook (1939: 266), nom. inval.
Cocos butyrosa Wendland in Kerchove de Denterghem (1878: 240), nom. nud.
Ethnora maripa (Martius) Cook (1940: 297), nom. inval.
Heptandra phalerata (Martius) Cook (1939: 277), nom. inval.
Lithocarpus cocciformis Targioni Tozzetti ex Steudel (1841: 56), nom. illeg.
Maximiliana argentinensis Spegazzini (1917: 169). This name is not included by Glassman (1999) and is listed under
Maximiliana attaleoides Barbosa Rodrigues (1875: 41). Englerophoenix attaleoides (Barbosa Rodrigues) Barbosa
Rodrigues (1903a: 76). Attalea attaleoides (Barbosa Rodrigues) Wessels Boer (1965: 157). Type:—BRAZIL.
Pará: monte Curumú, no date, J. Barbosa Rodrigues 355 (holotype destroyed). Neototype (designated by Wessels
Boer 1965):—Barbosa Rodrigues 1903a: t. 60A. Glassman (1999) treated this as an uncertain species, based on
the lack of a type specimen and a confusing protologue. This is followed here.
Maximiliana jagua Seemann ex Wendland in Kerchove de Denterghem (1878: 251), nom. nud.
Maximiliana princeps Martius (1844: t. 4), nom. nud.
Maximiliana venatorum Wendland in Kerchove de Denterghem (1878: 251). Glassman (1999) considered this name
might apply to another genus. It is an unplaced name in Govaerts & Dransfield (2005).
Orbignya campestris Barbosa Rodrigues (1898b: 78). Type:—BRAZIL. Matto Grosso: Capão Bonito, no date, J.
Barbosa Rodrigues 240 (holotype destroyed). This was treated as a doubtful species by Glassman (1999) and this
is followed here.
Orbignya cuci Kunth ex Wendland in Kerchove de Denterghem (1878: 252), nom. nud.
Orbignya excelsa Barbosa Rodrigues (1891a: 32). This name is not included by Glassman (1999) and is listed under
Orbignya huebneri Burret (1929a: 501). Attalea huebneri (Burret) Zona (2002: 133). Type:—BRAZIL. Amazonas:
Lago Monduruçu, Rio Manacapuru, Solimões, no date, G. Huebner 64 (holotype, B n.v., B image!). Glassman
(1999) treated this as an uncertain species, based on an incomplete protologue and fragmentary type. This is
followed here.
Orbignya humilis Martius (1844: 129). Type:—BOLIVIA. Santa Cruz: Santa Ana, no date, A. d’Orbigny 22 (holotype
P, destroyed). Neotype (designated by Glassman 1999):—Martius 1845: t. 169–I. Glassman (1999) treated this as
an uncertain species, based on an incomplete protologue and lack of a type specimen. This is followed here.
Orbignya longibracteata Barbosa Rodrigues (1898b: 79). Type:—BRAZIL. Matto Grosso: Capão Bonito, no date, J.
Barbosa Rodrigues 239 (holotype destroyed). This was treated as a doubtful species by Glassman (1999) and this
is followed here.
Orbignya macrocarpa Barbosa Rodrigues (1898b: 74). Type:—BRAZIL. Matto Grosso: Capão Bonito, Serra Quebra
Cabeça, no date, J. Barbosa Rodrigues 217 (holotype destroyed). This was treated as a doubtful species by
Glassman (1999) and this is followed here.
Orbignya macropetala Burret (1929a: 507). Type:—GUYANA. Rupununi River, no date, R. Schomburgk (holotype B
n.v., B image!). This species was treated as doubtful by Glassman (1999) based on a fragmentary type specimen
and incomplete protologue, and this is followed here.
Orbignya speciosa Barbosa Rodrigues (1891a: 32). Uncertain name, Glassman (1999).
Scheelea cubensis Burret (1929a: 671). Type:—CUBA. No locality, no date, J. Gundlach s.n. (holotype B n.v.). This
was treated as a doubtful species by Glassman (1999). No species of Attalea is known from Cuba.
Scheelea excelsa Barbosa Rodrigues (1891a: 30), nom. illeg.
Scheelea excelsa Karsten (1856: 267). Type:—Karsten 1866: t. 176, figs. 10–11. This species was treated as doubtful
by Glassman (1999) based on an incomplete protologue and no type specimen, and this is followed here.
Scheelea imperialis Bull (1875: 589). This name is not included by Glassman (1999) and is listed under ‘Unplaced
Scheelea martiana Burret (1929a: 661). Proposed as a new name for Attalea excelsa Martius, here treated as a doubtful
name following Glassman (1999).
Scheelea unguis Nicholson (1886: 385). This name is not included by Glassman (1999) and is listed under ‘Unplaced
Temenia regia (Martius) Cook (1939: 276), nom. inval.
Appendix IV. Index of Names
Accepted taxa are bold face.
Attalea Kunth in Humboldt, Bonpland & Kunth ................................................................................................................................... 13

Attalea acaulis Burret ................................................................................................................................................................................. 37
Attalea acaulis Wendland ........................................................................................................................................................................... 70
Attalea agrestis Barbosa Rodrigues ............................................................................................................................................................ 70
Attalea allenii Moore ................................................................................................................................................................................. 16
Attalea amygdalina Kunth in Humboldt, Bonpland & Kunth .............................................................................................................. 17
Attalea amylacea (Barbosa Rodrigues) Zona ............................................................................................................................................. 50
Attalea anisitsiana (Barbosa Rodrigues) Zona ........................................................................................................................................... 50
Attalea apoda Burret ................................................................................................................................................................................... 35
Attalea attaleoides (Barbosa Rodrigues) Wessels Boer .............................................................................................................................. 71
Attalea barreirensis Glassman .................................................................................................................................................................. 19
Attalea bassleriana (Burret) Zona .............................................................................................................................................................. 20
Attalea blepharopus Martius ....................................................................................................................................................................... 20
Attalea boehmii Drude ................................................................................................................................................................................ 70
Attalea borgesiana Bondar ......................................................................................................................................................................... 40
Attalea borgesiana Hawkes ........................................................................................................................................................................ 40
Attalea brasiliensis Glassman ..................................................................................................................................................................... 28
Attalea brejinhoensis (Glassman) Zona ...................................................................................................................................................... 60
Attalea burretiana Bondar .......................................................................................................................................................................... 49
Attalea butyracea (Mutis ex Linnaeus f.) Wessels Boer .......................................................................................................................... 19
Attalea butyrosa Loddiges ex Wendland in Kerchove de Denterghem ...................................................................................................... 70
Attalea camopiensis (Glassman) Zona ........................................................................................................................................................ 40
Attalea camposportoana Burret .................................................................................................................................................................. 28
Attalea cephalotus Poeppig ex Martius .................................................................................................................................................... 23
Attalea cohune Martius ............................................................................................................................................................................. 25
Attalea colenda (Cook) Balslev & Henderson ......................................................................................................................................... 27
Attalea compta Martius ............................................................................................................................................................................. 28
Attalea compta var. acaulis Martius ........................................................................................................................................................... 40
Attalea concentrista Bondar ....................................................................................................................................................................... 49
Attalea concinna (Barbosa Rodrigues) Burret ............................................................................................................................................ 32
Attalea coronata Loddiges ex Wendland in Kerchove de Denterghem ...................................................................................................... 70
Attalea crassispatha (Martius) Burret ..................................................................................................................................................... 29
Attalea cryptanthera Wessels Boer ............................................................................................................................................................. 46
Attalea cuatrecasana (Dugand) Henderson, Galeano & Bernal ............................................................................................................ 30
Attalea dahlgreniana (Bondar) Wessels Boer ............................................................................................................................................. 46
Attalea degranvillei (Glassman) Zona ........................................................................................................................................................ 40
Attalea dubia (Martius) Burret ................................................................................................................................................................. 32
Attalea eichleri (Drude) Henderson ......................................................................................................................................................... 34
Attalea excelsa Martius ............................................................................................................................................................................... 70
Attalea exigua Drude ................................................................................................................................................................................. 35
Attalea fairchildensis (Glassman) Zona ...................................................................................................................................................... 20
Attalea ferruginea Burret ............................................................................................................................................................................ 54
Attalea funifera Martius ........................................................................................................................................................................... 37
Attalea funifera var. acaulis (Burret) Bondar ............................................................................................................................................. 37
Attalea geraensis (“ceraensis”) Barbosa Rodrigues .................................................................................................................................. 35
Attalea glassmanii Zona ............................................................................................................................................................................. 60
Attalea goeldiana Huber ............................................................................................................................................................................. 43
Attalea gomphococca Martius .................................................................................................................................................................... 50
Attalea grandis Wendland in Kerchove de Denterghem ............................................................................................................................. 70

Attalea guacuyule (Liebmann ex Martius) Zona ........................................................................................................................................ 25
Attalea guaranitica Barbosa Rodrigues ...................................................................................................................................................... 35
Attalea guianensis (Glassman) Zona ........................................................................................................................................................ 39
Attalea hoehnei Burret ................................................................................................................................................................................ 50
Attalea huebneri (Burret) Zona ................................................................................................................................................................... 71
Attalea humboldtiana Spruce ...................................................................................................................................................................... 20
Attalea humilis Martius ex Sprengel ........................................................................................................................................................ 40
Attalea iguadummat de Nevers ................................................................................................................................................................. 41
Attalea indaya Drude .................................................................................................................................................................................. 32
Attalea insignis (Martius) Drude in Engler & Prantl ............................................................................................................................. 43
Attalea kewensis (Hooker) Zona ................................................................................................................................................................. 20
Attalea lapidea (Gaertner) Burret ............................................................................................................................................................... 70
Attalea lauromuelleriana (Barbosa Rodrigues) Zona ................................................................................................................................. 50
Attalea leandroana (Barbosa Rodrigues) Zona .......................................................................................................................................... 20
Attalea liebmannii (Beccari) Zona .............................................................................................................................................................. 20
Attalea limbata Seemann ex Wendland in Kerchove de Denterghem ........................................................................................................ 70
Attalea luetzelburgii (Burret) Wessels Boer ............................................................................................................................................. 45
Attalea lundellii (Bartlett) Zona .................................................................................................................................................................. 20
Attalea lydiae (Drude) Barbosa Rodrigues ................................................................................................................................................. 60
Attalea macoupi Sagot ex Drude ................................................................................................................................................................ 70
Attalea macrocarpa (Karsten) Linden ........................................................................................................................................................ 20
Attalea macrolepis (Burret) Wessels Boer .................................................................................................................................................. 20
Attalea macropetala (Burret) Wessels Boer ................................................................................................................................................ 46
Attalea magdalenae Linden ........................................................................................................................................................................ 70
Attalea magdalenica (Dugand) Zona .......................................................................................................................................................... 20
Attalea manaca Linden ............................................................................................................................................................................... 70
Attalea maracaibensis Martius ................................................................................................................................................................... 19
Attalea maripa (Aublet) Martius .............................................................................................................................................................. 45
Attalea maripensis (Glassman) Zona .......................................................................................................................................................... 39
Attalea microcarpa Martius ........................................................................................................................................................................ 70
Attalea microclada Burret ........................................................................................................................................................................... 70
Attalea monogyna Burret ............................................................................................................................................................................ 28
Attalea monosperma Barbosa Rodrigues .................................................................................................................................................... 70
Attalea moorei (Glassman) Zona ................................................................................................................................................................ 23
Attalea nucifera Karsten ........................................................................................................................................................................... 47
Attalea oleifera Barbosa Rodrigues ......................................................................................................................................................... 49
Attalea osmantha (Barbosa Rodrigues) Wessels Boer ................................................................................................................................ 20
Attalea parviflora Barbosa Rodrigues ........................................................................................................................................................ 50
Attalea pacensis Moraes & Pintaud ............................................................................................................................................................ 50
Attalea peruviana Zona ............................................................................................................................................................................... 20
Attalea phalerata Martius ex Sprengel .................................................................................................................................................... 50
Attalea phalerata Martius ex Sprengel var. concinna Moreno Suárez & Moreno Suárez ......................................................................... 23
Attalea piassabossu Bondar ........................................................................................................................................................................ 39
Attalea pindobassu Bondar ....................................................................................................................................................................... 52
Attalea pixuna Barbosa Rodrigues .............................................................................................................................................................. 70
Attalea plowmanii (Glassman) Zona ........................................................................................................................................................ 53
Attalea polysticha (Burret) Wessels Boer ................................................................................................................................................... 56
Attalea princeps Martius ............................................................................................................................................................................. 50
Attalea purpurea Linden ............................................................................................................................................................................. 70
Attalea puruensis Linden ............................................................................................................................................................................ 70
Attalea pycnocarpa Wessels Boer ............................................................................................................................................................... 57
Attalea racemosa Spruce ........................................................................................................................................................................... 54
Attalea regia (Martius) Wessels Boer ......................................................................................................................................................... 45
Attalea rhynchocarpa Burret ...................................................................................................................................................................... 17
Attalea rosii Loddiges ex Loudon ............................................................................................................................................................... 70
Attalea rostrata Oersted .............................................................................................................................................................................. 20
Attalea sagotii (Trail ex Im Thurn) Wessels Boer ................................................................................................................................... 56
Attalea salazarii (Glassman) Zona ............................................................................................................................................................. 20
Attalea salvadorensis Glassman ................................................................................................................................................................. 41
Attalea seabrensis Glassman ...................................................................................................................................................................... 52
Attalea septuagenata Dugand ................................................................................................................................................................... 58
Attalea speciosa Martius ........................................................................................................................................................................... 60
Attalea spectabilis Martius ......................................................................................................................................................................... 70
Attalea spectabilis var. monosperma (Barbosa Rodrigues) Drude ............................................................................................................. 70
Attalea spectabilis var. polyandra Drude .................................................................................................................................................... 71
Attalea spectabilis var. typica Drude .......................................................................................................................................................... 71
Attalea spinosa Meyen ................................................................................................................................................................................ 71

Attalea tessmannii Burret .......................................................................................................................................................................... 62
Attalea tiasse Linden ................................................................................................................................................................................... 71
Attalea transitiva Barbosa Rodrigues ......................................................................................................................................................... 71
Attalea uberrima Dugand ........................................................................................................................................................................... 17
Attalea venatorum Martius ......................................................................................................................................................................... 71
Attalea victoriana Dugand .......................................................................................................................................................................... 17
Attalea vitrivir Zona .................................................................................................................................................................................... 60
Attalea wallisii Huber ................................................................................................................................................................................. 20
Attalea weberbaueri (Burret) Zona ............................................................................................................................................................. 50
Attalea wesselsboerii (Glassman) Zona ...................................................................................................................................................... 20
Attalea × minarum (Balick, Anderson & Medeiros-Costa) Zona ............................................................................................................... 29
Attalea × piasssabossu Bondar ................................................................................................................................................................... 39
Attalea × teixeriana (Bondar) Zona ........................................................................................................................................................... 34
Attalea × voeksii Noblick ex Glassman ...................................................................................................................................................... 39
Bornoa Cook ............................................................................................................................................................................................... 13
Bornoa crassispatha (Martius) Cook .......................................................................................................................................................... 71
Cocos butyracea Mutis ex Linnaeus f. ....................................................................................................................................................... 19
Cocos butyrosa Wendland in Kerchove de Denterghem ............................................................................................................................. 71
Cocos cocoyule Martius .............................................................................................................................................................................. 25
Cocos guacuyule Liebmann ex Martius ...................................................................................................................................................... 25
Cocos lapidea (Gaertner) Burret ................................................................................................................................................................. 70
Cocos lapidea Gaertner ............................................................................................................................................................................... 70
Cocos regia Liebmann in Martius .............................................................................................................................................................. 20
Englerophoenix Kuntze ............................................................................................................................................................................... 13
Englerophoenix attaleoides (Barbosa Rodrigues) Barbosa Rodrigues ....................................................................................................... 71
Englerophoenix caribaeum (Grisebach & Wendland) Kuntze ................................................................................................................... 45
Englerophoenix insignis (Martius) Kuntze ................................................................................................................................................. 43
Englerophoenix longirostrata (Barbosa Rodrigues) Barbosa Rodrigues ................................................................................................... 45
Englerophoenix maripa (Aublet) Kuntze ................................................................................................................................................... 45
Englerophoenix regia (Martius) Kuntze ..................................................................................................................................................... 45
Englerophoenix tetrasticha (Drude) Barbosa Rodrigues ............................................................................................................................ 50
Ethnora Cook .............................................................................................................................................................................................. 13
Ethnora maripa (Martius) Cook ................................................................................................................................................................. 71
Heptandra Cook .......................................................................................................................................................................................... 13
Heptandra phalerata (Martius) Cook ......................................................................................................................................................... 71
Lithocarpus Targioni Tozzetti ex Steudel ................................................................................................................................................... 13
Lithocarpus cocciformis Targioni Tozzetti ex Steudel ................................................................................................................................ 71
Markleya dahlgreniana Bondar .................................................................................................................................................................. 46
Markleya Bondar ......................................................................................................................................................................................... 13
Maximiliana Martius ................................................................................................................................................................................... 13
Maximiliana argentinensis Spegazzini ....................................................................................................................................................... 71
Maximiliana attaleoides Barbosa Rodrigues .............................................................................................................................................. 71
Maximiliana caribaea Grisebach & Wendland ex Grisebach ..................................................................................................................... 45
Maximiliana crassispatha Martius ............................................................................................................................................................. 29
Maximiliana elegans Karsten ..................................................................................................................................................................... 45
Maximiliana insignis Martius ..................................................................................................................................................................... 43
Maximiliana jagua W. Seem. ex Wendland in Kerchove de Denterghem .................................................................................................. 71
Maximiliana longirostrata Barbosa Rodrigues ........................................................................................................................................... 45
Maximiliana macrogyne Burret .................................................................................................................................................................. 46
Maximiliana macropetala Burret ................................................................................................................................................................ 46
Maximiliana maripa (Aublet) Drude .......................................................................................................................................................... 45
Maximiliana martiana Karsten ................................................................................................................................................................... 45
Maximiliana princeps Martius .................................................................................................................................................................... 71
Maximiliana regia Martius ......................................................................................................................................................................... 45
Maximiliana stenocarpa Burret .................................................................................................................................................................. 46
Maximiliana tetrasticha Drude ................................................................................................................................................................... 50
Maximiliana venatorum Wendland in Kerchove de Denterghem ............................................................................................................... 71
Orbignya Martius ex Endlicher .................................................................................................................................................................. 13
Orbignya agrestis (Barbosa Rodrigues) Burret .......................................................................................................................................... 70
Orbignya barbosiana Burret ....................................................................................................................................................................... 60
Orbignya brejinhoensis Glassman .............................................................................................................................................................. 60
Orbignya campestris Barbosa Rodrigues ................................................................................................................................................... 71
Orbignya cohune (Martius) Dahlgren ex Standley ..................................................................................................................................... 25
Orbignya crassispatha (Martius) Glassman ............................................................................................................................................... 29
Orbignya cuatrecasana Dugand ................................................................................................................................................................. 30
Orbignya cuci Kunth ex Wendland in Kerchove de Denterghem ............................................................................................................... 71
Orbignya dammeriana Barbosa Rodrigues ................................................................................................................................................ 25

Orbignya dubia Martius .............................................................................................................................................................................. 32
Orbignya eichleri Drude ............................................................................................................................................................................. 34
Orbignya excelsa Barbosa Rodrigues ......................................................................................................................................................... 71
Orbignya guacuyule (Liebmann ex Martius) Hernández Xolocotzi ........................................................................................................... 25
Orbignya huebneri Burret ........................................................................................................................................................................... 71
Orbignya humilis Martius ........................................................................................................................................................................... 71
Orbignya longibracteata Barbosa Rodrigues ............................................................................................................................................. 71
Orbignya luetzelburgii Burret ..................................................................................................................................................................... 45
Orbignya lydiae Drude ............................................................................................................................................................................... 60
Orbignya macrocarpa Barbosa Rodrigues ................................................................................................................................................. 71
Orbignya macropetala Burret ..................................................................................................................................................................... 71
Orbignya martiana Barbosa Rodrigues ...................................................................................................................................................... 60
Orbignya microcarpa (Martius) Burret ...................................................................................................................................................... 70
Orbignya oleifera Burret ............................................................................................................................................................................. 60
Orbignya phalerata Martius ....................................................................................................................................................................... 60
Orbignya pixuna (Barbosa Rodrigues) Barbosa Rodrigues ........................................................................................................................ 70
Orbignya polysticha Burret ......................................................................................................................................................................... 56
Orbignya racemosa (Spruce) Drude ........................................................................................................................................................... 54
Orbignya sabulosa Barbosa Rodrigues ...................................................................................................................................................... 56
Orbignya sagotii Trail ex Im Thurn ............................................................................................................................................................ 56
Orbignya speciosa (Martius) Barbosa Rodrigues ....................................................................................................................................... 71
Orbignya spectabilis (Martius) Burret ........................................................................................................................................................ 70
Orbignya urbaniana Dammer ..................................................................................................................................................................... 34
Orbignya × teixeriana Bondar..................................................................................................................................................................... 34
Palma maripa Aublet .................................................................................................................................................................................. 45
Parascheelea Dugand ................................................................................................................................................................................. 14
Parascheelea anchistropetala Dugand ....................................................................................................................................................... 45
Parascheelea luetzelburgii (Burret) Dugand .............................................................................................................................................. 45
Pindarea Barbosa Rodrigues ...................................................................................................................................................................... 13
Pindarea concinna Barbosa Rodrigues ....................................................................................................................................................... 32
Pindarea dubia (Martius) Hawkes .............................................................................................................................................................. 32
Pindarea fastuosa Barbosa Rodrigues ........................................................................................................................................................ 32
Sarinia Cook ............................................................................................................................................................................................... 14
Sarinia funifera (Martius) Cook ................................................................................................................................................................. 37
Scheelea Karsten ......................................................................................................................................................................................... 13
Scheelea amylacea Barbosa Rodrigues ...................................................................................................................................................... 50
Scheelea anizitziana Barbosa Rodrigues .................................................................................................................................................... 50
Scheelea attaleoides Karsten ...................................................................................................................................................................... 43
Scheelea bassleriana Burret ....................................................................................................................................................................... 20
Scheelea blepharopus (Martius) Burret ...................................................................................................................................................... 20
Scheelea brachyclada Burret ...................................................................................................................................................................... 20
Scheelea butyracea (Mutis ex Linnaeus f.) Karsten ex Wendland in Kerchove de Denterghem ............................................................... 19
Scheelea camopiensis Glassman ................................................................................................................................................................. 40
Scheelea cephalotes (Poeppig ex Martius) Karsten .................................................................................................................................... 23
Scheelea corumbaensis (Barbosa Rodrigues) Barbosa Rodrigues ............................................................................................................. 50
Scheelea costaricensis Burret ..................................................................................................................................................................... 20
Scheelea cubensis Burret ............................................................................................................................................................................ 71
Scheelea curvifrons Bailey .......................................................................................................................................................................... 20
Scheelea degranvillei Glassman ................................................................................................................................................................. 40
Scheelea dryanderae Burret ........................................................................................................................................................................ 20
Scheelea excelsa Barbosa Rodrigues .......................................................................................................................................................... 71
Scheelea excelsa Karsten ............................................................................................................................................................................ 71
Scheelea fairchildensis Glassman ............................................................................................................................................................... 20
Scheelea goeldiana (Huber) Burret ............................................................................................................................................................ 43
Scheelea gomphococca (Martius) Burret .................................................................................................................................................... 50
Scheelea guianensis Glassman ................................................................................................................................................................... 39
Scheelea huebneri Burret ............................................................................................................................................................................ 50
Scheelea humboldtiana (Spruce) Burret ..................................................................................................................................................... 20
Scheelea imperialis Bull ............................................................................................................................................................................. 72
Scheelea insignis (Martius) Karsten ........................................................................................................................................................... 43
Scheelea kewensis Hooker .......................................................................................................................................................................... 20
Scheelea lauromuelleriana Barbosa Rodrigues .......................................................................................................................................... 50
Scheelea leandroana Barbosa Rodrigues ................................................................................................................................................... 20
Scheelea liebmannii Beccari ....................................................................................................................................................................... 20
Scheelea lundellii Bartlett ........................................................................................................................................................................... 20
Scheelea macrocarpa Karsten .................................................................................................................................................................... 20
Scheelea macrolepis Burret ........................................................................................................................................................................ 20

Scheelea magdalenica Dugand ................................................................................................................................................................... 20
Scheelea maracaibensis (Martius) Burret ................................................................................................................................................... 19
Scheelea maripa (Aublet) Wendland in Kerchove de Denterghem ............................................................................................................ 45
Scheelea maripensis Glassman ................................................................................................................................................................... 39
Scheelea martiana Burret ........................................................................................................................................................................... 72
Scheelea microspadix Burret ...................................................................................................................................................................... 50
Scheelea moorei Glassman ......................................................................................................................................................................... 23
Scheelea osmantha Barbosa Rodrigues ...................................................................................................................................................... 20
Scheelea parviflora (Barbosa Rodrigues) Barbosa Rodrigues ................................................................................................................... 50
Scheelea passargei Burret ........................................................................................................................................................................... 20
Scheelea phalerata (Martius ex Sprengel) Burret ...................................................................................................................................... 50
Scheelea plowmanii Glassman .................................................................................................................................................................... 53
Scheelea preussii Burret .............................................................................................................................................................................. 20
Scheelea princeps (Martius) Karsten .......................................................................................................................................................... 50
Scheelea princeps var. corumbaensis Barbosa Rodrigues .......................................................................................................................... 50
Scheelea quadrisperma Barbosa Rodrigues ............................................................................................................................................... 50
Scheelea quadrisulcata Barbosa Rodrigues ................................................................................................................................................ 50
Scheelea regia Karsten ................................................................................................................................................................................ 20
Scheelea rostrata (Oersted) Burret ............................................................................................................................................................. 20
Scheelea salazarii Glassman ....................................................................................................................................................................... 20
Scheelea stenorhyncha Burret ..................................................................................................................................................................... 20
Scheelea tessmannii Burret ......................................................................................................................................................................... 20
Scheelea tetrasticha (Drude) Burret ........................................................................................................................................................... 50
Scheelea unguis Nicholson ......................................................................................................................................................................... 72
Scheelea urbaniana Burret ......................................................................................................................................................................... 20
Scheelea wallisii (Huber) Burret ................................................................................................................................................................. 20
Scheelea weberbaueri Burret ...................................................................................................................................................................... 50
Scheelea wesselsboerii Glassman ............................................................................................................................................................... 20
Scheelea zonensis Bailey ............................................................................................................................................................................ 20
Temenia Cook ............................................................................................................................................................................................. 13
Temenia regia (Martius) Cook .................................................................................................................................................................... 72
× Attabignya Balick, Anderson & Medeiros-Costa ..................................................................................................................................... 13
× Attabignya minarum Balick, Anderson & Medeiros-Costa .................................................................................................................... 14
× Maximbignya dahlgreniana (Bondar) Glassman .................................................................................................................................... 14
Ynesa Cook ................................................................................................................................................................................................. 14
Ynesa colenda Cook .................................................................................................................................................................................... 27
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. The curators
of the following herbaria sent these loans, or allowed me to study specimens in their care: AAU, BH, BM, BR, COAH,
COL, F, K, NY, MO, P, and US. I am particularly grateful to the curators of B and M for making type images available,
and to Paula Leitman of BR for supplying type images. Dr. Michael Balick of the New York Botanical Garden, Drs.
Rodrigo Bernal and the late Gloria Galeano of the Universidad Nacional in Colombia, and Renata Martins of the
Universidade de Brasília shared their knowledge of Attalea. Dr. Fred Barrie of the Missouri Botanical Garden provided
nomenclatural expertise. I am grateful to Drs. Larry Noblick of the Montgomery Botanical Center and Scott Zona of
Florida International University for reviewing earlier versions of the manuscript, and to two anonymous reviewers.

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Phytotaxa 444 (1): 001–076 ISSN 1179-3155 (print edition)

Accepted by Carlos Lehnebach: 23 Mar. 2020; published: 25 May 2020 1

FIRST PUBLISHED IN 2020 BY

© 2020 Magnolia Press

ISSN 1179-3155 (print edition)

2 • Phytotaxa 444 (1) © 2020 Magnolia Press HENDERSON

Key words: Maximiliana, Neotropics, Orbignya, Palmae, Scheelea

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Materials and Methods

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Distribution and Habitat

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Key to the species of Attalea

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FIGURE 6. Distribution maps of Attalea cephalotus, A. cohune, A. colenda, and A. compta.

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FIGURE 21. Attalea insignis, staminate inflorescence, infructescence (Amazonas, Brazil).

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Distribution and habitat:—Colombia (Santander) in the Magdalena valley at 227(200–300) m in lowland

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FIGURE 32. Distribution maps of Attalea speciosa and A. tessmannii.

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