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Title: Neotropical primates
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Title: Neotropical primates a newsletter of the Neotropical Section of the IUCNSSC Primate Specialist Group
Abbreviated Title: Neotrop. primates
Physical Description: v. : ill. ; 27 cm.
Language: English
Creator: IUCN/SSC Primate Specialist Group -- Neotropical Section
IUCN/SSC Primate Specialist Group -- Neotropical Section
Conservation International
Center for Applied Biodiversity Science
Publisher: Conservation International
Place of Publication: Belo Horizonte Minas Gerais Brazil
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Publication Date: June 2000
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Subject: Primates -- Periodicals -- Latin America   ( lcsh )
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Wildlife conservation -- Periodicals   ( lcsh )
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Full Text

ISSN 1413-4703


pnma es

JUNE 2000

A Journal and Newsletter of the
Neotropical Section of the IUCN/SSC
Primate Specialist Group

"An Assessment of the Diversity of
New World Primates"




Editors: Anthony B. Rylands and Emesto Rodrfguez-Luna
PSG Chairman: Russell A. Mittermeier
PSG Deputy Chairmen: Anthony B. Rylands and William R. Konstant

Neotropical Primates
A Journal and Newsletter of the Neotropical Section of the IUCN/SSC Primate Specialist Group

Center for Applied Biodiversity Science
Conservation International
2501 M. St. NW, Suite 200, Washington, DC 20037

ISSN 1413-4703 Abbreviation: Neotrop. Primates

Anthony B. Rylands, Center for Applied Biodiversity Science, Conservation International, Washington, DC
Ernesto Rodriguez-Luna, UniversidadVeracruzana, Xalapa, Mexico

Assistant Editor
Jennifer Pervola, Center for Applied Biodiversity Science, Conservation International, Washington, DC

Editorial Board
Hannah M. Buchanan-Smith, University of Stirling, Stirling, Scotland, UK
Adelmar E Coimbra-Filho,Academia Brasileira de Cincias, Rio de Janeiro, Brazil
Liliana Cort6s-Ortiz, Universidad Veracruzana, Xalapa, Mexico
Carolyn M. Crockett, Regional Primate Research Center, University of Washington, Seattle, WA, USA
Stephen E Ferrari, Universidade Federal do Para, Belem, Brazil
EckhardW Heymann, Deutsches Primatenzentrum, Gbttingen, Germany
William R. Konstant, Conservation International Washington, DC
Russell A. Mittermeier, Conservation International, Washington, DC
Marta D. Mudry, Universidad de Buenos Aires, Argentina
Horacio Schneider, Universidade Federal do Para, Belem, Brazil
Karen B. Strier, University ofWisconsin, Madison, Wisconsin, USA
Maria Emilia Yamamoto, Universidade Federal do Rio Grande do Norte, Natal, Brazil

Primate Specialist Group
Chairman Russell A. Mittermeier
Deputy Chairs Anthony B. Rylands & William R. Konstant
Co-Vice Chairs for the Neotropical RegionAnthony B. Rylands & Ernesto Rodriguez-Luna
Vice Chair forAsiaArdithA. Eudey
Vice Chair for Africa Thomas M. Butynski
Vice Chair for Madagascar Jbrg U. Ganzhorn

Design and composition:
Glenda P. Fabregas, Center for Applied Biodiversity Science, Conservation International, Washington, DC.

Front Cover:
The black-headed uacari, Cacajao melanocephalus ouakary.

This issue of NeotropicalPrimates was kindly sponsored by the Margot Marsh Biodiversity Foundation, 432 Walker Road, Great Falls, Virginia
22066, USA, and the Houston Zoological Gardens Conservation Program, General Manager Donald G. Olson, 1513 North MacGregor,



A DMsion of tlhe Houston
Parks and Recreation Department

Neotropical Primates 8(2), June 2000 61


Anthony B. Rylands
Center for Applied Biodiversity Science (CABS), Conservation International, 2015 M Street N.W., Washington, DC 20037,
Horacio Schneider
Universidade Federal do Para, Campus de Braganca, 68600-000 Braganca, Para, Brazil.
Alfredo Langguth
Departamento de Sistemitica e Ecologia CCEN, Universidade Federal da Parafba, 58059-900 Joao Pessoa, Parafba, Brazil.
Russell A. Mittermeier
Conservation International, 2015 M Street N.W., Washington, DC 20037, USA.
Colin P. Groves
Ari i, Ji. -.,- and Anirl'-..p..1..;,., Faculty of Arts, The Australian National University, Canberra, A.C.T. 0200, Australia.
Ernesto Rodriguez-Luna
Institute de Neuroetologfa, Universidad Veracruzana, Xalapa, 91000 Veracruz, Mexico.


An understanding of the full diversity of the primates is vital for priority-setting for conservation purposes, both for captive
breeding and in situ conservation measures. Here we provide a full, annotated listing of the Neotropical primate species and
subspecies resulting from a workshop, "Primate Taxonomy for the New Millennium", which brought together experts in
primate taxonomy and biogeography, morphologists and geneticists. It was organized by the IUCN/SSC Primate Specialist
Group (PSG) and held at the Disney Institute, Orlando, Florida, in February 2000. The list separates the Platyrrhini into five
families (Callitrichidae, Cebidae, Aotidae, Pitheciidae and Atelidae), 18 genera (Cebuella, Mico, Callithrix, Saguinus,
Leontopithecus, Callimico, Saimiri, Cebus, Aotus, Callicebus, Pithecia, Chiropotes, Cacajao, Alouatta, Ateles, Lagothrix, Oreonax
and Brachyteles), 110 species, and 205 species and subspecies.

Key Words Primates, taxonomy, Platyrrhini.


"Taxonomy precedes conservation.. .Without the formal struc-
ture of names and an agreed system of usage, there can be no
understanding of what exists to be conserved." (Collar, 1997).

The aims of this assessment of the diversity of the New World
primates are to have at hand a listing of primates as a basis for
conservation measures, both ex situ and in situ, and to stimu-
late further research into the systematics and taxonomy of
the group. For each genus we have attempted to provide a
summary of the different views regarding their systematics,
based on the diverse disciplines which identify characteristics
that contribute to the recognition of different forms: Mor-
phology, Genetics, Physiology and Behavior. Conservation
measures and the assessment of the conservation status, as well
as the taxonomic status, of any particular primate are compli-
cated when geographic distributions are poorly known. They
are poorly known for the majority. Many new taxa have been
discovered in the last two decades (see Table 1), and much
research is still required. For some, habitat destruction on a
grand scale has already denied us the possibility of ever un-
derstanding their original distribution patterns.

The taxonomic listing that we publish here is of course not
definitive, but will serve as a working basis for the action of

the IUCN/SSC Primate Specialist Group (PSG). It resulted
from a workshop "Primate Taxonomy for the New Millen-
nium", held at the Disney Institute, Orlando, Florida, 25-29
February 2000. It was organized by the PSG Chair Russell
A. Mittermeier in collaboration with Don Melnick, Execu-
tive Director of the Center for Environmental Research and
Conservation, Columbia University, NY, and John E Oates,
Professor at Hunter College, City University of New York,
NY, and was sponsored by the Margot Marsh Biodiversity
Foundation, Virginia, and the Disney Institute, Orlando,
Florida, which kindly provided the venue and accommoda-

The Workshop involved the collective brainstorming of ap-
proximately 25 field primatologists, taxonomists, biogeogra-
phers, morphologists and geneticists in discussions of two
major questions: 1) the identification of taxa for which little is
known or for which there is conflicting evidence and opin-
ions from the different disciplines regarding their systematics
and the validity or otherwise of described forms; and 2) the
establishment of a single taxonomic listing for the primates,
based on the evidence available today, most particularly from
morphological and genetic research. Those who contributed
to the group for the Neotropical primates included: Russell
A. Mittermeier (Conservation International, Washington,
DC), Colin P. Groves (Australian National University,

Our thanks especially to Rick Barongi, I i I I I, I ... 1 consultantt for Walt Disney's "Animal Kingdom" theme park, and currently Director of the Houston
Zoological Gardens, Houston, TX (for many years a sponsor ofNeotropicalPrimates), for hosting the meeting.

62 Neotropical Primates 8(2), June 2000

Canberra), Horacio Schneider (Universidade Federal do Pard,
Belkm, Brazil), Douglas Brandon-Jones (British Museum of
Natural History, London), Ernesto Rodrfguez-Luna
(Universidad Veracruzana, Xalapa, Mexico), Alfredo
Langguth (Universidade Federal do Parafba, Joao Pessoa, Bra-
zil), Peter Grubb (London, UK), and Anthony B. Rylands,
(Center for Applied Biodiversity Science, Conservation In-
ternational, Washington, DC). Particularly important was the
contribution of Colin Groves, whose book on primate tax-
onomy is soon to be published by the Smithsonian Institu-
tion Press, Washington, DC.

Taxonomy Families and Subfamilies

The taxonomy of the New World primates has undergone
considerable change over the last two decades, stimulated in
large part by the extensive revision of the callitrichids by Pro-
fessor Philip Hershkovitz (1977, 1979a, 1982), and his sub-
sequent reviews of the cebid genera, Aotus, Saimiri, Chiropotes,
Pithecia, Cacajao and Callicebus (1979b, 1983, 1984, 1985,
1987a, 1987b, 1990). Considerable attention has also been
given to the phylogeny and taxonomy at the family and sub-
family level, principally through morphological studies
(Rosenberger, 1980, 1981; Rosenberger and Coimbra-Filho,
1984; Rosenberger and Strier 1989; Rosenberger et al.
1990), but also more recently using chromosome and mo-
lecular genetics (Dutrillaux, 1988; Schneider et al., 1993,
1995, 1996). Cytotaxonomy is also becoming increasingly
important for systematics at the species and subspecies level
(see, for example the revisions of Aotus by Hershkovitz 1983;
Nagamachi et al. 1995, 1996, 1997, 1999).

Table 1. New World primates described since 1980.

Species name
1. Callithrix nigriceps Ferrari & Lopes, 1992
2. Callithrix argentata marcai Alperin, 1993


Platyrrhine systematics at the family and subfamily level was
reviewed by Rosenberger (1981). The predominant classifi-
cation during this century has involved the use of just two
families, the Callitrichidae (or, formerly, Hapalidae), with
Cebuella, C .-'. Saguinus and Leontopithecus, and Cebidae
(including the remaining genera), with Callimico being placed
in either of the two, or in its own family (Hill, 1957; Dollman,
1933; Hershkovitz, 1977). This system was maintained in all
of the major syntheses published over the last 75 years (for
example, Simpson, 1945; Hill, 1957, 1960, 1962; Cabrera,
1958; Napier and Napier, 1967; Simons, 1972; and
Hershkovitz, 1977). Hershkovitz (1977) placed the extant
cebids in seven subfamilies: Saimiriinae, Aotinae, Callicebinae,
Pitheciinae, Cebinae, Alouattinae and Atelinae (Table 2).
Groves (1989) divided the Cebidae into five families (Cebidae,
Aotidae, Atelidae, Callicebidae, and Pitheciidae), although
he subsequently (1993) modified this arrangement, defining
the following subfamilies: Alouattinae, Aotinae, Atelinae
(Ateles, Brachyteles, and Lagothrix), Callicebinae, Cebinae (in-
cluding Cebus and Saimiri, following Rosenberger 1981, see
below), and Pitheciinae (Cacajao, Chiropotes and Pithecia).

Rosenberger (1981) proposed an alternative classification
(Table 2), based on hypotheses concerning phylogenetic re-
lationships and a proposal for the scenario of their adaptive
radiation (see also, Rosenberger, 1980; Rosenberger and
Coimbra-Filho, 1984; Rosenberger and Strier, 1989;
Rosenberger et al., 1990). Rosenberger's (1981) scheme in-
cluded the use of tribes and subtribes (Table 2). It maintained
but redefined the use of the family name Cebidae to include
two subfamilies; the Cebinae (extant genera Cebus and Saimiri)
(see Harada et al., 1995) and Callitrichinae (extant genera

Common name
Black-headed marmoset

Marca's marmoset
Maues marmoset
Satere marmoset
Black-crowned dwarf marmoset
Manicore marmoset

Rio Acarf marmoset
Wied's black-tufted-ear marmoset

HernAndez-Camacho's black mantle

Callithrix mauesi Mittermeier, Schwarz & Ayres, 1992
Callithrix saterei Silva, Jr. & Noronha, 1998
Callithrix humilis Van Roosmalen, Van Roosmalen, Mittermeier & Fonseca, 1998
Callithrix manicorensis Van Roosmalen, Van Roosmalen, Mittermeier & Rylands, 2000
Callithrix acariensis Van Roosmalen, Van Roosmalen, Mittermeier & Rylands, 2000
Callithrix kuhlii Coimbra-Filho, 1985
Saguinus ./.s .' f,.'' hernandezi Hershkovitz, 1982

10. Leontopithecus caissara Lorini & Persson, 1990 Black-faced lion tamarin
11. Saimiri boliviensis peruviensis Hershkovitz, 1984 Peruvian squirrel monkey
12. Saimiri vanzolinii Ayres, 1981 Black-crowned squirrel monkey
13. Cebus olivaceus kaapori Queiroz, 1992 Kakapo capuchin
14. Aotus lemurinus brumbacki Hershkovitz, 1983 Brumback's night monkey
15. Aotus hershkovitzi Ramirez-Cerquera, 1983 Hershkovitz's night monkey
16. Aotus nancymaae Hershkovitz, 1983 Ma's night monkey
17. Callicebus barbarabrownae Hershkovitz, 1990 Northern Bahian blond titi
18. Callicebus coimbrai Kobayashi & Langguth, 1999 Coimbra's titi
19. Pithecia irrorata vanzolinii Hershkovitz, 1987 Vanzolini's bald-faced saki
20. Pithecia aequatorialis Hershkovitz, 1987 Equatorial saki
21. Chiropotes satanas utahicki Hershkovitz, 1985 Uta Hick's bearded saki
22. Cacajao calvus novaesi Hershkovitz, 1987 Novaes' bald-headed uacari

Neotropical Primates 8(2), June 2000 63

64 Neotropical Primates 8(2), June 2000

C .-' [includes CGbueIllaJ Saguinus, Leontopithecus, and
Cal/,I.co), and placed all other platyrrhines into a second
family, the Atelidae, which included two subfamilies: Atelinae
(Alouatta, Ateles, Brachyteles, Lagothrix) and Pitheciinae
(Pithecia, Chiropotes, Cacajao, Callicebus, Aotus). This classi-
fication was adopted by Kinzey (1997) in his review of the
evolution, ecology and behavior of the New World primates.

A molecular phylogeny, elaborated by Schneider et al. (1993,
1995, Schneider et al., 1993, 1995, 1996) (see below for a
summary of the molecular data), agreed in large part with
the classification of Rosenberger (1981), having only one major
difference; the alignment ofAotus with the Cebidae (subfamily
Aotinae) rather than the Pitheciinae. Tyler (1991) argued that
there are three well-defined clades among the extant platyr-
rhines: atelines, pitheciines, and callitrichids, but that evolu-
tionary relationships of the remaining genera (Aotus,
Callicebus, Cebus and Saimiri) was uncertain. Like Schneider
et al. (1993), Tyler (1991) argued that Aotus and Callicebus
are not closely related, but went even further, and, reviewing
morphological characters of the former, concluded that "Aotus
must be removed from other ceboids and made a sister group
of the entire Anthropoidea". Dutrillaux (1988) also argued
for the primitiveness of Aotus on the basis of its karyotype.
There is general consensus regarding the affinity of Callicebus
with the pitheciines (or pitheciids) (see Porter et al., 1999)
but the position of Aotus remains problematic. Tejedor (1998)
recently reviewed the morphological evidence for the phylo-
genetic placements of Callicebus and Aotus. He pointed out
that two Miocene genera from Patagonia, Homunculus and
Carlocebus show many primitive characters shared by both
Aotus and Callicebus (see also Kay, 1980; Rosenberger, 1981;
Ford, 1986, 1988; but see Rose and Fleagle, 1981;
Hershkovitz, 1970 who placed Homunculus in its own family
Homunculidae), and suggest they have a close common an-
cestor. Rosenberger in Schneider and Rosenberger (1996)
placed Callicebus and Aotus in the subtribe Homunculini with
the Pitheciinae. Gregory (1922), Miranda Ribeiro (1940),
Cabrera (1958), Simons (1972) and Fleagle (1988) placed
them in the same subfamily Aotinae. Fleagle (1999), how-
ever, taking note of Schneider et al. (1993) placed them in
separate subfamilies, as did Pocock (1925), Hill (1960),
Hershkovitz (1977) and Thorington and Anderson (1984).
Tejedor (1998) concluded that Aotus and Callicebus are an-
cient taxa and placed them in separate subfamilies within the
Atelidae but did not discount the possibility that Callicebus
belongs correctly in the subfamily Pitheciinae. In summary
the placements of Callicebus have included the subfamilies
Callicebinae, Cebinae and Pitheciinae. Aotus has been placed
in its own family Aotidae, in Atelidae and in Cebidae (sensu
Rosenberger, 1981), and in the subfamilies Aotinae and

The other differences between the schemes of Rosenberger
et al. (1990) and Schneider et al. (1993, 1995, 1996) are as
follows: 1) Schneider et al. place Saguinus in the Subtribe
Saguina rather than Leontocebina; 2) Schneider et al. sepa-
rate Leontopithecus from C .-( and Cebuella and place it

in its own Subtribe Leontopithecina; 3) Schneider et al. align
Callimico more closely with C .-'. and Cebuella and place
it in the Subtribe Callimiconina; 4) Schneider et al. align
Lagothrix with Brachyteles in the Subtribe Brachytelina,
whereas Rosenberger et al. place Lagothrix in the Subtribe
Atelina 5) Schneider et al. place Cacajao and Chiropotes in
the Subtribe Chiropotina separate from Pithecia; and 5) place
Callicebus in a separate Tribe rather than Subtribe of the

Hershkovitz (1977) placed Callimico in its own family;
Callimiconidae. Rosenberger (1981) argued for its alignment
as the subfamily Callimiconinae within the family Cebidae,
redefined as including also the subfamilies Cebinae (Cebus,
Saimiri) and Callitrichinae (Cebuella, Callithrix, Mico,
Saguinus, Leontopithecus) (see also Martin, 1992; Barroso,
1995; Pastorini et al., 1998; Canavez et al., 1999). Cronin
and Sarich (1978), Schneider et al. (1993, 1996), Pastorini
etal. (1998), Chaves etal. (1999) and Canavez etal. (1999a)
demonstrated that Callimico is more closely related to
C .-'. than it is to the tamarins.

Groves (in press) pointed out that the correct family name
for the night monkeys is Nictipithecidae Gray, 1870; Aotidae
Poche, 1908 being a junior synonym. Aotidae perhaps could
be maintained due its common usage, but this will require
clarification with the International Commission on Zoologi-
cal Nomenclature. Likewise, the correct name of the squirrel
monkey subfamily is Chrysotrichinae Cabrera, 1900; a se-
nior synonym of Saimirinae Miller, 1924 (Groves, in press).
As in the case of the night monkeys, we continue with the use
of Saimirinae for the squirrel monkeys because of its familiarity.

The placing of the seed-eating Cacajao, Pithecia and
Chiropotes, either as a subfamily of the Atelidae or in a sepa-
rate family is still under discussion. Although Schneider (in
Schneider and Rosenberger, 1996) argued for family status,
whereas Rosenberger in the same publication agreed to dif-
fer, placing it as a subfamily, Porter et al. (1999), with
Schneider as co-author, also argued for its subfamily status
following their study of the epsilon-globin gene.

We believe that the scheme of Schneider (in Schneider and
Rosenberger, 1996) represents the state of the art in
Platyrrhini classification since it is based on phylogenetic analy-
ses of morphological as well as molecular data. The precise
position of certain genera may need more information not
yet available but we can trust the general structure of that
proposal, and the species lists we present in the next section
follows the suprageneric classification of the above mentioned
authors (Table 2).


There are sixteen genera currently recognized for the
Platyrrhini. There is general consensus as to the validity of all
except for two. Since 1980, studies on the phylogenetic af-
finity of the pygmy marmoset, Cebuellapygmaea, to the Ama-

Neotropical Primates 8(2), June 2000 65

zonian marmosets (C .-'. ) have indicated that it could be
considered congeneric (for example, Rosenberger, 1981;
Barroso et al., 1997; Tagliaro et al., 1997; Porter et al.,
1997a). Here we maintain, however, the genus Cebuella, while
separating the Amazonian marmosets from the Atlantic for-
est marmosets, using C .-('. Erxleben, 1777 for the latter
and Mico Lesson, 1840 for the former (see section on Cebuella,

C. P. Groves (in press) has indicated that the yellow-tailed
woolly monkey, Lagothrix flavicauda, is sufficiently divergent
from other woolly monkeys as to warrant its own genus:
Oreonax Thomas, 1927. However, no DNA studies have yet
been carried out to support this view.

Species and Subspecies

The taxonomy within most of the platyrrhine genera is still
controversial, with discrepancies arising due to differing spe-
cies and subspecies concepts and to the characters used in
differentiating the forms. Tables 4-8 give a listing of the spe-
cies and subspecies. The lists do not reflect a taxonomic study
of our own, but an appraisal of the evidence available in the
literature mentioned in the text. We have taken, however,
the side of the splitterss" rather than the lumperss". This is
most evident in cases such as that of Saimiri, where current
evaluations have demonstrated the existence of five species
and 12 subspecies on the one hand (Hershkovitz, 1984,
1987b), and just two (or even one) species, on the other
(Costello et al., 1993). Alouatta is another case in point, where
new information is continuously being published and the tax-
onomy research is still in a state of flux (see Rylands and Bran-
don-Jones, 1998). Our aim is to provide an estimate of the
diversity of Neotropical primates, and while there is still dis-
cussion as to the validity or otherwise of numerous forms, we
prefer to maintain them since genetic evidence has confirmed
many formerly dubious forms and is demonstrating a large
hidden diversity. For conservation purposes it would be a se-
rious mistake to ignore the genetic and morphological diver-
sity of such wide-ranging genera as Saimiri. Even while split-
ting, it is highly probable that the list understates the diver-
sity of most Neotropical primate genera, most particularly in
such genera as Ateles, Alouatta, Cebus, and Lagothrix, which
have yet to receive modern systematic revisions, but also in,
for example C .-( for which new species have been de-
scribed recently, and more will undoubtedly be described in
the near future. Possible new forms of Lagothrix, Chiropotes,
Cacajao, Saguinus and C .-('. have been informally an-

The listings in Appendix 1 provide a total of five genera and
39 species with 61 species and subspecies of Callitrichidae,
11 species with 44 species and subspecies of Cebidae, 10 spe-
cies with 12 species and subspecies ofAotidae, 26 species with
44 species and subspecies of Pitheciidae, and 20 species with
45 species and subspecies of Atelidae. In all, we list 16 gen-
era, 107 species and 204 species and subspecies of Neotropi-
cal monkeys ranging from southern Mexico through to north-

ern Argentina and Paraguay, and possibly the northeastern
most tip of Uruguay (Villalba et al., 1995). Mittermeier
(1987), in his analysis of the diversity of Neotropical primates
and their distributions by country and the major phytogeo-
graphic regions of Gentry (1982), worked on the basis of
"some 65 species". The increase in species listed here comes
from both taxonomic revisions, which have raised subspecific
forms to full species and the descriptions of entirely new spe-
cies. The taxonomic revision of Callicebus by Hershkovitz
(1990) increased the number of species from just three to
13. All C .-'. forms are here listed as species (Rylands et
al., 1993), and such as Alouatta sara, Ateles chamek, A.
marginatus, and Brachyteles .'*,.... .-.'... are considered valid
species. Entirely new forms described recently include
C .-'. .. C. mauesi, C marcai, C saterei, C humilis,
C manicorensis and C. acariensis, Leontopithecus caissara, Aotus
hershkovitzi, Callicebus coimbrai and Cebus kaapori.

The Molecular Data

Molecular data suitable for cladistic analysis of the New World
monkeys were first obtained at the beginning of the 90's.
Their contribution to understanding the systematics at
supraspecific level and the phylogeny of New World primates
has been profound. Here we present a brief summary.

The first DNA dataset of aligned sequences of the nuclear
epsilon globin gene involving all 16 platyrrhine genera was
published by Schneider et al. (1993). Subsequently, a series
of datasets was generated, some including all extant New
World primate genera (Schneider et al., 1996; Canavez et
al., 1999a; von Dorman et al., 1999, Goodman et al., 1998),
while others concentrated on particular clades (Canavez et
al., 1999b, Chaves et al., 1999; Meireles et al., 1999; Porter
etal., 1997a, 1997b; Porter et al., 1999; Pastorini et al., 1998;
Tagliaro et al., 1999).

Table 3 summarizes the results involving four nuclear datasets
(IRBP, EPSILON, G6PD, and P-2M) for all extant New
World primates, and additional molecular data (vWF, up-
stream region EPSILON, y-Globin, ND4, and tRNA) includ-
ing specific clades. The four DNA datasets (EPSILON, IRBP,
G6PD and P-2M) were combined in tandem to perform a
total analysis using Maximum parsimony (MP) and Neigh-
bor-Joining (NJ) approaches using PAUP version 4.0b3a
(Swofford, 1998). MP and NJ trees (not shown) were almost
identical, differing only in the placement of Aotus. The MP
tree placed Aotus as the basal lineage of the callitrichine clade,
while NJ showed Aotus to occur in an unresolved trichotomy
with the Cebus-Saimiri clade and the callitrichines.

The four nuclear genes (P-2M, EPSILON, G6PD, and IRBP)
analyzed separately are not congruent with regard to the close
relationship among the three major clades (pitheciine, ateline,
callitrichine). EPSILON and P-2M support a pitheciine-
ateline clade, whereas G6PD groups the ateline with the
callitrichine clade, and IRBP shows the pitheciine clade to be
closer to the callitrichine clade. The topology resulting from

66 Neotropical Primates 8(2), June 2000

Table 3. DNA studies that elucidate the phylogeny of New World primates.
GENES Kb AUTHORS Pitheciine- Aotus, Cebus Callicebus as Lagothrix and Callitrichine dade
Ateline- and Saimiri as basal genus Brachyteles as Saguinus as Callimico and
Cebid closely linked to of the sister group in basal genus Callithrix as
relationship callitrichines pitheciine the ateline dade sister group
BS% BS% BS % BS % BS % BS %
EPSILON 1,8 Schneider et al., 1993 PA 79 94 99 86 77* 69
IRBP 1,9 Schneider etal. 1996 PC 96 99 100 77 100 84**
G6PD 1,2 Von Dornum & Ruvolo, AC 40 83 78 72 100 80
P-2M 1,7 Canavez et a, 1999a PA 72 56 87 96 100 92
VWF 0,7 Chaves etal., 1999 98 84
EPSILON and 5' region 1.8 to 4.3 Porter etal, 1997a PAC 100 100
EPSILON 5' region 1.2 -1.4 Porter etal., 1999 100
ND4, tRNAs 0,8 Pastorini etal., 1998 81* 80
Gamma-Globin 9,6 Meireles etal., 1999 100
BS: Bootstrap Support. Bootstrapping involves creating a new data set by sampling N characters randomly with replacement, so that the
resulting data set has the same size as the original, but some characters have been left out and others are duplicated. The random variation
of the results from analyzing these bootstrapped data sets can be shown statistically to be typical of the variation that you would get from
collecting new data sets (Felseinstein, 1985).
PA= Pitheciines x Atelines; PC= Pitheciines x Cebids; AC= Atelines x Cebids; PAC=unresolved trichotomy
*Saguinus and Leontopithecus as sister group
** Callimico and Leontopithecus as sister group of Callithrix

the total analysis, favors the ateline-pitheciine dade, but the
bootstrap support is not strong (78%) in the MP tree. In ad-
dition, the analysis of Porter et al. (1997b) using 5'upstream
region of epsilon gene in selected primate taxa showed the
three major clades in an unresolved trichotomy. The poor
resolution of the phylogenetic reconstruction of the three
major platyrrhine lineages, resulting from the most robust
molecular data set so far available for the New World mon-
keys strongly suggests that the pitheciines, atelines and cebids
emerged at almost the same time in the platyrrhine radiation.
These lineages are named by Schneider et al. (1996),
Schneider and Rosenberger (1996) and Goodman et al.
(1998), as the Families Atelidae, Pitheciidae and Cebidae.

In the Pitheciidae the four nuclear genes and the joint DNA
analysis significantly placed Callicebus as a sister group of the
traditional pitheciines, Pithecia, Chiropotes and Cacajao (boot-
strap values ranging from 78 to 100%). These results were
reinforced by the complementary study of Porter et al. (1999)
encompassing more than 4 Kb of the epsilon globin gene
and its upstream region.

The joint analysis strongly supports the monophyly of the
Cebidae, grouping Aotus, Cebus, and Saimiri with the small
callitrichines (bootstrap values from 83 to 100%). Addition-
ally, three of the four nuclear genes (IRBP, G6PD, P-2M),
and the epsilon upstream region of Porter et al. (1997a, 1999)
also indicate a close relationship between Cebus and Saimiri.
Among the callitrichines, additional molecular data of
Pastorini etal., (1998) using mitochondrial DNA and Chaves
et al. (1999), Canavez et al. (1999a, 1999b), using nuclear
DNA (intron 11 of von Willebrand, P-2-microglobulin
genes), strongly supported the reclassification of Cebuella as
belonging to the genus C .-'. (See also Porter et al.,
1997b), and also significantly placed Callimico as a sister
group of Callithrix, contrary to the traditional view of
Callimico as being basal to the callitrichines. Five of the six
genes, and the joint analysis placed Saguinus as the basal ge-
nus of the callitrichine tree with significant bootstrap values

(98-100%). Finally, four nuclear genes and the joint analysis
showed a congruent branching pattern in the atelid clade,
placing Alouatta as the most basal lineage and Brachyteles from
the Atlantic forest) and Lagothrix (from the Amazonian for-
est), as sharing the last common ancestor (bootstrap values
ranging from 78 to 100%), contrasting to the widely dissemi-
nated view that Ateles and Brachyteles share the last common
ancestor in the ateline clade. Furthermore, as shown in Table
3, the investigation by Meireles et al. (1999), encompassing
more than 9 Kb of the gamma globin gene complex strongly
supports the Lagothrix-Brachyteles clade (bootstrap value of

In summary, three evolutionary lineages are strongly estab-
lished for the New World monkeys: atelines, pitheciines, and
cebids. However, two major points regarding the branching
patterns of the most ancient lineages remain to be clarified:
(i) what is the exact branching pattern ofAotus, Cebus, Saimiri
and the small callitrichines? and (ii) which two of these three
lineages, pitheciines, atelines or cebids, are more closely re-

All the platyrrhine genera require further investigation at the
subgeneric level. Preliminary estimates indicate that, in the
majority, speciation seems to have occurred in the Pliocene-
Pleistocene scenario (5 Ma or less). However, because the in-
ternodes are usually very small when species and populations
are compared, even using highly variable mitochondrial DNA,
a "all-evidence" approach, using geomorphology, biogeogra-
phy, traditional morphology, behavior and cytogenetics, in
association with DNA sequencing, is required to better un-
derstand the recent evolutionary processes which have resulted
in the species diversity of New World monkeys existing today
(see, for example, Boinski and Cropp [1999], who used this
approach to examine the systematics of squirrel monkeys).

Neotropical Primates 8(2), June 2000 67

Table 4. Species and subspecies of the Family Callitrichidae.
Family Callitrichidae
1. Cebuella pygmaea pygmaea (Spix, 1823)
2. C pygmaea niveiventris Lonnberg, 1940
3. Mico argentatus (Linnaeus, 1766)
4. M. leucippe (Thomas, 1922)
5. M. melanurus (E. Geoffroy in Humboldt, 1812)
6. M. intermedius (Hershkovitz, 1977)
7. M. emiliae (Thomas, 1920)
8. M. nigriceps (Ferrari & Lopes, 1992)
9. M. marcai (Alperin, 1993)
10. M. humeralifer (E. Geoffroy in Humboldt, 1812)
11. M. chrysoleucus (Wagner, 1842)
12. M. mauesi (Mittermeier, Schwarz & Ayres, 1992)
13. M. humilis (Van Roosmalen, Van Roosmalen, Mittermeier & Fonseca, 1998)
14. M. saterei (Silva, Jr. & Noronha, 1998)
15. M. manicorensis (Van Roosmalen, Van Roosmalen, Mittermeier & Rylands, 2000)
16. M. acariensis. (Van Roosmalen, Van Roosmalen, Mittermeier & Rylands, 2000)
17. Callithrixjacchus (Linnaeus, 1758)
18. C penicillata (t. Geoffroy,1812)
19. C kuhlii Coimbra-Filho, 1985
20. C (E. Geoffroy in Humboldt, 1812)
21. C aurita (E. Geoffroy in Humboldt, 1812)
22. C. flaviceps (Thomas, 1903)
23. Saguinus ,...,,....',,. ......, (Spix, 1823)
24. S. .., ... hernandezi Hershkovitz, 1982
25. S. U,... (Jimenez de la Espada, 1870)
26. S. fuscicollisfuscicollis (Spix, 1823)
27. S. fuscicollis fuscus (Lesson, 1840)
28. S. fuscicollis avilapiresi Hershkovitz, 1966
29. S. fuscicollis cruzlimai Hershkovitz ,1966
30. S. fuscicollis leucogenys (Gray, 1866)
31. S. fuscicollis lagonotus (Jimenez de la Espada, 1870)
32. S. fuscicollis primitivus Hershkovitz, 1977
33. S. fuscicollis (Pucheran, 1845)
34. S. fuscicollis nigrifrons (I. Geoffroy, 1850)
35. S. fuscicollis weddelli (Deville, 1849)
36. S. fuscicollis melanoleucus (Miranda Ribeiro, 1912)
37. S. fuscicollis crandalli Hershkovitz, 1966
38. S. tripartitus (Milne-Edwards, 1878)
39. S. mystax mystax (Spix, 1823)
40. S. mystax pileatus (I. Geoffroy & Deville, 1848)
41. S. mystax pluto (Lonnberg, 1926)
42. S. labiatus labiatus (E. Geoffroy in Humboldt,1812)
43. S. labiatus thomasi (Goeldi, 1907)
44. S. labiatus rufiventer (Gray, 1843)
45. S. imperator imperator (Goeldi, 1907)
46. S. imperatorsubgrisescens (Lonnberg, 1940)
47. S. midas (Linnaeus, 1758)
48. S. ,.i., lE. Geoffroy, 1803)
49. S. inustus (Schwarz, 1951)
50. S. bicolor (Spix, 1823)
51. S. martinsi martinsi (Thomas, 1912)
52. S. martinsi ochraceus Hershkovitz, 1966
53. S. leucopus (Giinther, 1877)
54. S. oedipus (Linnaeus, 1758)
55. S. i" *(Pucheran, 1845)
56. Leontopithecus rosalia (Linnaeus, 1766)
57. L. chrysomelas (Kuhl, 1820)
58. L. chrysopygus (Mikan, 1823)
59. L. caissara Lorini & Persson, 1990
60. Callimico goeldii (Thomas, 1904)

Common name (following Hershkovitz, 1977)
Pygmy marmoset

Black-chinned emperor tamarin
Bearded emperor tamarin
Golden-handed tamarin
Black-handed tamarin
Mottled-face tamarin
Pied bare-face tamarin
Martin's bare-face tamarin
Ochraceous bare-face tamarin
Silvery-brown bare-face tamarin
Cotton-top tamarin
Geoffroy's tamarin
Golden lion tamarin
Golden-headed lion tamarin
Black lion tamarin
Black-faced lion tamarin
Goeldi's monkey

Silvery marmoset
Golden-white bare-ear marmoset
Black-tailed marmoset
Aripuana marmoset
Snethlage's marmoset
Black-headed marmoset
Marca's marmoset
Black and white tassel-ear marmoset
Golden-white tassel-ear marmoset
Mauds marmoset
Black-crowned dwarf marmoset
Saterd marmoset
Manicord marmoset
Rio Acarf marmoset
Common marmoset
Black-tufted-ear marmoset
Wied's black-tufted-ear marmoset
Geoffroy's tufted-ear marmoset
Buffy-tufted-ear marmoset
Buffy-headed marmoset
Spix's black mantle tamarin
Hernindez-Camacho's black mantle tamarin
Graell's black-mantle tamarin
Spix's saddle-back tamarin
Lesson's saddle-back tamarin
Avila Pires' saddle-back tamarin
Cruz Lima's saddle-back tamarin
Andean saddle-back tamarin
Red-mantle saddle-back tamarin
Saddle-back tamarin
Illiger's saddle-back tamarin
Geoffroy's saddle-back tamarin
Weddell's saddle-back tamarin
White saddle-back tamarin
Crandall's saddle-back tamarin
Golden-mantle saddle-back tamarin
Spix's mustached tamarin
Red-cap mustached tamarin
White-rump mustached tamarin
Red-bellied tamarin
Thomas' mustached tamarin

68 Neotropical Primates 8(2), June 2000

Table 5. Species and subspecies of the Family Cebidae.
Family Cebidae Common name
1. S. oerstedii oerstedi (Reinhardt, 1872) Black-crowned Central American squirrel monkey
2. S. oerstedii citrinellus Thomas, 1904 Grey-crowned Central American squirrel monkey
3. Saimiri boliviensis boliviensis (I. Geoffroy & de Bolivian squirrel monkey
Blainville, 1834)
4. S. boliviensis peruviensis Hershkovitz, 1984 Peruvian squirrel monkey
5. S. boliviensis pluvialis Lonnberg, 1940
6. S. boliviensis jaburuensis Lonnberg, 1940
7. S. vanzolinii Ayres, 1981 Black-headed squirrel monkey
8. S. sciureus sciureus (Linnaeus, 1758) Common squirrel monkey
9. S. sciureus ,:;,,..*,, (Von Pusch, 1941)
10. S. sciureus cassiguiarensis (Lesson, 1840) Humboldt's squirrel monkey
11. S. sciureus macrodon (Elliot, 1907) Ecuadorian squirrel monkey
12. S. ustus I. Geoffroy, 1843 Golden-backed squirrel monkey
13. Cebus apella apella (Linnaeus, 1758) Guianan brown capuchin
14. C. apella fatuellus (Linnaeus, 1766)
15. C apella macrocephalus Spix, 1823 Large-headed capuchin
16. C apella peruanus Thomas, 1901 Peruvian tufted capuchin
17. C apella tocantinus Lonnberg, 1939
18. C apella margaritae Hollister, 1914 Margarita Island capuchin
19. C libidinosus libidinosus Spix, 1823 Bearded capuchin
20. C libidinosus pallidus (Gray, 1866) Tambopata tufted capuchin, pale capuchin
21. C libidinosus paraguayanus Fischer, 1829 Paraguayan tufted capuchin
22. C libidinosus juruanus Lonnberg, 1939
23. C nigritus nigritus (Goldfuss, 1809) Black-horned capuchin
24. C nigritus robustus Kuhl, 1820 Crested capuchin
25. C nigritus cucullatus Spix, 1823
26. C xanthosternos Wied-Neuwied, 1826 Yellow-breasted capuchin
27. C ,:./',,.,. ,:.'.,., ,:' (Humboldt, 1812) White-fronted capuchin
28. C :. ,,/'..'' adustus Hershkovitz, 1949 Brown-faced capuchin
29. C :. ",.' aeguatorialis Allen, 1914 Ecuadorian capuchin
30. C ,:.,'J,,,.,' cesarae Hershkovitz, 1949
31. C ,:.'., ,."'. cuscinus Thomas, 1901 Shock-headed capuchin
32. C ,:././,.,*. malitiosus Elliot, 1909
33. C :. ,",.'.' trinitatis Von Pusch, 1941 Trinidad white-fronted capuchin
34. C ,:.!'.f,,,.. versicolor Pucheran, 1845 Varied capuchin
35. C. a.!', ..,, yuracus Hershkovitz, 1949 Andean white-fronted capuchin
36. C ,:.'.,,,.'. leucocephalus Gray, 1865
37. C capucinus capucinus (Linnaeus, 1758) White-faced capuchin
38. C capucinus limitaneus Hollister, 1914
39. C capucinus imitator Thomas, 1903 Panamanian white-throated capuchin
40. C capucinus curtus Bangs, 1905 Gorgona white-fronted capuchin
41. C olivaceus olivaceus Schomburgk, 1848 Wedge-capped capuchin
42. C olivaceus apiculatus Hershkovitz, 1949
43. C olivaceus brunneus Allen, 1914 Brown weeper capuchin
44. C. olivaceus castaneus I. Geoffroy, 1851 Chestnut capuchin
45. C olivaceus kaapori Queiroz, 1992 Ka'apor capuchin

Neotropical Primates 8(2), June 2000 69

Table 6. Species and subspecies of the Family Aotidae.

Family Aotidae
1. Aotus lemurinus lemurinus I. Geoffroy, 1846
2. A. lemurinus griseimembra Elliot, 1912
3. A. lemurinus zonalis Goldman, 1914
4. A. lemurinus brumbacki Hershkovitz ,1983
5. A. hershkovitzi Ramirez-Cerquera, 1983
6. A. trivirgatus (Humboldt, 1812)
7. A. vociferans (Spix, 1823)
8. A. miconax Thomas, 1927
9. A. nancymaae Hershkovitz, 1983
10. A. nigriceps Dollman, 1909
11. A. azarai azarai (Humboldt, 1812)
12. A. azarai boliviensis Elliot, 1907
13. A. azarai infulatus (Kuhl, 1820)

Common name
Colombian or lemurine night monkey
Grev-leaeed niaht monkey

Brumback's night monkey
Hershkovitz's night monkey
Douroucouli, owl monkey, night monkey

Andean night monkey
Ma's night monkey
Black-headed or Peruvian night monkey
Azara's night monkey
Bolivian night monkey
Feline night monkey

Table 7. Species and subspecies of the Family Pitheciidae.

Family Pitheciidae
1. ( ...... ,, modestus Lonnberg, 1939
2. C. donacophilus (D'Orbigny, 1836)
3. C. pallescens Thomas, 1907
4. C. olallae L6nnberg, 1939
5. C. oenanthe Thomas, 1924
6. C. cinerascens (Spix, 1823)
7. C .- 'Thomas, 1908
8. C. baptista Lonnberg, 1939
9. C moloch (H..f--.. 1.1. ..--. 1807)
10. C. brunneus (Wagner, 1842)
11. C. cupreus (Spix, 1823)
12. C. ornatus (Gray, 1866)
13. C. medemi Hershkovitz, 1963
14. C. torquatus torquatus (H. .f,.i. ,1.. ... 1807)
15. C. torquatus lugens (Humboldt, 1811)
16. C. torquatus lucifer Thomas, 1914
17. C. torquatus purinus Thomas, 1927
18. C. torquatus regulus Thomas, 1927
19. C. personatus (E. Geoffroy, 1812)
20. C. nigrifrons (Spix, 1823)
21. C. melanochir Wied-Neuwied, 1820
22. C. barbarabrownae Hershkovitz, 1990
23. C. coimbrai Kobayashi and Langguth, 1999
24. Pithecia pithecia pithecia (Linnaeus, 1758)
25. P. pithecia chrysocephala I. Geoffroy, 1850
26. P. monachus monachus (E. Geoffroy, 1812)
27. P. monachus miller Allen, 1914
28. P. monachus napensis Lonnberg, 1938
29. P. irrorata irrorata Gray, 1842
30. P. irrorata vanzolinii Hershkovitz, 1987
31. P. albicans Gray, 1860
32. P. aeguatorialis Hershkovitz, 1987
33. Chiropotes albinasus (I. Geoffroy & Deville, 1848)
34. C. satanas satanas (Hoffmannsegg, 1807
35. C. satanas chiropotes (Humboldt, 1811)
36. C. satanas utahicki Hershkovitz, 1985
37. Cacajao calvus calvus (I. Geoffroy, 1847)
38. C. calvus ucayalii (Thomas, 1928)
39. C. calvus novaesi Hershkovitz, 1987
40. C. calvus rubicundus (I. Geoffroy and Deville, 1848)
41. C. melanocephalus melanocephalus (Humboldt, 1811)
42. C. melanocephalus ouakary (Spix, 1823)

Common name

Beni titi monkey
Andean titi monkey

Hoffmann's titi monkey

Ornate titi monkey

Collared titi, widow monkey

Widow monkey

Northern masked titi
Black-fronted titi
Southern Bahian masked titi
Northern Bahian blond titi
Coimbra's titi
White-faced saki
Golden-faced saki
Geoffroy's monk saki
Miller's monk saki
Napo monk saki
Gray's bald faced saki
Vanzolini's bald-faced saki
White saki, buffy saki
Equatorial saki
White-nosed bearded saki
Bearded saki
Black saki
Uta Hick's bearded saki
White bald-headed uacari
Ucayali bald-headed uacari
Novaes' bald-headed uacari
Red bald-headed uacari
Humboldt's black-headed uacari
Spix's black-headed uacari

70 Neotropical Primates 8(2), June 2000

Table 8. Species and subspecies of the Family Atelidae.

Family Atelidae
1. Alouatta seniculus seniculus (Linnaeus, 1766)
2. A. seniculus insulanus Elliot, 1910
3. A. seniculus ssp. (formerly straminea 1)
4. A. seniculus amazonica Lonnberg, 1941
5. A. seniculus juara Elliot 1910
6. A. seniculus puruensis Lonnberg, 1941
7. A. seniculus arctoidea Cabrera, 1940
8. A. sara Elliot, 1910
9. A. nigerrima Lonnberg, 1941
10. A. belzebul belzebul (Linnaeus, 1766)
11. A. belzebul discolor (Spix, 1823)
12. A. belzebul ululata Elliot, 1912
13. A. guariba guariba (Humboldt, 1812)
14. A. guariba clamitans Cabrera, 1940
15. A. palliata palliata (Gray, 1849)
16. A. palliata mexicana (Merriam 1902)
17. A. palliata aequatorialis (Festa, 1903)

18. A. coibensis coibensis Thomas, 1902
19. A. coibensis trabeata Lawrenc,e 1933
20. A. caraya (Humboldt,1812)
21. A. pigra Lawrence, 1933
22. Ateles , i'Kuhl, 1820
23. A. i azuerensis (Bole, 1937)
24. A. i frontatus (Gray, 1842)
25. A. i grisescens Gray, 1866
26. A. 'i panamensis Kellogg & Goldman, 1944
27. A. i ornatus (Gray, 1870)
28. A. i vellerosus (Gray, 1866)
29. A. i yucatanensis Kellogg & Goldman, 1944
30. A. i fusciceps Gray, 1866
31. A. rufiventris Allen, 1914
32. A. chamek (Humboldt, 1812)
33. A. paniscus (Linnaeus, 1758)
34. A. marginatus (E. Geoffroy, 1809)
35. A. belzebuth (E. Geoffroy, 1806)
36. A. hybridus hybridus (I. Geoffroy, 1829)
37. A. hybridus brunneus Gray, 1872
38. Lagothrix lagothricha (Humboldt, 1812)
39. L. cana cana (E. Geoffroy in Humboldt, 1812)
40. L. cana tschudii Pucheran, 1857
41. L. poeppigii Schinz, 1844
42. L. lugens Elliot, 1907
43. Oreonax flavicauda (Humboldt, 1812)
44. Brachyteles arachnoides (E. Geoffroy, 1806)
45. B. ', .' (Kuhl, 1820)

Common name
Red howling monkey
Trinidad howling monkey
Guianan red howling monkey

Ursine howling monkey
Bolivian red howling monkey
Black howling monkey
Red-handed howling monkey

Northern brown howling monkey
Southern brown howling monkey
Golden-mantled howling monkey
Mexican howling monkey
South Pacific blackish howling monkey,
Ecuadorian mantled howling monkey
Coiba Island howling monkey
Azuero howling monkey
Black howling monkey
Black howling monkey
Geoffroy's spider monkey
Azuero spider monkey
Black-browed spider monkey
Hooded spider monkey

Red spider monkey
Ornate spider monkey
Mexican spider monkey
Yucatin spider monkey
Brown-headed spider monkey
Colombian black spider monkey
Black-faced black spider monkey
Red-faced black spider monkey
White-whiskered spider monkey
White-bellied spider monkey
Variegated spider monkey
Brown spider monkey
Humboldt's woolly monkey
Geoffroy's woolly monkey

Poeppig's woolly monkey
Colombian woolly monkey
Yellow-tailed woolly monkey
Southern muriqui
Northern muriqui

Neotropical Primates 8(2), June 2000 71

Table 9. A summary table of the New World primates by genus.

Family Callitrichidae Species Taxa
Cebuella 1 2
Mico 14 14

Family Cebidae

15 33

Saimiri 5 12
Cebus 7 33
Subtotal 12 45
Family Aotidae
Aotus 8 13
Family Pitheciidae
( a,:.... "19 23
Pithecia 5 9
Chiropotes 2 4
Cacajao 2 6
Subtotal 28 42
Family Atelidae
Alouatta 8 21
Ateles 6 16
Lagothrix 4 5
Oreonax 1 1
Brachyteles 2 2
Subtotal 21 45
Total Neotropical 110 205

CEBUELLA Gray, 1886

Cebuella pygmaea pygmaea (Spix, 1823)
C. pygmaea niveiventris Lbnnberg, 1940

Cebuella was proposed by Gray (1866) as a subgenus of Hapale
(later C .-'' ( and soon after as a distinct genus (Gray,
1870). Rosenberger (1981), Rosenberger and Coimbra-Filho
(1984), and Natori (1994) argued, on morphological terms,
that the pygmy marmoset should correctly be included in
the genus C .-'. This was not maintained in Rosenberger
et al. (1990), although Barroso (1995), Barroso et al. (1997),
Moreira (1996), Schneider et al. (1996), Schneider and
Rosenberger (1996), Tagliaro et al. (1997), Porter et al.
(1997a), and Canavez et al. (1999a) also argued for this ar-
rangement on the basis of molecular genetics. The 1997 AZA
studbook for the pygmy marmoset placed it in the genus
C .-'. (Arden, 1998). The argument that Cebuella should
be included in the genus C .-'. centers on the conclu-
sion, from both morphological and genetic studies, that the
pygmy marmoset is more closely related to the Amazonian
marmosets (the argentata group of Hershkovitz [1977]) than
the latter are to the Atlantic forest marmosets (the jacchus
group of Hershkovitz [1977]). Schneider et al. (1993) and
Schneider and Rosenberger (1996), however, also concluded
that their molecular genetic data are compatible with jacchus
and pygmaea being congeneric. Although closely related to
the Amazonian marmosets, we believe that Cebuellapygmaea
deserves generic status. The species shows very distinct fea-

C .-.'. jacchus (Linnaeus, 1758)
C penicillata (E. Geoffroy, 1812)
C kuhlii Coimbra-Filho, 1985
C geoffroyi (E. Geoffroy in Humboldt, 1812)
C aurita (E. Geoffroy in Humboldt, 1812)
C. flaviceps (Thomas, 1903)

Mittermeier and Coimbra-Filho (1981) maintained that the
marmosets comprising Hershkovitz's (1975) "Jacchus" group,
should be considered good species rather than subspecies of
C jacchus, even though fertile hybrids had been produced in
captivity (Hill, 1957; Coimbra-Filho, 1970, 1971, 1973,
1974, 1978; Mallinson, 1971; Hampton et al., 1971;
Coimbra-Filho and Mittermeier, 1973b; Coimbra-Filho et
al., 1993b), and Hershkovitz (1975, 1977) and Alonso et al.
(1987) had provided evidence of intergradation in the wild.

tures, unique among the callitrichids, involving not only its
diminutive size but also the tree-gouging specialization not
observed to nearly the same extent in its Amazonian sister

If Cebuella is maintained as a separate genus, then the group
would be paraphyletic unless the argentata group marmosets
and thejacchus group marmosets are placed in different gen-
era. This was the option adopted, with the Amazonian mar-
mosets being placed in the genus Mico Lesson, 1840 (see be-
low). This decision was also based on the clear identity of the
three genera shown in all morphological and molecular stud-
ies available.

In 1940, Lbnnberg described the subspecies Cebuella
pygmaea niveiventris from Lago Ipixuna, south bank of the
Rio Solim6es, based on its sharply contrasting whitish chest,
belly, and inner surface of arms and legs. The ventral surface
of C. p. pygmaea is ochraceous. Cruz Lima (1945) and Napier
(1976) also recognized and described the two subspecies.
Cebuella pygmaea pygmaea (Spix, 1823) occurs in the state of
Amazonas, Brazil, north of the Rio Solim6es and south of the
Rio Japuri, southern Colombia north of the Rios Maranon
and Putumayo and south of the Rio CaquetA (Japuri), east-
ern Ecuador, and eastern Peru south of Rio Putumayo, north
of the Rio Maran6n and east of the Rio Pastaza, whereas
Cebuellapygmaea niveiventris Lbnnberg, 1940 occurs in east-
ern Peru south of the Rfo Maran6n and east of the Rfo
Huallaga, and in the state of Amazonas, Brazil, south of the
Rio Solim6es and west of the Rio Purus. It occurs as far south
as the Rio Muyumanu in the Pando of northern Bolivia
(Buchanan-Smith et al., 2000). Hershkovitz (1977) argued
that the color of the underparts is individually and locally
variable and not justifying the subspecific status of niveiventris.
Van Roosmalen andVan Roosmalen (1997), however, recently
presented evidence for the validity of the two subspecies based
on field observations and captive specimens of C. p.
niveiventris, and confirmed its presence in the interfluvium
of the Rios Purus and Madeira.

CALLITHRIX Erxleben, 1777

72 Neotropical Primates 8(2), June 2000

However, Mittermeier and Coimbra-Filho (1981) argued that
the issue of species or subspecies designations for the non-
Amazonian marmosets was controversial and depended on
the resolution of three questions: 1) Do the forms naturally
overlap in their ranges without interbreeding? 2) What is the
correct taxonomic interpretation of the intergrades reported
by Hershkovitz (1975, 1977), considering they might be
merely individual or ontogenetic variants rather than hybrids?
and 3) Presuming natural zones of intergradation do exist,
are they regions of broad clinal variation or narrow contact
zones? Mittermeier et al. (1988, p.21) provided answers to
these questions, which reinforced the argument that at least
the forms aurita, geoffroyi, penicillata, jacchus and kuhlii
should be considered valid species flavicepss may be subspe-
cific to aurita, see below), even though it would seem that
none of the "Jacchus" group marmosets overlap in their geo-
graphic distributions without interbreeding. Evidence is now
available for a number of natural hybrid zones either at the
distributional limits of the various forms or due to introduc-
tions. They are reviewed by Coimbra-Filho et al. (1993b),
who considered hybridization at natural distributional limits
(C penicillata x C. geoffroyi, C. penicillata x C. kuhlii, C.
geoffroyi x C. flaviceps, C. aurita x C. flaviceps) and where one
(or both) forms had been introduced (C. jacchus x C.

The important feature is that, in all cases, the documented
contact zones are narrow or confined and clinal variation is
not evident (Vivo, 1991; Coimbra-Filho et al., 1993b). Vivo
(1991) classified all the "Jacchus" group marmosets as species
(except for kuhlii which he did not recognize as distinct from
C penicillata), arguing that allopatry or parapatry alone can-
not be used to determine subspecific or specific status and
that there is no evidence for widespread intergradation or
clinal variation, and protesting that the use of subspecific clas-
sifications based not on spatial distribution of characters but
merely on the degree of similarity in pelage between forms is
inadequate. On the basis of pelage color and patterns alone,
Rosenberger (1984) also argued that they should be consid-
ered species rather than subspecies, but that more informa-
tion is needed from other systems, genetic and morphological.

The authorship of the name C .-' kuhlii is the subject
of some debate. Hershkovitz (1975, p.142) was the first to
indicate that Wied-Neuwied (1826) had referred to the mar-
moset of south-east Bahia as "Hapalepenicillata Kuhlii" (sic).
However, Hershkovitz (1975, 1977) argued at length that
kuhlii was not a valid taxon, being merely an intergrade
between C. j. penicillata and C. j. geoffroyi. Vivo (1991,
pp.80-81), on the other hand, argued that Wied-Neuwied
(1826) had not intentionally given it this name, merely, and
incorrectly, ascribing the authorship of the name penicillata
to Kuhl. The first person to intentionally use the name kuhlii
to describe the marmosets from south-east Bahia was
Hershkovitz (1975), but his argument that it was not a valid
taxonomic entity disqualifies the possibility of him being
attributed authorship. This is therefore given to Coimbra-
Filho (1985).

Groves (1989, 1993) listed the "Jacchus" group marmosets
as separate species (see also Marroig, 1995). Natori (1986,
1990) and Natori and Shigehara (1992) in their studies of
the dental morphology, and Natori (1994) in his
craniometrical study, also argued for their ranking as species,
based however on compliance with the separation of C.
argentata and C humeralifera as distinct species. Natori (1986)
examined six dental characters and tooth size in C
On the basis of molar tooth size alone, he found that the dif-
ference between the "Jacchus" group marmosets was greater
than between the Amazonian C argentata and C humeralifer.
He argued that if the latter are to be considered separate spe-
cies, then so should the "Jacchus" group marmosets. The same
conclusion was drawn by Natori (1994) in his study of 19
cranial measures. On the basis of Q-mode correlation of these
measures, the distances between the "Jacchus" group mem-
bers were greater than those between the Amazonian
argentata and humeralifer, and, excepting C jacchus and C
penicillata, were greater than between Cebuella pygmaea and
argentata and between Cebuella and humeralifer.

Cytogenetics and molecular genetics have to date been inde-
cisive in their contribution to the debate concerning the taxo-
nomic status of the "Jacchus" group marmosets. Peixoto
(1976) and Peixoto and Pedreira (1982) compared the chro-
mosomes of C jacchus, C. penicillata and C geoffroyi and re-
corded clear differences in G-banding indicating paracentric
inversions not found in later studies by Seuinez et al. (1988)
and Nagamachi (1995). Nagamachi (1995) and Nagamachi
et al. (1997) carried out a study of the chromosome mor-
phology of C. kuhlii and the other "Jacchus" group marmo-
sets except C. flaviceps. All of the eastern Brazilian marmosets
have a diploid chromosome number of 46, with 30 two-
armed and 14 acrocentric autosomes, a conservative submeta-
centric X chromosome, and a Y chromosome which is highly
variable in size and morphology. In C. kuhlii the Y chromo-
some is small and two-armed metacentricc). An analysis of
the G-banding patterns demonstrated a lack of any chromo-
somal rearrangements to differentiate their karyotypes. C-
banding, likewise, demonstrated no differences between the
species. Heterochromatin was found to occur in small quan-
tities in the centromeric regions of all the chromosomes, in
the telomeric region of the short arm of pair 6 and the
telomeric region of the long arm of chromosome 22. Ag-NOR
staining marked secondary constrictions of the small arms of
the acrocentric chromosomes. Nagamachi (1995) and
Nagamachi et al. (1997) concluded that the five species they
studied were extremely homogeneous in their karyotypes, ex-
cept for the size and morphology of the Y chromosome, which
in the case of C. jacchus was variable even between popula-
tions. They concluded that nothing can be said as a result
concerning the taxonomic status of each species.

Tagliaro et al. (1997) analyzed mitochondrial DNA control
region sequences in all the 'Jacchus" group marmosets, except
for C. flaviceps. In reconstructing the phylogeny of these
marmosets from their findings, they concluded that "Our trees

Neotropical Primates 8(2), June 2000 73

certainly do not come down in strong support of a
monophyletic C kuhlii, although their paraphyly is also only
weakly supported (i.e. a monophyletic C kuhlii adds only
one substitution to the MP tree)" (p.682), and later (p.683):
"our data do not support a clear taxonomic distinction
between C. kuhlii, C. penicillata, and possibly C. jacchus,
which [...] we regard as a tentative proposal but one that needs
to be further explored...". They found, on the contrary, strong
support that both C. aurita and C geoffroyi are "distinct
evolutionary entities". Genetic studies by Meireles et al.
(1992) concluded that M. humeralifer and M. emiliae are
subspecies of M. argentatus; and that C jacchus, C penicillata,
and C geoffroyi are subspecies; but that C geoffroyi is the most
primitive of the "jacchus" Group (most similar genetically to
M. argentatus), which should, therefore, be referred to as the
"geoffroyi" Group, with C. jacchus and C. penicillata as
subspecific to C geoffroyi. Meireles et al. (1998) concluded
that the forms kuhlii and penicillata are conspecific.

The primitiveness (Hershkovitz, 1977) of C geoffroyi was also
advocated by Natori (1986, 1994) and Natori and Shigehara
(1992), and agrees with the low degree of morphological ad-
aptation for tree-gouging observed in this species. Dental and
behavioral adaptations for tree-gouging are most advanced
in C jacchus and C penicillata.

Coimbra-Filho (1984, 1985, 1990), Rylands (1989b),
Mittermeier et al. (1988), Natori (1990), Rylands et al.
(1993), and Groves (1993, in press) recognize as a valid form.
Hershkovitz (1977) regarded C .-'. kuhlii from south-
ern Bahia to be a hybrid between C. j. geoffroyi and C. j.
penicillata. Vivo (1991) considered it to be indistinguishable
from C penicillata from northern Minas Gerais. Coimbra-
Filho (1984, 1985, 1990), Rylands (1989b), Mittermeier et
al. (1988), Natori (1990), Rylands et al. (1993), and Groves
(1993, in press) recognize it as a valid species. Groves (1993)
and Eisenberg and Redford (1999) gave the distribution as
Bolivia evidently a typographic error.

Coimbra-Filho (1986a, 1986b, 1990) and Coimbra-Filho et
al. (1991) argued that Callithrixflaviceps should be
considered subspecific to C aurita. The similarities between
C flaviceps and C aurita in pelage patterns such as the ear
tufts and the skull-like facial mask, ecological adaptations,
ontogeny, (the infants are extremely similar), vocalizations and
clinal variation in overall pelage color, indicate to us that
flaviceps might well be best considered a subspecies of aurita
(Coimbra-Filho, 1986a, 1986b; Coimbra-Filho et al.,

The following publications list all marmosets as species:
Mittermeier et al. (1992a), Vivo (1985, 1991), and Groves
(1993, in press).

MICO Lesson, 1840

1. Mico argentatus (Linnaeus, 1766)
2. M. leucippe (Thomas, 1922)

3. M. melanurus (E. Geoffroy in Humboldt, 1812)
4. M. intermedius (Hershkovitz, 1977)
5. M. emiliae (Thomas, 1920)
6. M. nigriceps (Ferrari & Lopes, 1992)
7. M. marcai (Alperin, 1993)
8. M. humeralifer (E. Geoffroy in Humboldt, 1812)
9. M. chrysoleucus (Wagner, 1842)
10.M. mauesi (Mittermeier, Schwarz & Ayres, 1992)
11.M. humilis (Van Roosmalen, Van Roosmalen,
Mittermeier & Fonseca, 1998)
12.M. saterei (Silva, Jr. & Noronha, 1998)
13.M. manicorensis (Van Roosmalen, Van Roosmalen,
Mittermeier & Rylands, 2000)
14.M. acariensis (Van Roosmalen, Van Roosmalen,
Mittermeier & Rylands, 2000)

As discussed in the section dealing with Cebuella, the phylo-
genetic closeness of the pygmy marmoset to the other Ama-
zonian marmosets has indicated that they could be consid-
ered to be of the same genus. The alternative, to recognize
the distinctiveness of pygmaea and avoid paraphyletic genera,
is to place the Amazonian and Atlantic forest marmosets into
separate genera. The oldest generic name applicable to the
Amazonian marmosets alone is Mico Lesson, 1840 (type spe-
cies Mico argentatus). The name Mico is used as a subgenus
for the Amazonian marmosets by Groves (in press).

Coimbra-Filho (1990) argued that the three subspecies of
Mico humeralifer listed by Hershkovitz (1977) (C .-A' h.
humeralifer, C h. chrysoleuca, and C h. intermedius) should
be considered species, following De Boer (1974) and Vivo
(1991). Mico intermedius was described as an intermediate
color form of M. h. humeralifer and M. h. chrysoleuca by
Hershkovitz (1977). Pelage color and pattern, and its geo-
graphic distribution, however, would place it as a subspecies
of M. argentatus if, following Hershkovitz (1977), the Ama-
zonian marmosets are considered subspecies of just two spe-
cies, M. argentatus and M. humeralifer (see Rylands et al.,
1993). Meireles et al. (1992) indicated that M. humeralifer
and M. "emiliae" from Rondonia are subspecific to M.
argentatus (see below).

Hapale emiliae was first described by Thomas (1904) from
the Rio Irirf, southern Pari. It was not recognized by
Hershkovitz (1977) who regarded it as a dark form of
C .-'. argentata. Vivo (1985, 1991) revalidated this form
on the basis of specimens from the state of Rondonia. How-
ever, Rylands et al. (1993) indicated that the Rondonia mar-
mosets described by Vivo should be considered a distinct form
based on the fact that its distribution and that of the Hapale
emiliae Thomas, 1904 from Maloca on the Rio Curui (see
Vivo 1985; Avila Pires 1986) are disjunct, and separated by
M. melanurus. M. emiliae was not listed by Groves (1993).
That the Rondonia "emiliae" is a distinct form from emiliae
Thomas 1904 was confirmed by Sena (1998) and Ferrari et
al. (1999), who found M. emiliae to be more similar to M.
argentatus than the "emiliae" from Rondonia. These authors
argue that the Rio Tapaj6s separates two distinct, if very re-

74 Neotropical Primates 8(2), June 2000

cent, radiations of Mico. Alperin (1995) argued that Mico
nigriceps (Ferrari and Lopes, 1992) and "emiliae" from
Rondonia belong to the same species.

C .-'. argentata marcai was described by Alperin 1993.
It is listed here as a species to conform with the view that all
Amazonian marmosets should be considered species (Vivo,
1985, 1991; Mittermeier et al., 1992a). It is known only from
the type locality, "Foz do Rio Castanho (= Rio Roosevelt),
afluente esquerda do Rio Aripuana. Estado do Amazonas,
Brasil" (Alperin 1993). The type locality as described by
Alperin is confused in that the Rio Castanho is not a syn-
onym of the Rio Roosevelt, and is a left bank affluent of the
Rio Roosevelt, not the Rio Aripuana. Confused also is the
map and gazetteer of localities provided in Alperin (1993)
which do not correspond.

The karyotype of M. saterei (Sousa e Silva Jr. & Noronha,
1998) was studied by Barros et al. (1996) and found to be
typical of the "argentata" marmoset group and very similar to
that of M. chrysoleuca.

SAGUINUS Hoffmannsegg, 1807

1. Saguinus I ... ...I (Spix, 1823)
2. S. ... hernandezi Hershkovitz, 1982
3. S. g:wI/I (Jimenez de la Espada, 1870)
4. S. fuscicollis fuscicollis (Spix, 1823)
5. S. fuscicollis fuscus (Lesson, 1840)
6. S. fuscicollis avilapiresi Hershkovitz, 1966
7. S. fuscicollis cruzlimai Hershkovitz, 1966
8. S. fuscicollis leucogenys (Gray, 1866)
9. S. fuscicollis lagonotus (Jimenez de la Espada, 1870)
10. S. fuscicollis primitivus Hershkovitz, 1977
11.S. fuscicollis I. (Pucheran, 1845)
12. S. fuscicollis nigrifrons (I. Geoffroy, 1850)
13. S. fuscicollis weddelli (Deville, 1849)
14. S. fuscicollis melanoleucus (Miranda Ribeiro, 1912)
15. S. fuscicollis crandalli Hershkovitz, 1966
16.S. tripartitus (Milne-Edwards, 1878)
17. S. mystax mystax (Spix, 1823)
18. S. mystaxpileatus (I. Geoffroy & Deville, 1848)
19.S. mystax pluto (Lbnnberg, 1926)
20.S. labiatus labiatus (E. Geoffroy in Humboldt, 1812)
21. S. labiatus rufiventer (Gray, 1843)
22. S. labiatus thomasi (Goeldi, 1907)
23. S. imperator imperator (Goeldi, 1907)
24. S. imperator subgrisescens (Lbnnberg, 1940)
25. S. midas (Linnaeus, 1758)
26. S. niger (E. Geoffroy, 1803)
27. S. inustus (Schwarz, 1951)
28.S. bicolor (Spix, 1823)
29.S. martins martinsi (Thomas, 1912)
30.S. martinsi ochraceus Hershkovitz ,1966
31. S. leucopus (Giinther, 1877)
32. S. oedipus (Linnaeus, 1758)
33. S. geoffroyi (Pucheran, 1845)

The taxonomy of Saguinus ..., follows Hershkovitz
(1982). S. n. g:.wll: is listed as a full species by Hernandez-
Camacho and Cooper (1976) and Defler (1994), on the basis
that it is sympatric with a population of S. ...I in the
region of Puerto Legufzamo in southern Colombia.
Hernandez-Camacho and Defler (1991) listed it as a subspe-
cies of S. .... in conformity with Hershkovitz (1977).
Although Defler (1994) indicated that further studies are
needed to prove the species' status of this form, it was consid-
ered a distinct species by Groves (in press).

The taxonomy of Saguinus fuscicollis is based on Hershkovitz
(1977); see also Cheverud and Moore (1990). S. f
melanoleucus, S. f acrensis, and S. f crandalli were listed as
subspecies of S. melanoleucus by Coimbra-Filho (1990).
Saguinus fuscicollis acrensis Carvalho 1957 is not considered a
valid form but a hybrid between S. f fuscicollis and S. f
melanoleucus on the upper Rio Jurui, following Peres (1993).
S. f cruzlimai and S. f crandalli are of unknown provenance
(Hershkovitz, 1977), and the latter at least may well also be a
hybrid of S. f fuscicollis x S. f melanoleucus. Hernindez-
Camacho and Cooper (1976) indicated the existence of an
undescribed S. fuscicollis subspecies in the region of San Jose
de Guaviare, Colombia (also cited by Defler, 1994). In a
molecular genetic study, Cropp et al. (1999) argued that the
form fuscus was closer to S. ...I than to S. fuscicollis and
gave it species status. Rylands et al. (1993) argued that S. f
melanoleucus should be considered a species distinct from S.
fuscicollis. This arrangement was adopted by Groves (in press),
with the form crandalli (of unknown provenance) as a sub-

Hershkovitz (1977) listed Saguinus tripartitus as a subspecies
of S. fuscicollis. Thorington (1988) argued for its species sta-
tus (see also Albuj a, 1994). It was listed as a species by Rylands
et al. (1993) and Groves (in press).

Saguinus labiatus rufiventer (Gray, 1843) was listed as a jun-
ior synonym of S. 1. labiatus by Hershkovitz (1977). He rec-
ognized that red-bellied tamarins in the north of their range
were distinct but argued that it was probably a clinal varia-
tion. It was considered a valid subspecies, however, by Groves
(in press), with a provisional distribution (also indicated by
Hershkovitz) which extends south from the Rio Solim6es be-
tween the Rios Madeira and Purus to the Rio Ipixuna, an east
bank tributary of the Rio Purus.

Groves (in press) found that, while S. m mystax and S. m.
pluto are very similar to each other), the form pileatus is quite
distinct, and he listed it as a good species, S. pileatus.

Hershkovitz (1977) and Eisenberg (1989) listed the black
handed tamarin as a subspecies of S. midas. On the basis of
morphometric studies of the postcanine dentition, Natori and
Hanihara (1992) found S. m. midas to be more similar to S.
bicolor than to S. m. niger and argued that S. m. niger should
be raised to species status (Rylands et al., 1993), and also

Neotropical Primates 8(2), June 2000 75

noted that there was a need to modify the species' group ar-
rangements of Hershkovitz (1977), that is, placing the S. midas
Group in the Bare-face Tamarin section. Melo et al. (1992),
on the other hand, examined 20 blood genetic systems in the
two subspecies and obtained results compatible with their clas-
sification as subspecies, not being sufficiently divergent to war-
rant species status. Canavez et al. (1999a) using molecular
data (nuclear DNA) found that S. midas midas and S. bicolor
are more closely related to each other than to Saguinus midas
niger. Molecular data from mitochondrial DNA, including
the two S. midas (S. m. midas, S. m. niger) and the three S.
bicolor subspecies (S. b. bicolor, S. b. martinsi and S. b.
ochraceus), strongly support (Bootstrap values of 98%) S. midas
(S. m. midas, S. m. niger) and S. bicolor (S. b. bicolor, S. b.
martinsi, and S. b. ochraceus) as two distinct clades (Cropp et
al., 1999).

Hernindez-Camacho and Defler (1991) indicated the prob-
able existence of two subspecies of S. inustus in Colombia.

Hershkovitz (1977) considered Saguinus geoffroyi to be a sub-
species of S. oedipus. Comparative morphological studies by
Hanihara and Natori (1987), Moore and Cheverud (1992)
and Skinner (1991) argued for them being separate species.
Eisenberg (1989), Rylands (1993). Hernindez-Camacho and
Cooper (1976), Mittermeier and Coimbra-Filho (1981),
Hernindez-Camacho and Defler (1988, 1991), Mittermeier
et al. (1988), Rylands et al. (1993), Groves (1993, in press),
Mast et al. (1993) and Defler (1994) all list S. geoffroyi and S.
oedipus as separate species.


1. Leontopithecus rosalia (Linnaeus, 1766)
2. L. chrysomelas (Kuhl, 1820)
3. L. chrysopygus (Mikan, 1823)
4. L. caissara Lorini & Persson, 1990

The lion tamarins, Leontopithecus, are given as separate spe-
cies following Della Serra (1951), Rosenberger and Coimbra-
Filho (1984), Mittermeier et al. (1988), Natori (1989), and
Rylands et al. (1993). They have been listed as subspecies of
L. rosalia by Coimbra-Filho and Mittermeier (1972, 1973a),
Hershkovitz (1977), Mittermeier and Coimbra-Filho (1981),
Forman et al. (1986) and Seuinez et al. (1988), the latter
two publications on the basis of identical chromosome mor-
phologies. L. caissara Lorini and Persson 1990 is regarded as
a subspecies of L. chrysopygus by Coimbra-Filho (1990). Al-
though some species hybridize in captivity (Coimbra-Filho
and Mittermeier, 1976), no contact or hybridization zones
are found today in thew wild.

CALLIMICO Miranda-Ribeiro, 1912

1. Callimico goeldii (Thomas, 1904)

To date, only a single species has been described for this ge-
nus, although no evaluation of geographic or population varia-
tion has been carried out since the 1977 treatise by

AOTUS Illiger, 1811

1. Aotus lemurinus lemurinus (I. Geoffroy, 1846)
2. A. lemurinus griseimembra (Elliot, 1912)
3. A. lemurinus zonalis Goldman, 1914
4. A. lemurinus brumbacki Hershkovitz, 1983
5. A. hershkovitzi Ramirez-Cerquera, 1983
6. A. trivirgatus (Humboldt, 1812)
7. A. vociferans (Spix, 1823)
8. A. miconax Thomas, 1927
9. A. nancymaae Hershkovitz, 1983
10.A. nigriceps Dollman, 1909
11.A. azarai azarai (Humboldt, 1812)
12.A. azarai boliviensis Elliot, 1907
13.A. azarai infulatus (Kuhl, 1820)

The taxonomy of Aotus follows Hershkovitz (1983), modi-
fied by Groves (in press). A. hershkovitzi was described by
Ramirez-Cerquera in 1983 (Abstracts. IX Cong.
Latinoamericano de Zoologia, Arequipa, Peru, p.148) (see also
Defler 1994). Mentioned only in an abstract, this form re-
quires a proper description.

Hernindez-Camacho and Cooper (1976) referred to A.
trivirgatus zonalis (Goldman 1914) in Panama and Colom-
bia, which is considered a synonym of A. t. griseimembra by
Hershkovitz (1949) and Hill (1960), but as a synonym of A.
1. lemurinus by Hershkovitz (1983), and a synonym of A.
lemurinus by Groves (1993). The status recommended by
Hershkovitz (1983) was accepted by Hernindez-Camacho
and Defler (1991), but Groves (in press) listed it as a valid
subspecies of A. lemurinus. Further modifications by Groves
(in press) of Hershkovitz's (1983) appraisal ofAotus taxonomy
are as follows: the form brumbacki Hershkovitz, 1983 is sub-
specific to A. lemurinus, and the form infulatus (Kuhl, 1820)
is subspecific to A. azarai (see below). The name azarae was
changed by Groves to azarai to conform to the International
Code of Zoological Nomenclature.

Pieczarka and Nagamachi (1988) found that the karyotypes
ofA. infulatus and A. azarae boliviensis are closer to each other
than to the karyotype of A. a. azarae, and argue for the need
to further revise the systematic classification of the genus.
Recent cytogenetic studies by Pieczarka et al. (1992) have
reinforced the validity of the species A. vociferans and A.
nancymaae. A phylogeny based on karyotypes is provided by
Galbreath (1983).

Groves (1993) suggested that A. nancymaae rather than A.
nancymai (see Hershkovitz 1983) is the correct name in ac-
cordance with Article 31 (a(ii)) of the International Code of
Zoological Nomenclature (International Trust for on Zoologi-
cal Nomenclature, 1985). D. Brandon-Jones (in litt., 2000)
argues, however, that surnames have no intrinsic gender, and
by default, when latinized should have a masculine ending.
When there is doubt, the spelling used in the original de-
scription (nancymai Hershkovitz, 1983 and azarae Humboldt,

76 Neotropical Primates 8(2), June 2000

1812, in this case) should be maintained, also following the
directive of the International Code of Zoological Nomencla-
ture. A detailed review of Aotus taxonomy is given by Ford

SAIMIRI Voigt, 1831

1. Saimiri boliviensis boliviensis (I. Geoffroy & de Blainville,
2. S. boliviensis peruviensis Hershkovitz, 1984
3. S. boliviensis pluvialis Lbnnberg, 1940
4. S. boliviensis jaburuensis Lbnnberg, 1940
5. S. vanzolinii Ayres, 1985
6. S. sciureus sciureus (Linnaeus, 1758)
7. S. sciureus macrodon (Elliot, 1907)
8. S. sciureus cassiquiarensis (Lesson, 1840)
9. S. sciureus .. (von Pusch, 1941)
10.S. oerstedii oerstedi (Reinhardt, 1872)
11.S. oerstedii citrinellus Thomas, 1904
12.S. ustus I. Geoffroy, 1843

Saimiri taxonomy follows Hershkovitz (1984) and Groves (in
press). Hershkovitz (1987b, footnote page 22) also recognized
Saimiri boliviensisjaburuensis and S. b. pluvialis (given as jun-
ior synonyms of S. b. boliviensis in Hershkovitz 1984) as valid
subspecies. They are not listed by Groves (in press).
Hershkovitz (1987b) referred to S. vanzolinii Ayres, 1985 as
a subspecies of S. boliviensis.

An alternative taxonomy is presented by Thorington (1985)
as follows: S. sciureus sciureus (includes the forms . .
macrodon, and ustus recognized by Hershkovitz, [1984]), S.
sciureus boliviensis (includes the forms pluvialis Lbnnberg,
1940 and jaburuensis Lbnnberg, 1940 recognized by
Hershkovitz [1987a]), S. sciureus cassiquiarensis, S. sciureus
oerstedii (includes the form citrinellus recognized by
Hershkovitz [1984]), and S. madeirae (given as a junior syn-
onym of S. ustus by Hershkovitz [1984]). Hernindez-
Camacho and Defler (1991) recognize S. sciureus caquetensis
Allen 1916, given as a junior synonym of S. sciureus macrodon
by Hershkovitz (1984). Groves (1993) recognized just five
species of Saimiri (S. boliviensis, S. oerstedii, S. sciureus, S. ustus
and S. vanzolinii). He listed S. b. peruviensis, S. b. jaburuensis
and S. b. pluvialis as synonyms of S. boliviensis, S. o. citrinellus
as a synonym of S. oerstedii, all subspecies of S. sciureus listed
here as synonyms, and S. madeirae recognized by Thorington
(1985) as a synonym of S. ustus. The most recent review (mor-
phology, genetics and behavior) of squirrel monkey taxonomy
by Costello et al. (1993) argued for the recognition of just
two species: S. sciureus in South America, and S. oerstedii in
Panama and Costa Rica. Silva et al. (1993) also gave evidence
for just one large polytypic South American species, Saimiri
sciureus. On the other hand, recent studies of Boinski and
Cropp (1999) using mtDNA, behavioral and morphological
data, and Cropp and Boinski (1999) using two nuclear genes
(IRBP and ZFX) and one mitochondrial (D-Loop) strongly
support the Hershkovitz (1984) taxonomy, advocating four
distinct species: Saimiri sciureus, S. boliviensis, S. oerstedii and

S. ustus.

CEBUS Erxleben, 1777

1. Cebus apella apella (Linnaeus, 1758)
2. C. apella fatuellus (Linnaeus, 1766)
3. C. apella macrocephalus Spix, 1823
4. C. apella peruanus Thomas, 1901
5. C. apella tocantinus Lbnnberg, 1939
6. C apella margaritae Hollister, 1914
7. C libidinosus libidinosus Spix, 1823
8. C libidinosus pallidus Gray, 1866)
9. C libidinosus paraguayanus Fischer, 1829
10. C libidinosusjuruanus Lbnnberg, 1939
11. C nigritus nigritus (Goldfuss, 1809)
12. C nigritus robustus Kuhl, 1820
13. C nigritus cucullatus Spix, 1823
14. C xanthosternos Wied-Neuwied, 1826
15. C albifrons albifrons (Humboldt, 1812)
16. C j .... adustus Hershkovitz, 1949
17. C albifrons aequatorialis Allen, 1914
18. C albifrons cesarae Hershkovitz, 1949
19. C albifrons cuscinus Thomas, 1901
20. C .... malitiosus Elliot, 1909
21. C albifrons trinitatis Von Pusch, 1941
22. C albifrons versicolor Pucheran, 1845
23. C albifrons yuracus Hershkovitz, 1949
24. C j ... leucocephalus Gray, 1865
25. C capucinus capucinus (Linnaeus, 1758)
26. C capucinus limitaneus Hollister, 1914
27. C capucinus imitator Thomas, 1903
28. C capucinus curtus Bangs, 1905
29. C olivaceus olivaceus Schomburgk, 1848
30. C olivaceus apiculatus Hershkovitz, 1949
31. C olivaceus brunneus Allen, 1914
32. C olivaceus castaneus I. Geoffroy, 1851
33. C olivaceus kaapori Queiroz, 1992

As recorded by Hill (1960, pp.405-6) "The classification of
the numerous forms of the present genus [Cebus] is one of the
most vexatious problems in Primate taxonomy and has been so
since early times. The fact has been commented on by every
student of the group, including particularly E Cuvier (1819-
24), I. Geoffroy Saint-Hilaire (1851), Burmeister (1854), Gray
(1870) and Cabrera (1917). The main cause of the confusion
is undoubtedly the proneness of Cebus monkeys to individual
variation, more particularly in certain populations. This is so
much so that scarcely any two individuals are alike. Some of
the variations are due to changes with age; a few are sexual
differences; some may be due to hybridization between adja-
cent populations, but by far the majority are purely individual
and appear quite fortuitously in members of the same stock
brought up under identical environmental conditions..."

Hill's (1960) lamentations are as valid today as they were over
30 years ago, with natural variability exacerbated by hybridiza-
tion caused by the transportation of animals as pets, and the
subsequent release of many in locations outside their natural

Neotropical Primates 8(2), June 2000 77

ranges. This process began with Indians even before Euro-
pean colonization, and continues today due to casual releases
and illegal commerce, and even, paradoxically, with environ-
mental authorities releasing confiscated animals without con-
sideration of their origin.

At species level there is not much disagreement, the difficulty
is in understanding regional variation in order to establish sub-
species. The current taxonomy of the capuchin monkeys has
followed the division established by Hershkovitz (1949, 1955)
of three "untufted" species, the white-throated or white-faced
capuchin Cebus capucinus (Linnaeus, 1758) of Colombia and
Central America, the weeper or wedge-capped capuchin, Cebus
nigrivittatus Wagner, 1848 (referred to as Cebusgriseus E Cuvier,
1819 by Hill [1960]); and Cebus olivaceus Schomburgk, 1848
by Husson [1978], and Robinson [1986]; see also Hershkovitz
[1958]) of the Brazilian Amazon, the Guianas, and Venezuela,
and the white-fronted capuchin, Cebus albifrons (Humboldt,
1812) of the Amazonian regions of Colombia, Ecuador, Peru,
Bolivia, and Brazil, and a single "tufted" species, the brown or
tufted capuchin, Cebus apella (Linnaeus, 1758), occurring
throughout the Amazon and Brazil, as well as northern Ven-
ezuela, northern Argentina and eastern Paraguay, and the
Yungas region of southern Bolivia.

The taxonomy of the tufted capuchins adopted here is that of
the most recent revision by Colin P. Groves (in press). He did
not, however, list any subspecies of C capucinus nor C olivaceus,
and only albifrons, unicolor, cuscinus, trinitatis, aequatorialis,
and versicolor for C. albifrons. Those listed by Rylands et al.
(1995) are however maintained here pending a detailed sys-
tematic revision these forms.

The C apella taxonomy of Hill (1960) was based on personal
communications from Kellogg, who was revising the taxonomy
but died before completing it. Apart from those listed above,
Hill (1960) gives the following subspecies: Cebus apella fatuellus
Colombian brown capuchin, from northern Colombia, type
locality unknown; Cebus apella tocantinus from type locality
Rio Tocantins no clear difference from peruanus; Cebus apella
magnus from the Rio Putumayo; Cebus apella juruanus from

the Rio Jurua; Cebus apella cay Azara's capuchin from Para-
guay; Cebus apella frontatus the white-cheeked capuchin, no
type locality, but argued by Vieira (1955) to occur in the Serra
da Paranapiacaba, Alto da Serra, Canandia, Santa Catarina,
and Rio Grande do Sul; Cebus apella vellerosus from Argentina
is given by Hill (1960) as a synonym of C. a. nigritus.
Hernindez-Camacho and Cooper (1976) argued that C.
apella north of the Amazon in Colombia exhibit no pheno-
typic distinctions sufficient to warrant subspecies. They give
the subspecies as C. a. apella (see also Hernandez-Camacho
and Defler, 1991; Defler, 1994). Hershkovitz in his studies of
Cebus taxonomy was apparently maintaining as valid the sub-
species C a. maranonis, C. a. macrocephalus, C a. pallidus, and
C a. peruanus (apud Encarnaci6n, 1993). All these subspecies
were listed by Aquino and Encarnaci6n (1994).

The subspecies in the Pando Department of Bolivia is uncer-
tain (Buchanan-Smith et al. 2000). According to Hill (1960),
C. apella pallidus Gray, 1985 occurs south of the Rio Madre
de Dios, indicating a distribution in north and central Bo-
livia and adjacent Brazil in the north-west of the Mato Grosso
(as described by Cabrera, 1958). Aquino and Encarnaci6n
(1994) placed C a. pallidus south of the Rio Madre de Dios
in the basin of the Rio Tambopata in Peru. According to
Cabrera (1958) and Hill (1960), macrocephalus Spix, 1823
occurs from the north of Bolivia through the upper Amazon
south of the Rio Amazonas-Solim6es in Brazil, west as far as
the Rfo Ucayali and along the left (west) bank of the Purus
(see Aquino and Encarnaci6n, 1994); this may also be the
form in the Pando. A third candidate is C a. peruanus Tho-
mas, 1901. Cabrera (1958) listed this form as a junior syn-
onym of C a. macrocephalus. Hill (1960) indicated it was
restricted to the upper Rio Jurui and the basin of the Rio
Inambari, and Aquino and Encarnaci6n (1994) include the
Tambopata basin and both sides of the Rio Madre de Dios in
Peru, west as far as the border with Bolivia. Groves (in press)
listed the form pallidus as a subspecies of C. libidinosus. Ander-
son (1997) listed C. apella pallidus for northern and central
Bolivia, with C a. paraguayanus occurring in the south east
of the Andes.

Table 10. A comparison of the classifications of Cebus apella by Vicira (1955), Cabrera (1958) and Hill (1960).

Vieira (1955) (Brazil)
C. nigritus (Goldfuss, 1809)
C. libidinosus pallidus Gray, 1865
C. frontatus Kuhl, 1820
C. vellerosus I.Geoffroy, 1851
C. robustus Kuhl, 1820
C. variegatus E. Geoffroy, 1812
C. libidinosus libidinosus Spix, 1823
C. libidinosus versutus Elliot, 1910
C. i. Fischer, 1829
C. libidinosuspallidus Gray, 1865
C. libidinosusjuruanus L6nnberg, 1941
C. apella apella (Linnaeus, 1758)
C. fatuellus macrocephalus Spix, 1823
C. fatuellus tocantinus L6nnberg, 1939
C. fatuellus peruanus Thomas 1901
C. fatuellus trepidus (Linnaeus, 1766)

Cabrera (1958)
C. apella apella (Linnaeus, 1758)
C. apella libidinosus Spix, 1823
C. apella macrocephalus Spix 1823
C. apella margaritae (Goldfuss, 1809)
C. apella nigritus (Goldfuss, 1809)
C. apella pallidus Gray, 1865
C. apella paraguayanus Fischer, 1829
C. apella robustus Kuhl, 1820
C. apella vellerosus I.Geoffroy, 1851
C. apella versutus Elliot, 1910
C. apella xanthosternos Wied, 1820

Hill (1960)
C. apella apella (Linnaeus, 1758)
C. apella margaritae (Goldfuss, 1809)
C. apella fatuellus (Linnaeus, 1766)
C. apella tocantinus L6nnberg, 1939
C. apella macrocephalus Spix 1823
C. apella magnus von Pusch, 1941
C. apella juruanus L6nnberg, 1939
C. apella maranonis von Pusch, 1941
C. apella peruanus Thomas, 1901
C. apella pallidus Gray 1865
C. apella cay Illiger, 1815
C. apella libidinosus Spix, 1823
C. apella xanthosternos Wied-Neuwied, 1820
C. apella robustus Kuhl, 1820
C. apella frontatus Kuhl, 1820
C. apella nieritus (Goldfuss, 1809)

Neotropical Primates 8(2), June 2000

Kinzey (1982) reviewed the southern forms of C apella, and
did not recognize C a. paraguayanus Fischer, 1829 from Ar-
gentina and Paraguay, considered by him to be a junior syn-
onym of C a. libidinosus. Bodini and Perez-Hernindez (1987)
and Bodini (1989) list only two subspecies for Venezuela: C a.
margaritae (an outlier and probably introduced to the Island
of Margarita) and C a. apella (Federal Territory of Amazonas)
(see also Sanz and Mirquez, 1994).

C xanthosternos, formerly considered a subspecies of Cebus
apella, is genetically quite distinct (Seuinez et al., 1986), and
is, therefore, listed as a separate species (Mittermeier etal., 1988;
see also Coimbra-Filho et al., 1991).

Torres (1988) carried out an incomplete revision. She identi-
fied different forms either side of the Rio Sao Francisco, Rio
Doce, and Rio Parana, and six core areas, five of which have
easily distinguished phenotypes. She indicated that C a. mac-
rocephalus from the upper Amazon may be a valid form, that
C a. apella occurs in the lower Amazon basin and Guianas,
and that C a. xanthosternos may be a valid species with the Rio
Sao Francisco separating C a. xanthosternos and C a. libidinosus
in the past. Jose de Sousa e Silva Jr. (a graduate student at the
Museu Nacional, Rio de Janeiro) is currently carrying out a
systematic revision of the species. The classification of the tufted
capuchins adopted by Groves (in press) is based on the studies
of Torres (1988).

Ten subspecies are listed for Cebus j .. The taxonomy
and distributions of the white-fronted capuchins are confused
and require revision. Groves (in press) listed only six subspe-
cies, not recognizing the forms adustus, cesarae, pleei, malitiosus

and leucocephalus recognized by Hershkovitz (1949). The sub-
species listed here are based on Hershkovitz (1949) and
Hernindez-Camacho and Cooper (1976) (see also
Hernindez-Camacho and Defler, 1985, 1991). Hernindez-
Camacho and Defler (1991) mentioned six subspecies in Co-
lombia and listed five: C a. malitiosus, C a. cesarae, C a.
versicolor, C a. albifrons, and C a. yuracus. They regarded C
a. unicolor as possibly a junior synonym of C. a. j .. ,
although, later, Defler (1994) also included this subspecies
for Colombia. Defler and Hernandez-Camacho (in press) dis-
cussed the taxonomy and nomenclature of C a. j .... and
C a. unicolor and concluded that C a. unicolor is a junior
synonym. Defler (1994) listed six subspecies for Colombia,
three endemic: C a. albifrons, C a. cesarae (endemic), C a.
versicolor (endemic), C a. malitiosus (endemic), C a. unicolor,
and C a. yuracus. C a. pleei Hershkovitz, 1949 and C a.
leucocephalus Gray, 1865 (listed by Hershkovitz 1949) are
regarded by Hernindez-Camacho and Cooper (1976),
Hernindez Camacho and Defler (1991), and Defler (1994)
as junior synonyms of C a. versicolor. Bodini and Perez-
Hernindez (1987) listed three subspecies for Venezuela: C
a. adustus, C a. leucocephalus (regarded a junior synonym of
C a. versicolor by Hernindez-Camacho and Cooper [1976]),
and C a. unicolor. Bodini (1989) pointed out that Hill (1960)
gives four subspecies for Venezuela, C a. cesarae, C a. adustus,
C a. j ... and C a. unicolor. Bodini and PNrez-Hernindez
(1987) give the form in the Federal Territory of Amazonas as
C a. unicolor, in agreement with Hershkovitz (1949), al-
though Hershkovitz was uncertain of the limits of both this
subspecies and C a. unicolor. Hernindez-Camacho and Coo-
per (1976) indicated that the Amazonian form in Venezuela
was C a. j .. based on their belief that C a. unicolor is a
junior synonym. Defler and Hernindez-Camacho (in prep.)

Table 11. A comparison of the classifications of the untufted capuchins, Cebus, by Cabrera (1958), Hill (1960) and Groves (in press).

Cabrera (1958)
C .,:./.,.,* adustus Hershkovitz, 1949
C .,:., ,."',.. aeguatorialis Allen, 1914
C :..p',,,..' '..:.,.,,,.', (Humboldt, 1812)
C ,:.hp,,.'' cesarae Hershkovitz, 1949
C .,:.,I,. .'' cuscinus Thomas, 1901
C :.!.,,,..,. hypoleucus (Humboldt, 1812)
C ,:.! .[,,.' leucocephalus Gray, 1865
C ,:.;, ,'., .pleei Hershkovitz, 1949
C ,:.,.' ,,,. unicolor Spix, 1823
C ,:./h,. '.', versicolor Pucheran, 1845
C. ,:.'f .' yuracus Hershkovitz, 1949

C nigrivittatus apiculatus Hershkovitz, 1949
C nigrivittatus brunneus Allen, 1914
C. nigrivittatus castaneus I. Geoffroy, 1851
C. nigrivittatus nigrivittatus Wagner, 1848
C. nigrivittatus olivaceus Schomburgk, 1848

Hill (1960)
C ,:.,.,,,,.., hypoleucus (Humboldt, 1812)
C ,*:. ./ ,,.,. malitiosus Elliot, 1909)
C .:,./',,.:' cesarae Hershkovitz, 1949
C ,:.h, ,'.,. pleei Hershkovitz, 1949
C ,:.,,,.. ',' versicolor Pucheran, 1845
C ,:./,, ,.,' leucocephalus Gray, 1865
C .,:,./ .,:' adustus Hershkovitz, 1949
C :.!./,,...: ,'.,*,,',,.'. (Humboldt, 1812)
C ,:.h, p,.,' unicolor Spix, 1823
C ,:.!., .,' yuracus Hershkovitz, 1949
C :.'.J,,,.,' cuscinus Thomas, 1901
C ,:../, ,.:' aequatorialis Allen, 1914
C .,:.,'.,,.', trinitatisVon Pusch, 194

C capucinus limitaneus Hollister, 1914
C capucinus imitator Thomas, 1903
C capucinus capucinus (Linnaeus, 1758)
C capucinus nigripectus Elliot, 1909
C capucinus curtus Bangs, 1905

Groves (in press)
C. .:.,", ',,.' .:. ,,t.p,.', (Humboldt, 1812)
C. ,:.,, *.' unicolor Spix, 1823
C. ':.,./.' cuscinus Thomas, 1901
C. :.,./, ,.' trinitatis Von Pusch, 1942
C .:.,,./,,.' aeguatorilais Allen, 1914
C .:,,/'..* ', versicolor Pucheran, 1845

C. olivaceus Schomburgk, 1848

C kaapori Queiroz, 1982

C capucinus (Linnaeus, 1758)

C griseus griseus Cuvier, 1819
C griseus castaneus I. Geoffroy, 1851
C griseus brunneus Allen, 1914
C vriseus leporinus Von Pusch, 1941

Neotropical Primates 8(2), June 2000 79

in redescribing the type species of Simia j ... Humboldt,
1812, demonstrated finally that the form C a. unicolor (Spix,
1823) is a junior synonym, and is therefore not listed here.

C a. hypoleucus (Humboldt, 1812) from northern Colombia
was not recognized as a valid form by Hershkovitz (1949)
who regarded it as a synonym of C a. albifrons, neither by
Hernindez-Camacho and Cooper (1976), Hernindez
Camacho and Defler (1991), and Defler (1994).

Hill (1960) listed five subspecies of Cebus capucinus. C. c.
nigripectus from the upper Rio Cauca in Colombia, C. c.
capucinus (Colombia), C c. imitator (Costa Rica, including
Coiba Island, and Panama), C c. limitaneus (Honduras and
Nicaragua), and C. c. curtus (Colombia). Hernindez-
Camacho and Cooper (1976) argued that variability in popu-
lations of these capuchins on the upper Rio Cauca did not
support the validity of C c. nigripectus, and HernAndez-
Camacho and Defler (1991) and Defler (1994) listed just
two subspecies of C capucinus for Colombia: C c. capucinus
and C c. curtus. Defler (1994) includes C c. nigripectus un-
der the name of C c. curtus. C c. curtus is endemic to the
Colombian Island of Gorgona. It was possibly introduced in
the 16th or 17th Centuries (Hernindez-Camacho and Defler,
1991) and is very similar to Panamanian C capucinus (R. A.
Mittermeier, pers. obs.). Groves (in press) found the species
to be monotypic.

The correct scientific name for the weeper capuchin, has been
the cause for some discussion. Two names in current use are
nigrivittatus Wagner, 1848 (adopted by Cruz Lima, 1945;
Hershkovitz, 1949; Cabrera, 1958) and olivaceus Schomburgk
1848 (adopted by Tate, 1939, and Husson, 1957, 1978). A
third name, griseus E Cuvier, 1819, was used by Hill (1960).

Hill (1960) argued that the first name properly given to this
capuchin monkey is Cebus griseus E Cuvier, 1819, based on a
color plate drawn from a live animal. In a footnote, Hill (1960,
p.429) explained that different copies of Cuvier's plate vary in
the depth of the color of the upper parts of the monkey, and
that this may have caused the doubts expressed by Hershkovitz
(1949) who said that although the head clearly depicts the dis-
tinctive wedge-shaped black patch on the crown, the remain-
der of the body resembles Cebus albifrons, the confusion as such
making the identity of the animal uncertain and the name in-
valid. Hill (1960) insisted that the copy of the plate in his pos-
session depicted the animal which Hershkovitz refers to as
nigrivittatus. There is no preserved type specimen for C griseus,
and the type locality was restricted by Hill to French Guiana
(1958a in Hill 1960). However, Hershkovitz (1959) demon-
strated that Frederic Cuvier did not in fact attribute the name
of griseus to the sajou mile" depicted, and that it was only his
son in 1842 that referred to it as griseus, by which time the
same name had been used by Desmarest for Buffon's "sajou
gris", a tufted capuchin (Groves, in press).
Hershkovitz (1949) opted for the name of Cebus nigrivittatus
Wagner, 1848, although recognizing that Schomburgk had de-
scribed the same animal as C olivaceus in the same year, and

that nigrivittatus may be a junior synonym if it can be proved
that olivaceus predates it. Husson (1957, 1978) argued, how-
ever, for the name of Cebus olivaceus Schomburgk, 1848, on
the basis that the use of the name nigrivittatus is not valid ac-
cording to the International Rules of Zoological Nomencla-
ture. The problem arises from a taxonomic confusion caused
by Von Pusch (1941) who combined the genera Saimiri (the
squirrel monkeys) and Cebus, resulting in two species having
the same name of nigrivittatus, C'. ...-'. .-.-.. Wagner,
1846 (now considered a junior synonym of Saimiri sciureus)
and Cebus nigrivittatus Wagner, 1848. Regarding both as be-
longing to the same genus, Von Pusch (1941), correctly under
the circumstances, rejected the name Cebus nigrivittatus be-
cause it was predated by Chrysothrix nigrivittatus, the former
then being considered a junior secondary homonym. Von Pusch
(1941) renamed the capuchin Cebus capucinus leporinus. Al-
though now in separate genera, Husson (1957), citing Follett
(1955), argued that the International Rules of Zoological No-
menclature do not permit the revalidation of the name, having
been, as it was, invalidated before 1951. The International Rules
published in 1985 state "A junior secondary homonym replaced
before 1961 is permanently invalid" (Article 59b). Hershkovitz
(1955), on the other hand, merely referred to Cebus capucinus
leporinus Von Pusch as a junior synonym of nigrivittatus, and
argued that the ephemeral, and now discarded, homonymy
between the squirrel monkey and the capuchin was unworthy
of consideration (Hershkovitz, 1958).

In summary, C griseus is the earliest name, the validity of which
depends on the interpretation of a color plate, while C.
nigrivittatus and C olivaceus were names published by sepa-
rate authors in the same year, with controversy concerning the
fact that the former, even if it was published earlier, was invali-
dated by Von Pusch (1941), and such an invalidation is deemed
permanent according to the International Trust for Zoological
Nomenclature (1985).

Husson (1957, 1978), Eisenberg (1989), Groves (1993, in
press) and Emmons and Feer (1997), and a number of recent
authors who have studied this animal in the wild (for example,
Fragaszy, 1986; Robinson, 1986, 1988; Miller, 1996, 1998)
refer to the Guiana wedge-capped capuchin as Cebus olivaceus.
A recent study of the chromosomes of a Venezuelan form of
this species refers to it as C nigrivittatus (see Martinez et al.,
1999). It would seem likely, however, that the correct name is
C griseus. Under any circumstances, a modern taxonomic revi-
sion is urgently needed for this species (Bodini, 1989).
Hershkovitz (1949), Cabrera (1958) and Hill (1960) are still
the best sources of reference for the subspecific variation.
Hershkovitz (1949) listed five subspecies which he condition-
ally regarded as valid: nigrivittatus Wagner, 1848, from the
upper Rio Branco Brazil (given as a junior synonym of Cebus
apella by Elliot [1913], and as a junior synonym of C griseus
leporinus Von Pusch, 1941 by Hill, [1960]); olivaceus
Schomburgk, 1848, from the southern foot of Monte Roraima,
Brazil (given as a junior synonym of C, .. .. I-.) Elliot [1913],
and as a junior synonym of C griseus griseus by Hill [1960]);
castaneus I. Geoffroy, 1851 described from Cayenne, French

8o Neotropical Primates 8(2), June 2000

Guiana (listed as a full species by Elliot [1913]; listed by
Hill [1960], but considered of doubtful validity); apiculatus
Elliot, 1907 (listed as a subspecies of Cebus apella by Tate
[1939], and listed as a junior synonym of C griseus griseus
by Hill [1960]), from La Uni6n, Rfo Cuara, Venezuela;
and brunneus Allen, 1914 from northern Venezuela (listed
by Hill [1960], but also considered of doubtful validity).
Hershkovitz (1949) gave the type localities for each of the
forms, but unfortunately not the distributional limits.
A sixth, undescribed form was mentioned by Bodini and
Perez-Hernindez (1987) from central Venezuela, north
of the Rio Orinoco, and a seventh weeper capuchin, de-
scribed as a species but probably only subspecifically dif-
ferent (Harada and Ferrari, 1996), was described from the
eastern Amazon in the state of Maranhao and eastern part
of the state of Pard in 1992: C kaapori Queiroz, 1992.
The distribution of this form is now quite well known, oc-
curring east of the lower Rio Tocantins to the right bank
of the Rio Pindard and the lower Rio Mearim (Queiroz,
1992; Ferrari and Lopes, 1996; Silva, Jr. and Cerqueira,
1998; Carvalho, Jr. et al., 1999).

Bodini and Perez-Hernindez (1987) listed five subspe-
cies for Venezuela: C. n. brunneus, C n. apiculatus, C. n.
nigrivittatus (Amazonian), C. n. olivaceus and a fifth form
which has yet to be classified (widely distributed through-
out central and northern Venezuela, to the north of Rfo
Orinoco). Bodini (1989) pointed out that the subspecies
of C. nigrivittatus are not well defined.

Groves (in press) listed no subspecific forms of C.
olivaceus, although he noted a "differentiation between
paler, more buffy colors in Venezuela and the Demerara
region of Guyana, and dark brown in the hinterland of
Guyana", but he had not seen sufficient specimens from
Brazil to allocate subspecific names to them. The type of
apiculatus is pale and that of castaneus dark.

CALLICEBUS Thomas, 1903

1. Callicebus modestus Lbnnberg, 1939
2. C. donacophilus (D'Orbigny, 1836)
3. C. pallescens Thomas, 1907
4. C. olallae Lbnnberg 1939
5. C. oenanthe Thomas, 1924
6. C. cinerascens (Spix, 1823)
7. C. hoffmannsi Thomas, 1908
8. C. baptista Lbnnberg, 1939
9. C. moloch (Hoffmannsegg, 1807)
10. C. brunneus (Wagner, 1842)
11. C. cupreus (Spix, 1823)
12. C. ornatus (Gray, 1866)
13. C. medemi Hershkovitz, 1963
14. C. torquatus torquatus (Hoffmannsegg, 1807)
15. C torquatus lugens (Humboldt, 1811)
16. C. torquatus lucifer Thomas, 1914
17. C. torquatus purinus Thomas, 1927
18. C. torquatus regulus Thomas, 1927
19. C. personatus (E. Geoffroy, 1812)

20. C. nigrifrons (Spix, 1823)
21. C. melanochir Wied-Neuwied, 1820
22. C. barbarabrownae Hershkovitz, 1990
23. C. coimbrai Kobayashi and Langguth, 1999

Callicebus taxonomy according to Hershkovitz (1988,
1990), modified by Kobayashi and Langguth (1999) and
Groves (in press). Hershkovitz (1990) placed the Callicebus
species into four groupings: 1) C. modestus (C modestus); 2)
C. donacophilus (C olallae, C. donacophilus, C oenanthe);
3) C moloch (C. brunneus, C caligatus, C. cupreus, C dubius,
C hoffmannsi, C moloch, C. cinerascens and C. personatus);
and 4) C. torquatus (C. torquatus). A recent phylogenetic
study, based on cranial measurements, by Kobayashi (1995)
resulted in the following five groupings: 1) C donacophilus
(C. modestus, C. ollalae, C. donacophilus); 2) C cupreus (C.
caligatus, C. cupreus); C. moloch (C. brunneus, C hoffmannsi,
C. moloch, C cinerascens); 4) C. personatus (C. personatus);
and 5) C. torquatus (C. torquatus). Kobayashi (1995) did
not evaluate C. oenanthe and C. p. barbarabrownae, and
the position of C dubius was uncertain. Kobayashi (1995)
indicated that C caligatus is a synonym of C cupreus. Groves
(1992, 1993) indicated that C brunneus and C. hoffmannsi
may be subspecies of C moloch (C. h. baptista is a synonym
of C. hoffmannsi), that C. caligatus and C. dubius may be
synonyms of C cupreus, that C cupreus discolor and C. c.
ornatus are synonyms of C. cupreus, that C. donacophilus
pallescens is a synonym of C. donacophilus, and that all sub-
species of C torquatus and C personatus listed here are syn-
onyms of their respective species. However, in a later publi-
cation and following further study of museum specimens,
Groves (in press) advocated the taxonomy adopted here.
The changes from the Hershkovitz (1990) taxonomy are as
follows: The form pallescens Thomas, 1907 is considered a
separate species rather than subspecific to C donacophilus;
the form baptista Lbnnberg, 1939 is considered a separate
species rather than subspecific to C. hoffmannsi; the forms
discolor (I. Geoffroy & Deville, 1848), dubius Hershkovitz,
1990, and caligatus are not valid; the forms ornatus (Gray,
1866) and medemi Hershkovitz, 1963, are considered sepa-
rate species rather than subspecific to C. cupreus and C.
torquatus, respectively; all the personatus subspecies listed
by Hershkovitz (1990) are considered distinct species, fol-
lowing Kobayashi and Langguth (1999).

Kobayashi and Langguth (1994a, 1994b, 1999) described
Callicebus coimbrai from the coast of the state of Sergipe,
Brazil. On the basis of their study of this new species, along
with previous craniometric studies by Kobayashi (1995),
Kobayashi and Langguth (1999a, 1999b) argued that the
titis of the Atlantic forest should be considered species rather
than subspecies of C personatus. They listed C. nigrifrons,
C. personatus, C. melanochir, and C. barbarabrownae be-
sides C. coimbrai.

PITHECIA Desmarest, 1804

1. Pithecia pithecia pithecia Linnaeus, 1766

Neotropical Primates 8(2), June 2000 81

P. pithecia chrysocephala I. Geoffroy, 1850
P. monachus monachus (E. Geoffroy, 1812)
P. monachus miller Allen, 1914
P. monachus napensis Lbnnberg, 1938
P. irrorata irrorata Gray, 1842
P. irrorata vanzolinii Hershkovitz, 1987
P. albicans Gray, 1860
P. aequatorialis Hershkovitz, 1987

Taxonomy and distributions of Pithecia follow those of
Hershkovitz (1987a) and Eisenberg (1989) except in the
recognition of 1? monachus napensis Lbnnberg, 1939. A
phylogeny at the generic level was recently proposed by
Schneider et al. (1995).

Hershkovitz carried out two revisions of Pithecia, the first
he published in 1979(b) that recognized only four spe-
cies (P pithecia, P. hirsuta, P. monachus and P. albicans)
was superceded by a second in 1987(a). Hershkovitz
(1987a) placed hirsuta Wagner, 1840, as a junior syn-
onym of monachus E. Geoffroy, 1812. The form napensis
Lbnnberg, 1939 was considered by Hershkovitz to be a
local population of hirsuta in 1979(b), and then also jun-
ior synonym of monachus in 1987(a). However, P. Grubb
(pers. comm. 2000) pointed out that the forms napensis
and possibly hirsuta are distinct, and have defined geo-
graphical distributions. P. monachus napensis from the Rfo
Napo, Ecuador, is listed here and, pending further revi-
sion, it is quite possible that hirsuta should also be consid-
ered a valid subspecies of P monachus.

CHIROPOTES Lesson, 1840

1. Chiropotes albinasus (I. Geoffroy & Deville, 1848)
2. C satanas satanas (Hoffmannsegg, 1807)
3. C. satanas chiropotes (Humboldt, 1811)
4. C. satanas utahicki Hershkovitz, 1985

The taxonomy of Chiropotes follows Hershkovitz (1985).
J. P. Boubli (pers. comm.) observed an undescribed
bearded saki from the upper Rio Negro, in Brazil, which
was also confirmed by Marc G. M. van Roosmalen (pers.
comm., 1999).

CACAJAO Lesson, 1840

1. Cacajao calvus calvus (I. Geoffroy, 1847)
2. C. calvus ucayalii Thomas, 1928
3. C. calvus novaesi Hershkovitz, 1987
4. C. calvus rubicundus (I. Geoffroy and Deville, 1848)
5. C melanocephalus melanocephalus (Humboldt, 1811)
6. C. melanocephalus ouakary (Spix, 1823)

The taxonomy of Cacajao follows Hernindez-Camacho
and Cooper (1976), Hershkovitz (1987b), Eisenberg
(1989) and Boubli (1994). Szalay and Delson (1979)
gave C c. rubicundus full species status. The geographic
range of C. c. novaesi is broader than was previously

thought, and a new population of white uakaris has been
recorded from the Rio Jurupari, a tributary of the Rio
Envira, well to the south of C. c. calvus, and separated by
the red C c. novaesi, which may indicate the need for a
reappraisal of the taxonomy and systematics of the species
(Silva, Jr. and Martins, 1999). In a full review of the ecol-
ogy and biogeography of the genus, Barnett and Bran-
don-Jones (1997) argued that Cacajao may be congeneric
with Chiropotes. Morphologically they are separated by the
smaller skull and longer tail of the latter, differences con-
sidered by them to be insufficient to separate them at any
more than the subgeneric level.

ALOUATTA Lacdp&de, 1799

1. Alouatta seniculus seniculus (Linnaeus, 1766)
2. A. seniculus insulanus Elliot, 1910
3. A. seniculus (formerly stramineus)
4. A. seniculus amazonica Lbnnberg, 1941
5. A. seniculus juara Elliot, 1910
6. A. seniculus puruensis Lbnnberg, 1941
7. A. seniculus arctoidea Cabrera, 1940
8. A. sara Elliot, 1910
9. A. nigerrima Lbnnberg, 1941
10.A. belzebul belzebul (Linnaeus, 1766)
11. A. belzebul discolor (Spix, 1823)
12.A. belzebul ululata Elliot, 1912
13.A. guariba guariba (Humboldt, 1812)
14.A. guariba clamitans Cabrera, 1940
15.A. palliata palliata (Gray, 1849)
16.A. palliata mexicana (Merriam, 1902)
17.A. palliata aequatorialis (Festa, 1903)
18.A. coibensis coibensis Thomas, 1902
19.A. coibensis trabeata Lawrence, 1933
20.A. caraya (Humboldt, 1812)
21.A. pigra Lawrence, 1933

Hill (1962) and Stanyon et al. (1995) list nine subspecies of
A. seniculus: A. s. seniculus, A. s. arctoidea, A. s. stramineus, A.
s. macconelli, A. s. insulanus, A. s. amazonica, A. s. juara, A. s.
puruensis, A. s. sara. Cytogenetic studies have been carried
out on; A. s. seniculus by Chu and Bender (1961), Bender
and Chu (1963), and Yunis et al. (1976); A. s. stramineus by
Lima and Seuanez (1991); A. s. macconnelli by Lima et al.
(1990); A. s. sara by Minezawa et al. (1985) and Stanyon et
al. (1995); and A. s. arctoidea by Stanyon et al. (1995). A.
sara is recognized as a full species here following Minezawa
et al. (1985; see also Groves, 1993; Stanyon et al. 1995).
Stanyon et al. (1995) concluded that the number of chro-
mosomal differences between A. s. sara and A. s. arctoidea
was on a similar scale to those found between A. s. sara and
A. s. seniculus by Minezawa et al. (1985). Stanyon et al.
(1995) noted that "as for other organisms in tropical forest
regions, it is probable the biological diversity and number
of species have been underestimated". Hill (1962) argued
that the description of A. s. juara by Elliot (1910) confused
the name and type locality. Hill (1962) demonstrated that
it came in fact from the Rio Jurui in Brazil, whereas Elliot

Neotropical Primates 8(2), June 2000

Table 12. A comparison of the classifications of the spider monkeys, Ateles, by Kellogg and Goldman (1944), Cruz Lima (1945),
Froehlich et al. (1991), Collins (1999a, 1999b) and Collins and Dubach (2000a, 2000b).

Kellogg & Goldman (1944)
A. paniscus paniscus (Linnaeus, 1758)
A. paniscus chamek (Humboldt, 1812)
A. belzebuth belzebuth E. Geoffroy, 1806
A. belzebuth marginatus E. Geoffroy, 1809
A. belzebuth hybridus I. Geoffroy, 1829
A. fusciceps fusciceps Gray, 1866
A. fusciceps robustus Allen, 19142
A. i '-- 1 Kuhl, 1820
A. vellerosus Gray 1866
A. 'yucatanensis Kellogg & Goldman, 1944
A. p an Schlegel, 1876
A. frontatus (Gray, 1842)
A. ornatus Gray, 1870
A. p ianamensis Kellogg & Goldman, 1944
A. i azuerensis Bole, 1937
A. gi 'risescens Gray, 1866
? A. rufiventris Sclater, 1871
Cruz Lima (1945) (Brazilian Amazon)
A. paniscus paniscus (Linnaeus, 1758)
A. paniscus longimembris Allen, 1914
A. marginatus E. Geoffroy, 1809
A. ,. ..,, Hasselt & Kuhl in Kuhl 1820 1
A. behebuth E. Geoffroy, 1806
A. variegatus Wagner, 1840

gave the Rio Juara in the Peruvian Amazon. Three years later,
Elliot (1913) referred to the Rio Juara in the Brazilian, not
Peruvian, Amazon. Hill (1962) indicated that A. s.juarais prob-
ably a junior synonym of A. s. seniculus.

The holotype of the form straminea Humboldt, 1812 is a
female A. caraya, and is, therefore, a junior synonym, and not
available for the red howlers. Further research into the sys-
tematics of the red howlers from northern South America is
necessary to establish the true name for those in the north-
eastern Amazon and Venezuela. A number of names will need
to be considered, including such as Mycetes auratus Gray,
1845 and M. laniger Gray, 1845 (Rylands and Brandon-Jones,
1998). Bonvicino et al. (1996) argued that red howlers east
of the Rio Trombetas are distinct from those to the west, and
gave them the name of macconnelli (see, however, Rylands
and Brandon-Jones, 1998). Groves (in press) listed the Guiana
red howler as A. macconnelli Elliot, 1910.

An undescribed subspecies of A. seniculus north of the Rio
Orinoco was reported by Bodini and Perez-Hernindez (1987).
This may be one of the forms described by Gray (1845).

The taxonomy and distributions of A. belzebul follow Hill
(1962), Langguth et al. (1987), and Bonvicino et al. (1989).
Cruz Lima (1945) listed the form nigerrima L6nnberg 1941
as a full species. Cytogenetic studies have indicated the prob-
ability that A. b. nigerrima is sufficiently distinct as to warrant
species status (Armada et al., 1987; see also Lima and Seuinez

Froehlich et al. (1991)

A. paniscus (Linnaeus, 1758)
A. belzebuth belzebuth E. Geoffroy, 1806
A. belzebuth marginatus E. Geoffroy, 1809
A. belzebuth chamek (Humboldt, 1812)
A. hybridus I. Geoffroy, 1829
A. robustuss Allen, 1914
A. 'fusciceps Gray, 1866
A. i' (subspecies of Kellogg & Goldman, 1944)
Collins (1999a, 1999b; Collins and Dubach, 2000a,
A. paniscus (Linnaeus, 1758)
A. hybridus I. Geoffroy, 1829
A. belzebuth belzebuth E. Geoffroy, 1806
A. belzebuth marginatus E. Geoffroy, 1809
A. behebuth chamek (Humboldt, 1812)
A. i ,' (subspecies of Kellogg & Goldman, 1944,
but including the forms fusciceps and robustus)

1989), and more closely related to seniculus than to belzebul
(see Oliveira, 1996). The possibility remains it may in fact be a
subspecies of A. seniculus.

Hershkovitz (1963, p.397) claimed that, although pre-dat-
ing Stentorfuscus E. Geoffroy, 1812 by two months (as re-
lated by Thomas, 1913), Simia guariba Humboldt, 1812 is a
primary homonym of E. Geoffroy's (1806) guariba, which
Hershkovitz (1963), therefore, regarded as a junior objective
synonym of Alouatta belzebul (Linnaeus, 1766). However,
unlike "simia belzebuth" and "simia seniculus", the name
"guariba" is not mentioned binominally by E. Geoffroy
(1806), who was evidently proposing it only as a vernacular
name with which to distinguish the howler, Alouatta belzebul,
from the spider monkey, Ateles belzebuth. Simia guariba
Humboldt, 1812, therefore,_does not have an available se-
nior homonym, and Hill (1962) and Hirsch et al. (1991),
following Cabrera (1958), were correct in employing it as
the species name for the Atlantic forest brown howling mon-
key. Stentorfuscus E. Geoffroy, 1812 is a junior synonym. The
correct name for the Atlantic forest brown howling monkey
is Alouatta guariba (Humboldt, 1812) (Rylands and Brandon-
Jones 1998). The occurrence of A. guariba in Argentina was
reviewed by Bitteti et al. (1994). Cytogenetic studies were car-
ried out by Lima and Seuinez (1991) and Oliveira etal. (1998).

Villalba et al. (1995) have raised the possibility that A. caraya
extends as far south as northern Uruguay.

Neotropical Primates 8(2), June 2000 83

Table 13. A comparison of the classifications of the woolly monkeys, Lagothrix, by Vieira (1955), Cruz Lima (1945), Hill
(1962), Cabrera (1958) and Fooden (1963).

Vieira (1955) (only Brazil) and Cruz Lima (1945)
(only Brazilian Amazon)
L. lagotricha lagotricha (Humboldt, 1812)
L. lagotricha puruensis Lonnberg, 1940
L. ubericola Elliot, 1909
L. infumata (Spix, 1823)
L. caroarensis Lonnberg, 1940

Hill (1962)
L. cana cana E. Geoffroy, 1812
L. cana lugens Elliot, 1907
L. cana olivacea (Spix, 1823)
L. cana poeppigii Schinz, 1844
L. lazothricha (Humboldt, 1812)

Cabrera (1958)

L. cana cana E. Geoffroy, 1812
L. cana lugens Elliot, 1907
L. cana olivacea (Spix, 1823)
L. cana poeppigii Schinz, 1844
L. lagothricha (Humboldt, 1812)
L. hendeei Thomas, 1927
Fooden (1963)
L. lagotricha cana E. Geoffroy in Humboldt, 1812
L. lagotricha lugens Elliot, 1907
L. lagotricha poeppigii Schinz, 1844
L. lagotricha lagotricha (Humboldt, 1812)
L. flavicauda (Humboldt, 1812)

L. (Oreonax) hendeei Thomas, 1927

The taxonomic status and distribution of Alouatta pigra fol-
lows Smith (1970), Horwich (1983), Horwich and Johnson
(1984). Alouatta palliata luctuosa Lawrence, 1933 listed by
Hill (1962) for Belize was not recognized by Froehlich and
Froehlich (1986). Alouatta pigra luctuosa was listed by Dahl
(1987) for the primates of Belize.

Rodrfguez-Luna et al. (in prep.) have found mixed
populations of Alouatta palliata mexicana and A. pigra in
reduced habitat fragments in Tabasco, Mexico that are
sympatric. Individuals of these populations have been
captured for a comparative molecular genetic analysis of A.
palliata and A. pigra to clarify this intergradation of
populations and possible hybridization processes (Cortes Ortiz
et al, in prep.). Canales-Espinosa et al. (in prep.) are also
studying evident morphological differences in these

Using mtDNA markers, Cortes-Ortiz et al. (in prep.) found
that Alouatta palliata and A. coibensis comprise a very closely
related and monophyletic group of mtDNA lineages. The
mitochondrial DNA divergence between the nominal spe-
cies A. palliata and A. coibensis is very low, showing only 0.1%
sequence divergence, more than an order of magnitude fewer
nucleotide substitutions than were observed between any
other pair of Alouatta species. Divergence between A. palliata
and A. coibensis is similar to mitochondrial DNA distances
observed between geographically-separated populations
within each of these two species.

Groves (1993) lists the following species: A. belzebul (discolor,
nigerrima, and ululata synonyms), A. caraya, A. coibensis
(trabeata synonym), A. fusca clamitanss synonym), A. palliata
(aequatorialis, mexicana synonyms), A. pigra (A. p. luctuosa syn-
onym), A. seniculus, and A. sara. Groves (in press) listed the
form nigerrima as full species.

ATELES E. Geoffroy, 1806

1. Ateles geoffroyi geoffroyi Kuhl, 1820

2. A. geoffroyi azuerensis (Bole, 1937)
3. A. geoffroyi fontatus (Gray, 1842)
4. A. geoffroyi grisescens Gray, 1866
5. A. geoffroyi panamensis Kellogg & Goldman, 1944
6. A. geoffroyi ornatus Gray, 1870
7. A. geoffloyi vellerosus Gray, 1866
8. A. geoffroyi yucatanensis Kellogg & Goldman, 1944
9. A geoffloyi fusciceps Gray, 1866
10.A. geoffroyi rufiventris Allen, 1914
11.A. chamek (Humboldt, 1812)
12.A. paniscus (Linnaeus, 1758)
13.A. marginatus E. Geoffroy, 1809
14.A. belzebuth E. Geoffroy, 1806
15.A. hybridus hybridus I. Geoffroy, 1829
16.A. hybridus brunneus Gray, 1872

The taxonomy of the spider monkeys, Ateles, follows Kellogg
and Goldman (1944) and Konstant et al. (1985), except in
the recognition of the forms hybridus, chamek and marginatus
as distinct species. Hill (1962) followed Kellogg and Goldman
(1944). Heltne and Kunkel (1975) argued that A. fusciceps
robustus Allen, 1914 is a junior synonym of A. f rufiventris
Sclater, 1871. De Boer and Bruijn (1990), Froehlich et al.
(1991), and Medeiros (1994) argued that A. paniscus is a
distinct form with no subspecies. De Boer and Bruijn (1990)
indicated that chamek should be considered a full species,
and likewise Sampaio et al. (1993) argued that the forms
paniscus and chamek should be considered distinct species on
the basis of the genetic distance between them. Ateles chamek
was found to be closer to belzebuth than topaniscus. Froehlich
et al. (1991) indicated that the form chamek is subspecific to
A. belzebuth rather than A. paniscus. Hernindez-Camacho
and Cooper (1976), Hernindez-Camacho and Defler (1991)
refer to all Ateles as subspecies of A. paniscus, following
Hershkovitz (1972). Defler (1994), on the other hand, fol-
lows Kellogg and Goldman (1944) and refers to three spe-
cies in Colombia: A. belzebuth, A. fusciceps, and A. geoffroyi.
Groves (1993) recognized the following species: chamek
Humboldt 1812, paniscus (Linnaeus, 1758), marginatus E.
Geoffroy 1809, fusciceps Gray, 1866, geoffroyi Kuhl, 1820,

84 Neotropical Primates 8(2), June 2000

and belzebuth E. Geoffroy, 1806. He placed the forms
hybridus I. Geoffroy, 1829, and brunneus Gray, 1870 as syn-
onyms of A. belzebuth. Heltne and Kunkel (1975) provided
additional taxonomic notes concerning pelage coloration in
A. paniscus (including chamek) and A. fusciceps. Kunkel et al.
(1980), reported on chromosomal variation and zoogeogra-
phy in the genus.

Hernindez-Camacho and Defler (1988, 1991) and Defler
(1994, in press) argue for the validity of the form brunneus
Gray, 1872 from the Departments of Bolivar, Antioquia and
Caldas, between the lower Rios Cauca and Magdalena in Co-
lombia, and it is listed here as a subspecies of A. hybridus as
recommended by Defler (in press).

A. rufiventris Sclater, 1871, from the Rio Atrato, Darien, Co-
lombia, is listed as incertae sedis by Kellogg and Goldman

Silva-L6pez et al. (1996) argued against the validity of the
subspecies of spider monkey A. geoffroyi pan. In their review
of the taxonomic status of Guatemalan spider monkeys they
concluded that 1) the coloration pattern of A. g. vellerosus
includes a broader spectrum of color and tones than was pre-
viously considered; 2) A. g. yucatanensis may be considered a
valid taxon, but only after more evidence can be obtained
from several localities in the known geographical range of
the subspecies; and 3) based on the available evidence (maps,
vegetation types, and existing records) and supporting the
observations of Konstant et al. (1985) that A. g. pan is prob-
ably merely a variation of the highly variable A. g. vellerosus.

Medeiros et al. (1997) confirmed the conclusions of Froehlich
et al. (1991) in indicating that the forms marginatus (E.
Geoffroy, 1809) and chamek (Humboldt, 1812) should be
considered subspecific to A. belzebuth, and that hybridus I.
Geoffroy, 1829 is better placed as a subspecies of A. geoffroyi.
Medeiros et al. (1997) indicate, however, that from the cyto-
genetic viewpoint rufiventris Sclater, 1871, may well be re-
productively isolated, although a hybrid zone between this
form and A. g. panamensis was reported by Rossan and Baerg
(1977). Collins (1999a, 1999b; Collins and Dubach, 2000a,
2000b) argued strongly for the species status of the form
hybridus I. Geoffroy, 1829, but otherwise agreeing with the
classification proposed by Froehlich et al. (1991).

LAGOTHRIXE. Geoffroy in Humboldt, 1812

Lagothrix lagothricha (Humboldt, 1812)
L. cana cana (E. Geoffroy in Humboldt, 1812)
L. cana tschudii Pucheran, 1857
L. poeppigii (Schinz, 1844)
L. lugens (Elliot, 1907)

The Lagothrix taxonomy follows Fooden (1963), Hernandez-
Camacho and Cooper (1976), Eisenberg (1989), Hernandez-
Camacho and Defler (1991) and Aquino and Encarnaci6n

(1994), and Groves (in press). The taxonomy of Groves (in
press is adopted here, but the genus is in need of revision.
While Fooden (1963) listed all the woolly monkeys except
for L. flavicauda as subspecies of L. lagothricha, Groves gives
them species status and lists two subspecies of cana, the nomi-
nate and L. c. tschudii Pucheran, 1857, a dark form from
Peru and Bolivia. Although for many years there were no
records of woolly monkeys in Bolivia, the recent discovery of
a population in Madidi (Wallace and Painter, 1999) may well
be referable to this subspecies (see Fooden, 1963) or may rep-
resent a new taxon. Fooden (1963) considered the form
olivaceus (Spix, 1823) to be a junior synonym of L. 1. cana.
The form infumata (Spix, 1823) is considered a junior syn-
onym ofL. 1. lagothricha and L. 1. poeppigii, by Fooden (1963).
Vieira (1955) recorded its distribution as eastern Peru, Ecua-
dor, and Amazonas, Brazil from the Rio Jurua. L. caroarensis
Lbnnberg, 1940, known only from its type locality, the Lago
Caroaro, lower Rio Ia on the frontier of Brazil and Peru, is
considered by Fooden (1963) to be a junior synonym of L. 1.
lagothricha. L. ubericola Elliot, 1909 from eastern Peru and
Brazil (Igarap6 Grande, Rio Jurui) and L. 1. puruensis from
Brazil, (Redencao, Lago Beruri, Aruma, Rio Purus), are con-
sidered junior synonyms of L. 1. cana by Fooden (1963).

The juvenile, orange-colored woolly monkey illustrated by
Cruz Lima (1945), of unknown provenance and given merely
as Lagothrix sp. has recently been located on the upper Rio
Jutaf by Carlos Peres (pers. comm. 1999).

OREONAX Thomas, 1927

1. Oreonax flavicauda (Humboldt, 1812)

As a result of his comparative studies of cranial morphology in
the atelines, Groves (in press) concluded that the yellow-tailed
woolly monkey should properly be considered a monotypic
genus, quite separate from the woolly monkeys, Lagothrix.


1. Brachyteles arachnoides (E. Geoffroy, 1806)
2. Brachyteles '-,.... .'.. (Kuhl, 1820)

Vieira (1944) recognized two subspecies of Brachyteles. Recent
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et al. (1991) and Lemos de SA and Glander (1993) indicates
that Vieira's original (1944) standing was valid, but that differ-
entiation is even more extreme and justifies the classification of
the two forms as separate species (see also Coimbra-Filho et al.,
1993). Groves (in press) lists the two muriquis as separate species.


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References. Examples of house style can be found throughout
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Journal article. Stallings, J. D. and Mittermeier, R. A. 1983.
The black-tailed marmoset (C .-'. argentata melanura)
recorded from Paraguay. Am. J. Primatol. 4: 159-163.

Chapter in book. Brockelman, W. Y. and Ali, R. 1987. Meth-
ods of surveying and sampling forest primate populations.
In: Primate Conservation in the Tropical Rain Forest, C. W.
Marsh and R. A. Mittermeier (eds.), pp. 23-62. Alan R. Liss,
New York.

Book. Napier, P. H. 1976. Catalogue of Primates in the British
Museum (Natural History). Part 1: Families Callitrichidae and
Cebidae. British Museum (Natural History), London.

Thesis/Dissertation. Wallace, R. B. 1998. The behavioral
ecology of black spider monkeys in north-eastern Bolivia. Doc-
toral thesis, University of Liverpool, Liverpool, U.K.

Report. Muckenhirn, N. A., Mortensen, B. K., Vessey, S.,
Frazer, C. E. 0. and Singh, B. 1975. Report on a primate
survey in Guyana. Unpublished report, Pan American Health
Organization, Washington, DC.

Neotropical Primates is produced in collaboration
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plied Biodiversity Science, 2501 M. St. NW, Suite
200, Washington, DC 20037.

Neotropical Primates
Journal and Newsletter of the IUCN/SSC Primate Specialist Group
Vol. 8(2), August, 2000


An Assessment of the Diversity of New World Primates
Anthony B. Rylands, Horacio Schneider, Alfedo Langguth, Russell A. Mittermeier,
Colin P Groves and Ernesto Rodriguez-Luna ............................................. .................. 61-93

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