Front Cover
 Back Matter
 Back Cover

Title: Neotropical primates
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00098814/00032
 Material Information
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
Belo Horizonte Minas Gerais Brazil
Publication Date: March 2000
Frequency: quarterly
Subject: Primates -- Periodicals -- Latin America   ( lcsh )
Primates -- Periodicals   ( lcsh )
Wildlife conservation -- Periodicals   ( lcsh )
Genre: review   ( marcgt )
periodical   ( marcgt )
Spatial Coverage: Brazil
Additional Physical Form: Also issued online.
Language: English, Portuguese, and Spanish.
Dates or Sequential Designation: Vol. 1, no. 1 (Mar. 1993)-
Issuing Body: Issued jointly with Center for Applied Biodiversity Science, <Dec. 2004->
General Note: Published in Washington, D.C., Dec. 1999-Apr. 2005 , Arlington, VA, Aug. 2005-
General Note: Latest issue consulted: Vol. 13, no. 1 (Apr. 2005).
 Record Information
Bibliographic ID: UF00098814
Volume ID: VID00032
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: oclc - 28561619
lccn - 96648813
issn - 1413-4705


This item has the following downloads:

NP8 ( PDF )

Table of Contents
    Front Cover
        Front Cover
        Page 1
        Page 2
        Page 3
        Page 4
        Page 5
        Page 6
        Page 7
        Page 8
        Page 9
        Page 10
        Page 11
        Page 12
        Page 13
        Page 14
        Page 15
        Page 16
        Page 17
        Page 18
        Page 19
        Page 20
        Page 21
        Page 22
        Page 23
        Page 24
        Page 25
        Page 26
        Page 27
        Page 28
        Page 29
        Page 30
        Page 31
        Page 32
        Page 33
        Page 34
        Page 35
        Page 36
        Page 37
        Page 38
        Page 39
        Page 40
        Page 41
        Page 42
        Page 43
        Page 44
        Page 45
        Page 46
        Page 47
        Page 48
        Page 49
        Page 50
        Page 51
        Page 52
        Page 53
        Page 54
        Page 55
        Page 56
        Page 57
        Page 58
        Page 59
        Page 60
    Back Matter
        Back Matter
    Back Cover
        Back Cover
Full Text

ISSN 1413-4703



MARCH 2000

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





Ediors: Anthony B. Rylonds and Erneslo Rodriguez-Luna
PSG Chairman: Russell A. Miltermeier
PSG Depuly Chairmen: Anthony B. Rylands and Williom 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 Rodrfguez-Luna, Universidad Veracruzana, 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 Brasileirs de Citncias, Rio de Janeiro, Brazil
Liliana Cortds-Ortiz, Universidad Veracruzana, Xalapa, Mexico
Carolyn M. Crockett, Regional Primate Research Center, University ofWashington, Seattle, WA, USA
Stephen E Ferrari, Universidade Federal do Pari, Belem, Brazil
EckhardW Heymann, Deutsches Primatenzentrum, G6ttingen, Germany
William RK Konstant, Conservation International Washington, DC
RussellA. Mittermeier, Conservation International, Washington, DC
Marta D. Mudry, Universidad de Buenos Aires, Argentina
Horacio Schneider, Universidade Federal do Para, Belim, Brazil
Karen B. Strier, University of Wisconsin, 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 Region Anthony B. Rylands & Ernesto Rodriguez-Luna
Vice Chair for AsiaArdith A. Eudey
Vice Chair for Africa Thomas M. Butynski
Vice Chair for Madagascar Jorg U. Ganzhorn

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

Front Covern
The black-headed uacari, Cacajao melanocephalus ouakary

This issue of Neotropical Primates 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,



Pads and Ree

Neotropical Primates 8(1), March 2000


NeotropicalPrimates has taken on a new role doubling as a journal while maintaining its main function as a newsletter for the
Primate Specialist Group membership as well as for Neotropical primate researchers, zookeepers and conservationists worldwide.
Our intention is to include up to two or three peer-reviewed articles. These will be limited to aspects directly dealing with or
linked to the systematics and taxonomy, biogeography, ecology and conservation of the platyrrhines. For this reason, we are
pleased to welcome a number of renowned Neotropical primatologists with enormous experience in these areas as our new
Editorial Board. They have kindly accepted to play a special role in helping us to glean articles and information of importance for
Neotropical primate studies and conservation, and to maintain, raise even, the standards we are hoping to achieve.

Besides the change into part-journal, moving the editorial office to the Center for Applied Biodiversity Science (CABS) at
Conservation International (CI), Washington, DC, has been the reason for the delays in the publication of NeotropicalPrimates
- for which we apologize. Remedying this situation, we are also pleased to welcome on board Jennifer Pervola, who is now the
Assistant Editor and, by the way, is also helping Gustavo Fonseca with the editing of another SSC Specialist Group newsletter
for the Neotropics Edentata. Please send your contributions, and news items and announcements relevant to Neotropical
primates (as well as sloths, armadillos and anteaters) to her at CABS/CI.

In this issue, we are publishing the descriptions of two new marmoset species, following six which have already been
described from the basin of the Rio Madeira in the Brazilian Amazon in recent years: Callithrix intermedius in 1977, C.
nigriceps and C. mauesi in 1992, C marcai in 1993, C. saterei in 1998, and C. humilis in 1999. There are a further two
articles, one by Alejandro Estrada and co-workers on a population of black howler monkeys, Alouatta pigra, at Parque
Nacional Zona Arqueol6gica de Palenque in Chiapas, Mexico, and a second by Robert Wallace and co-workers on the
primates of the Rios Blanco y Negro Wildlife Reserve in Bolivia.

The following issue, 8(2), of Neotropical Primates will be dedicated to a taxonomic listing of the Platyrrhini; results of the
workshop "Primate Taxonomy for the New Millennium", organized by the PSG, and kindly hosted by the Disney Institute in
Orlando, Florida in February of this year. The aims of this Workshop were to provide the fullest assessment of primate diversity
with our current knowledge, and especially considering the numerous contributions and revelations of genetic studies in the last
decade. A full listing of the primates, be they species or subspecies, is a vital first step for the establishment of conservation
priorities and the full assessment of the Order for the IUCN/SSC Red List. In no way are we imposing a "definitive taxonomy"
and wherever there is disagreement or doubt, we hope it will stimulate substantiated discussion and further research, especially
into such poorly understood genera as Alouatta and Cebus.

Please send us short articles, as well as your publications, information about events, research programs, field sites, announcements,
theses and dissertations, and Society activities, so that we can fulfill our mission of disseminating a wide range of valuable
information for the conservation of primates and their forests in the New World.

Anthony B. Rylands
Ernesto Rodriguez-Luna

2 NeotropicalPrimates 8(1), March 2000


Marc G. M. van Roosmalen1, Tomas van Roosmalen2, Russell A. Mittermeier3, and Anthony B. Rylands4
'Instituto Nacional de Pesquisas da Amazonia, Caixa Postal 478, Manaus 69083-000, Amazonas, Brazil
2Colgate University, Hamilton, New York, USA
3Conservation International, 2501 M. Street NW, Washington, DC 20037, USA
4Center for Applied Biodiversity Science, Conservation International, 2501 M. Street NW Washington, DC 20037, USA.


Two new species of marmoset, Callithrix manicorensis sp. n. and Callithrix acariensis sp. n., are described; the first from the
interfluvium of the Rios Madeira, Aripuana and Manicord, and the second from the interfluvium of the Rios Acari and
Sucundurf, south of their confluence with the Rio Canuma, both in the state of Amazonas, south central Amazonia, Brazil.
The new species are members of the Callithrix argentata group of marmosets, sensu Hershkovitz (1977). Callithrix manicorensis
appears to be most closely related to C. nigriceps, C. marcai and C aff. emiliae, its neighbors to the south, and Callithrix
acariensis appears to be most closely related to Callithrix saterei, its neighbor to the north, and Callithrix melanura to the
south, but they are easily distinguished from them. The most noticeable features of Callithrix manicorensis are the almost
white or silvery upper parts including the arms, hands, chest, mande, nape and forehead, and the indistinct light gray (rather
than black) cap on the head. The back is grayish and transversely white or silvery striped or mottled. The under parts are
yellowish or orange. The legs and feet are orange. The tail is black. Those of Callithrix acariensis include the almost entirely
snow white upper parts including the upper arms, chest, mantle, nape and forehead, and the lack of a dark cap on the head.
The back is grayish. The under parts are white. The legs and feet are bright orange. The tail is dark black with a bright orange
dorsal patch at the proximal end. Most strikingly, this species shows a broad white hip and thigh stripe running from the mid
dorsum (almost continuing across the back) to the knee. The phylogenetic status of the two new marmoset taxa is presented
based on 902 base pairs (bp) of the mitochondrial control region and combined with previously determined sequences from
related species. Additionally, 635 bp of intron 2 of the nuclear b,-microglobulin gene was sequenced for C. manicorensis and
combined with sequences from related species. Callithrix manicorensis is shown to be a member of the C argentata clade
according to mitochondrial control region and b2-microglobulin intron 2 sequences. Callithrix acariensis is found to be most
closely related to C mauesi and C. humeralifera (the tufted-ear marmoset clade) based on mitochondrial control sequence. The
phylogeography of the marmosets is discussed, as well as possible scenarios for historic patterns of dispersal and speciation.
With the discovery of these two marmosets, the number of species of Callithrix increases to 21, of which 20 are endemic to
Brazil (considering C aff. emiliae from Rond6nia a valid species as it is widely separated from C emiliae). The total number of
primate species for the country as a whole increases to 79, 41 of which are endemic.

Key Words Primates, Callitrichidae, marmosets, Callithrix manicorensis sp. n., Callithrix acariensis sp. n., phylogeny,
phylogeography, Brazil, Amazonia.


Descreve-se duas novas especies de sagiii, Callithrix manicorensis sp. n., do interfltvio entire os Rios Madeira, Aripuana e
Manicore, e Callithrix acariensis sp. n., do interfluvio entire os Rios Acarf e Sucunduri, afluentes do Rio Canuma, estado do
Amazonas, Amaz6nia Central Sul, Brasil. As novas especies sao alocadas no grupo de sagiiis Callithrixargentata, sensu Hershkovitz
(1977). A nova espdcie, Callithrix manicorensis, parece ter maior parentesco corn C nigriceps, C. marcai, e C. aff. emiliae, os
seus vizinhos ao sul, ainda que muito distinta destas. Callithrix acariensis parece ter maior parentesco com C saterei, o seu
vizinho ao note, e C. melanura ao sul da sua distribuiqdo. As diferencas mais notAveis em Callithrix manicorensis sao a metade
anterior do corpo de cor branco ou branco-argenteo, incluindo os membros anteriores, as maos, garganta, peito, manto, e
nuca, e a presenca de uma mancha cinza clara ou cinza-argentea (nao preta) no v6rtice. 0 dorso e cinzento ou marron-claro
e transversalmente listrada ou mosqueado com branco ou branco-argenteo. 0 venture e os lados interiores do corpo sao amarelados
ou alaranjados. As pernas e os pes sao alaranjadas. A cauda 6 lisa, nio anelada e preta. As diferencas mais notaveis da nova
esp6cie Callithrix acariensis sio a metade anterior do corpo quase branco neve inclusive os bracos anteriores, peito, juba, nuca
e vertice, sem uma mancha cinza ou preta no vertice. 0 dorso e cinzento. 0 venture e branco. As pernas e os pes sio claramente
alaranjadas. A cauda 6 preta, proximalmente com uma mancha alaranjada. Esta especie mostra uma mancha larga branca na
coxa do joelho quase at6 o dorso. Apresenta-se uma filogenia que inclui as duas novas especies, baseada em 902 pares de bases
da regio control mitocondrial e 635 pares de bases do intron 2 do gene nuclear b2-microglobulin. 0 arranjo cladistico
sugere que Callithrix manicorensis seja um membro do grupo C. argentata, e que C acariensis seja mais pr6xima a C mauesi

Neotropical Primates 8(1), March 2000

e C. humeralifera (baseada na anglise da regido control mitocondrial). Discuta-se a filogeografia dos sagiiis amaz6nicos tanto
quanto cenirios possfveis para os padres hist6ricos de dispersio e especiaio. Corn essas descobertas, acrescenta-se mais duas
espdcies ao genero Callithrix, sendo agora um total de 21, com 20 endfmicas ao Brasil (considerando Callithrix aff. emiliae de
Rond6nia uma espdcie vilida, geograficamente isolada e separada de Callithrix emiliae). 0 ndmero de especies de primates
existentes no Brasil elevou-se para 79. Destas, 41 sao endemicas ao pais.
Palavras-Chave Primatas, Callitrichidae, sagiis, Callithrix manicorensis sp. n., Callithrix acariensis sp. n., filogenia, filogeografia,
Brasil, Amaz6nia.


All but one of the 19 marmosets, Callithrix Erxleben, 1777,
recognized to date are endemic to Brazil. The black-tailed
marmoset, Callithrix melanura, ranges into Bolivia and Para-
guay (Stallings and Mittermeier, 1983; Brown and Rumiz,
1986). Two major revisions of this genus have been carried
out in the last two decades, one by Hershkovitz (1977), who
recognized just three species and 11 taxa, and the other by
De Vivo (1988, 1991), who, listed an additional form, C.
emiliae, and elevated all the forms to species status.
Mittermeier etal. (1988) agreed with the taxonomy suggested
by De Vivo, but also recognized a sixth eastern Brazilian spe-
cies, C. kuhlii, which was considered a hybrid by Hershkovitz
(1977) and as merely a pelage color variant of C penicillata
by De Vivo (see Rylands et al., 1993). Mittermeier et al.
(1992) followed De Vivos decision on the Callithrix argentata
group, elevating all forms to full species status, and listed nine
species within this group.

In this paper we describe two new species from the basin of
the Rio Madeira, state of Amazonas, Brazil, increasing the
number of Callithrix species to 21 (see Table 1), the number
of Brazilian primate species to 79, 41 (52%) of them being
endemic, and the number of callitrichid taxa to 59 (see Van
Roosmalen et al., 1998). Figure 1 (after Van Roosmalen et aL,
1998) shows the distribution of 14 species of the Amazonian
Callithrix argentata group, including the two described here,
C manicorensis and C acariensis, and also Callithrix humilis
M. G. M. van Roosmalen, T. van Roosmalen, Mittermeier
and Fonseca, 1998.

C manicorensis is from the region confined by the Rios Ma-
deira, Aripuana and Manicor6, and C. acariensis occurs be-
tween the Rios Acari and Sucunduri, both in the state of
Amazonas in south central Amazonia, Brazil (Fig. 2). In its
pelage C. manicorensis is most similar to C. aff. emiliae from
Rond6nia (see footnotes to Table 1), C nigriceps, and C marcai
(Figs. 3 and 4), with which it is parapatric. However, C.
manicorensis is quite distinct, and clearly deserves recogni-
tion as a full species. C acariensis is most similar to C. melanura
to the south, C intermedia along the left (west) bank of the
upper Rio Aripuana, and C. saterei to the north (between
the Rios Canumd and Abacaxfs), all four sharing a very dis-
tinct pale thigh and hip stripe (Fig. 3).

Callithrix manicorensis was first discovered by Marc G. M.
and Tomas van Roosmalen on 5 July, 1996, during a field
trip to the region in search of a new species of dwarf marmo-

set, Callithrix humilis (Van Roosmalen et al, 1998). The ani-
mal was being kept as a pet and, as they were informed, had
been collected on the Rio Mariepaud. Callithrix acariensis
was first discovered by Marc G. M. van Roosmalen on 29
August, 1996, when he obtained a baby pet monkey from
locals living along the right bank of the lower Rio Acarf.

Callithrix manicorensis sp. n.

Holotype: INPA 2511, juvenile female, one of two juveniles
obtained alive by Marc G. M. van Roosmalen on 30 Decem-
ber 1996, from the vicinity of the town of Manicord. The
specimen INPA 2511 died soon after being acquired and is
deposited in the mammal collection of the Instituto Nacional
de Pesquisas da Amaz6nia (INPA) Museum, Manaus,
Amazonas, Brazil.

Paratype: A juvenile male, obtained alive together with the
holotype INPA 2511 by Marc G. M. van Roosmalen on 30
December, 1996, from the vicinity of the town of Manicore.
It is still alive in captivity at the time of this publication. It will
eventually be placed in the mammal collection of the Instituto
Nacional de Pesquisas da Amaz6nia (INPA), Manaus,
Amazonas, Brazil.

Type locality: Seringal Sao Luis, east bank of the middle Rio
Madeira, in the vicinity of the town of Manicore, state of
Amazonas, Brazil (Fig. 2). This region is located in south cen-
tral Amazonia, Brazil, south of the Rio Amazonas, east of the
Rio Madeira, and west of the lower Rio Aripuana. Coordi-
nates for the type locality are 050 50' 28" S, 610 18' 19" W
Altitude 45 m.

Other specimens examined: A specimen collected alive by
Marc G. M. van Roosmalen on 5 July, 1996. It was reported
to have been captured along the lower Rio Mariepaui. This
specimen, an adult male, died a month later and only the skull
and skeleton was preserved (see Table 3 for cranial measure-
ments; Table 2 for intermembral, brachial and crural indices).
It is deposited in the mammal collection of the Instituto
National de Pesquisas da Amaz6nia (INPA), Manaus, Amazonas,
Brazil, under the number INPA 2512. It is depicted in Figure
5. Furthermore, a skull and a complete skeleton of an adult
female that was kept for several years in the breeding center of
the first author, has been preserved and is deposited in the
mammal collection of the Instituto Nacional de Pesquisas da
Amaz6nia (INPA), Manaus, Amazonas, Brazil, under the num-
ber INPA 3930. Figure 9 shows the skull of this specimen, and
Table 4 gives the cranial and dental measurements.

4 NeotropicalPrimates 8(1), March 2000

Table 1. The taxonomic arrangements for the marmosets Cebuella and Callithrix in Hershkovitz (1977), De Vivo (1988, 1991), Mittermeier
et al (1988), Rylands et al. (1993), Van Roosmalen et al. (1998), and in this paper.
Hershkoitz 119"-) De Vivo 11988, 19911 Mittermeier t ,dl. 19831 Van Roosmalrn et al. This paper
Rflands et al. l19931 1 19981
Pygmy marmosets C. '../., ,An-'..n, in t)Clombia. Peru Br.i,! R .lin
C pygmaea' C pygmaea C pygmaea C p. pygmaea2 C p. pygmaea
C p. niveiventris C. p. niveiventris
Dwarf marmoset Callithrix humilis group (Brazilian Amazonia)
C humilis3 C. humilis
Marmosets Callithrix jacchus clade (Eastern Brazil)
C. jacchus jacchus C jacchus C jacchus C jacchus C. jacchus
C. j. Penicillata C penicillata C penicillata C penicillata C penicillata
C. j. geoffroyi C geoffroyi C geoffroyi C geoffroyi C. geoffroyi
C. j. aurita C aurita C aurita C aurita C aurita
C. j. flaviceps C flaviceps C flaviceps C flaviceps C flaviceps
C kuhlil C kuhlii C kuhlii
Marmosets Callithrix argentata clade (Brazilian Amazonia, Bolivia and Paraguay)
C argentata argentata C argentata C a. argentata C argentata C argentata
C a. leucippe C leucippe C a. leucippe C leucippe C leucippe
C a. melanura C melanura C a. melanura C melanura C melanura
C emiliae' C emiliae C aff. emiliae C aff. emiliae
C emiliae5 C emiliae
C. nigriceps' C niriceps C nigriceps
C marcai C marcai
C sateret C saterei
C manicorensis
C acariensis
C humeralifer humeralifer C humeralifera C h. humeralifer C humeralifera C humeralifera
C h. intermedius' C intermedia C h. intermedius C intermedia C intermedia
C h. chrysoleuca C chrysoleuca C h. chrysoleuca C chrysoleuca C chrysoleuca
C mauesi"' C mauesi

1 Rosenberger (1981 see also Rosenberger and Coimbra-Filho, 1984) argued, on morphological terms, that the pygmy marmoset should
correctly be included in the genus Callithrix. This argument was not maintained in Rosenberger et al. (1990), although recently
Nagamachi etal. (1992), Barroso (1995), Schneider et al. (1996), Barroso etal. (1997), Tagliaro (1997), and Tagliaro etal. (1997) have
also argued for this arrangement on the basis of their karyotype and molecular genetics.
2 Lonnberg (1940) argued for two subspecies of Cebuellapygmaea, C.p. pygmaea and C p. niveiventris. Although they were listed by Napier
(1976), Hershkovitz (1977) did not recognize them. Van Roosmalen and Van Roosmalen (1997) and Van Roosmalen etal. (1998) argued
that the two forms are distinct and valid subspecies. J. Hernmndez-Camacho (pers. comm.) believes they should be regarded as distinct
3 Described by Van Roosmalen etal. (1998).
4 Coimbra-Filho (1984, 1985, 1990), Rylands (1989), Mittermeier etal. (1988), Natori (1990), Rylands etal. (1993), and Groves (1993)
recognize Callithrix kuhlii from southern Bahia as a valid form. Hershkovitz (1977) regarded it as a hybrid between C j. geoffroyi and C
j. penicillata. De Vivo (1991) considered it to be indistinguishable from C penicillata.
5 Callithrix emiliae was first described by Thomas (1920) from the Rio Irirf, southern Pari. It was not recognized by Hershkovitz (1977)
who regarded it as merely a dark form of C argentata argentata. De Vivo (1985; see also De Vivo, 1991), revalidated this form on the basis
of specimens from the state of Rond6nia. However, Rylands et al. (1993) argued that the Rond6nia marmosets described by De Vivo
(1985) should be considered a distinct species based on the fact that the distribution of this form and that of the C emiliae described by
Thomas (1920) from Maloca on the Rio Curui (see De Vivo, 1985; Avila Pires, 1986) are disjunct, and separated by C melanura. C
emiliae was not listed by Groves (1993). Here we recognize the Rond6nia marmoset (C aff. emiliae of De Vivo, 1985) and, provisionally,
C emiliae Thomas, 1920 from the Rio Irirf until further research is carried out on these marmosets.
6 Described by Ferrari and Lopes (1992).
7 Callithrix marcai was described as a subspecies of Callithrix argentata by Alperin (1993). It is listed here as a species to conform with the
view that all Callithrix should be considered species (De Vivo, 1985, 1991; Mittermeier et al., 1992). 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 synonym of the Rio Roosevelt, and is a left bank affluent
of the Rio Roosevelt, not the Rio Aripuana.
8 Described by Silva Jr. and Noronha (1998).
9 Callithrix intermedia was described as an intermediate color form of C humeralifer by Hershkovitz (1977). Pelage color and pattern, and
its geographic distribution, however, would place it as a subspecies of C argentata if, following Hershkovitz (1977), the Amazonian
marmosets are considered subspecies of just two species, C argentata and C humeralifer (see Rylands et al., 1993).
"Described by Mittermeier et al. (1992).

Neotropical Primates 8(1), March 2000 5

Figure 1. Distribution of 14 species of the Amazonian Callithrix argentata group, including
C. manicorensis and C acariensis described in this paper, and also of Callithrix humilis Van
Roosmalen et al., 1998. Map by Stephen D. Nash.

62' 58

Figure 2. Distribution of Callithrix manicorensis and Callithrix acariensis. Map by Stephen D. Nash.

6 Neotropical Primates 8(1), March 2000

Table 2. The intermembral (humerus + radius)/(femur + tibia) x 100), brachial (radius/humerus) x
100) and crural (tibia/femur) x 100) indices of one adult Callithrix manicorensis compared with
those of Callithrix argentata, Callithrix humilis and Cebuella pygmaea (measurements taken by Lesa
A. Davis and Susan Ford).

Intermembral Index
C argen..;..
C manic. ..
C humi,..
Cebuella i,(-:a..


C %".d .. ~..


(-rural Inde%.

I ,.5


Table 3. Cranial measurements (mm) of the adult
male (Registration number INPA 2512) of Callithrix
manicorensis from the Rio Mariepaui.

Skull and mandible
Greatest Skull Length (SL)
Condylobasal Length
Zygomatic Width (ZW)
Biorbital Width (BW)

Braincase Length
Braincase (Postorbital) Width (PW)
Across First Molars (AM)
Length of Mandible (ML)
Coronoidal Height


pm -m 9.27
i-mi2 15.62
c-m2 11.72
pm2-m2 10.07
i-m2 15.80
c-m2 12.56

Geographic distribution: The species is known from the west
bank of the lower Rio Aripuana from the mouth, south to
the confluence with the Rio Roosevelt, and along the east
bank of the Rio Madeira from the mouth of the Rio Aripuana,
just south of the town of Novo Aripuana, south to the mouth
and right bank of the Rio Manicore. The southern limit for
the species is probably the headwaters of the Rio Manicord,
close to the Rio Roosevelt.

Habitat: The various groups observed in the wild by Marc
G. M. van Roosmalen were in dense primary terra firme
rainforest and in secondary forest surrounding plantations
and fields. Very high densities were found in so-called seringal,
a high riverbank forest, managed since the early days of rub-
ber exploitation in the Amazon, where the density of rubber
trees was increased artificially. Seringais are an early form of
agroforestry, in which areas of high riverbank forests were
turned into productive multi-species forests, rich in a num-
ber of commercially valuable and edible fruit producing na-
tive trees, including rubber (Hevea brasiliensis), Brazil nut
(Bertholletia excelsa), 'orelha de macaco' (Enterolobium
schomburgkii), wild cocoa (Theobroma spp.), 'bacurf' (Rheedia
spp., Platonia insignis), 'ingo' (Inga spp.), 'taperebi' (Spondias
mombin), 'biribi' (Rollinia squamosa), soursopp' (Annona
spp.), and a number of tree species belonging to the
Sapotaceae and Arecaceae (palm) families. The seringais ap-

pear to offer optimal habitat and a year-round food supply
for small monkeys, such as marmosets, pygmy marmosets,
dwarf marmosets, titi and night monkeys.

Diagnosis: A marmoset of the Callithrix argentata group
(sensu Hershkovitz, 1977), characterized by a black tail and a
grey cap on the head, but quite distinct from C. nigriceps, C.
marcai, and C aff. emiliae. The lightest of these four species
and most advanced phaeomelanically (sensu Hershkovitz,
1977), with the least pigmented face and ears. The cap on
the head is light gray instead of black in the other members
of this subgroup, and the naked face and eyes are surrounded
by white fur. The general appearance of this monkey when
spotted in the field is an overall drab whitish marmoset with
orange legs and a pinkish to orange naked face and ears, ir-
regularly brown mottled in the adults, and a slightly trans-
versely- striped gray back. This marmoset is easy to distin-
guish from the recently described black-capped dwarf mar-
moset (Van Roosmalen et al., 1998), with which it is sympat-
ric in the northern part of its distribution. The dwarf mar-
moset is almost one-third the size of Callithrix manicorensis,
has an overall dark brown appearance with light brown to
orange brown under parts. It spends more time gouging bark
on tree trunks, in an upright squirrel-like posture, than true
marmosets. The differences between C. manicorensis,
Callithrix humilis and C marcai, can be seen in the color
illustrations in Figures 3 and 4.

Description of the holotype: General coloration of upper parts
white, transversely mottled with gray; tail black; crown on
head triangular, gray; sides of upper arms, shoulders and neck
light grayish white, not mottled; back and rump grayish white,
mottled with darker gray, giving a dark gray, slightly
transversally striped appearance; outer sides of legs and feet
orange; upper surface of lower arms and hands slightly or-
ange, lower surface of lower arms ochraceous to orange; lower
surface of legs orange; white rim on forehead above the eyes
continuing onto the temples; face naked; facial skin and cheeks
orange, pigmented dark brown around the eyes and on the
lower lip and chin; nose and circumbuccal area naked, pink
to flesh-colored, dark brown mottled around the nostrils, with
a narrow dark brown vertical nose stripe, without moustache;
auricle non-pigmented, pink or flesh-colored, slightly hairy
(only in juveniles); throat, ventral surface of neck and chest
white; belly and inner surface of legs bright orange; exposed
genitals non-pigmented, white, hypertrophied (Fig. 6); tail
black, proximal end slightly mottled with orange and gray;
eyes with black pupil and dark brown iris. Weight of juvenile

Neotropical Primates 8(1), March 2000

female holotype: 135 g. Body measurements: head-body
length 160 mm; tail length 270 mm; total length 430
mm; hind foot 51 mm; hand 34 mm; hallux 8 mm, other
toes 12-15 mm, pollex 9 mm, other fingers 13-15 mm.

Origin of the name: This marmoset was first collected and
observed in the wild along the right bank of the Rio Manicord,
municipality of Manicord. It is therefore named after the river
Manicore, a right bank tributary of the Rio Madeira, south
central Amazonia. The name places it geographically relative
to its congeners.

Vernacular name: This marmoset is referred to as sagii, sauim
(souim), or 'branquinho' (= little white fellow, in Portuguese)
by the local people. For an English name, we suggest the Rio
Manicord marmoset.

Callithrix acariensis sp.n.

Holotype: A live female infant obtained by Marc G. M. van
Roosmalen on 29 August 1996 on the right bank of the Rio
Acarf near its confluence with the Rios Sucunduri and
Canumi. It was being kept as a pet and was only 1-2 months
old. It was kept in captivity in the endangered primate breed-
ing center in Manaus (managed by the first author). It died
when adult and is deposited in the mammal collection of the
Institute Nacional de Pesquisas daAmaz6nia (INPA), Manaus,
Amazonas, Brazil, registration number INPA 3931. The skull
is shown in Figure 10; cranial and dental measurements are
given in Table 4.

Type locality: A small settlement on the right bank of the
lower Rio Acarf close to the confluence with the Rios
Sucundurf and Canumi, state of Amazonas, south central
Amazonia, Brazil. Coordinates for the type locality are 050 07'
08" S, 60 01' 14" W.

Geographical distribution: The species is known from the
right bank of the lower Rio Acarf, and assumed to occur in
the entire interfluvium of the Rios Acarf and Sucundurf, south
perhaps to a contact zone with Callithrix melanura somewhere
between the Rio Aripuani and the Rio Juruena.

Habitat: This new species has not been observed in the wild,
but according to local settlers living along the right bank of
the Rio Acari it is found mainly in dense primary terra firme
rainforest and in secondary forest surrounding their planta-
tions and manioc fields.

Diagnosis: A marmoset of the Callithrix argentata group
(sensu Hershkovitz, 1977), it is one of the most colorful of the
Amazonian marmosets with its bright orange lower back,
under parts, legs and proximal end of the black tail, and pre-
dominantly white upper parts and chest contrasting with the
partly black pigmented muzzle, narrowly triangular nose
patch and ocular rings of the otherwise pink face. Its ears are
dark pigmented, partially covered with white hairs (Figs. 7
and 8). It is the lightest of its closest relatives, C saterei and C.
melanura, easy to distinguish from its neighbor to the north,

C saterei, which has a much darker back, lower arms and
legs, an almost totally naked head, asymmetrically black and
non-pigmented naked ears, a narrower yellowish orange thigh
and hip stripe, and a large black triangular patch on and above
the nostrils; its black tail lacks the orange proximal ring or
spot (Fig. 3). It is also very easy to distinguish from C. melanura,
its neighbor to the south, which is predominantly gray and
brown lacking orange and white in its pelage, and lacking the
white fur surrounding the face (Fig. 3).

Description of the holotype: General coloration of upper parts
white, distally grading into gray to dark gray mixed with or-
ange; tail black with a very distinct orange proximal end or
spot; crown on head light gray or white; sides of upper arms,
shoulders, and neck white or white with grayish hair tips; outer
sides of legs and feet bright orange, with a strikingly white, 2
cm wide thigh and hip stripe; upper surface of lower arms
and hands gray mixed with orange, lower surface of lower
arms white with orange; lower surface of legs white distally
grading into orange; forehead white continuing onto the
temples, nape, chest and upper arms; face naked; facial skin
pink or flesh-colored, with black pigmented patches between
the eyes, aside of the nostrils, and on the lower lip and chin;
nose and circumbuccal area naked, pink to flesh-colored, with
a narrowly triangular black patch on the nose, without mous-
tache; auricle dark pigmented, grayish, covered with quite a
few white hairs; throat, ventral surface of neck and chest white,
giving the appearance of a white mantle; belly and inner sur-
face of legs bright orange; exposed genitals non-pigmented,
white, pudenda hypertrophied, 2.2 x 2 x 1.5 cm (Fig. 7); tail
black, proximal end bright orange forming a 2 cm wide ring;
eyes with black pupil and orange brown iris. Weight of adult
female holotype: 420 g. Body measurements: head-body
length 240 mm; tail length 350 mm; total length 590
mm; hind foot 63 mm; hand 45 mm; hallux 10 mm,
other toes -15-20 mm; pollex -12 mm, other fingers 15-20

Origin of the name: This marmoset was collected along the
right bank of the Rio Acari. It is therefore named after this
river, the only reference to this remote, largely uninhabited
part of south central Amazonia, in order to place it geographi-
cally relative to its congeners.

Vernacular name: This marmoset is referred to as sagiii or sauim
(souim) by the local people. For an English name, we suggest
the Rio Acarf marmoset.

Phylogenetic Placement of Callithrix manicorensis and
Callithrix acariensis Based on Nuclear and Mitochondrial

Materials and Methods

Hair, skin, and liver samples were obtained from deceased
specimens from the Breeding Center for Endangered Ama-
zonian Primates in Manaus, Amazonas, Brazil, which were
deposited at the zoological museum of the Instituto Nacional
de Pesquisas da Amaz6nia (INPA), Manaus, Amazonas, Bra-

Neotropical Primates 8(1), March 2000

zil. Sampled marmosets included three Callithrix humilis, one
Callithrix manicorensis, one Callithrix acariensis, and one
Cebuella pygmaea niveiventris. DNA was extracted from the
tissues following the protocol provided by Gibco BRL
DNAzol extraction kit. The mitochondrial control region (also
known as the D-loop) and intron 2 of the P2 -microglobulin
gene were chosen for amplification because they had previ-
ously been characterized for various related callitrichids, and
because they arguably proffered some of the greatest phylo-
genetic resolution for mitochondrial and nuclear genomes in
these organisms, respectively. Primers were designed based
on previously sequenced callitrichid mitochondrial control
regions (Tagliaro et al., 1997). A heavy strand primer was
designed in the CSB1 region of the control region, as well as
a shorter primer in the same region for sequencing purposes.
In addition, primer L15926 (Kocher et al., 1989) was used
for light strand amplification. Primers used for b2-
microglobulin intron 2 PCR amplification were the same used
in Canavez et al., 1999. Polymerase chain reaction conditions
consisted of: 80 C (1 min), 94 C (2 min), 35 cycles of {940
C (1 min), 500 C (1 min 30 sec), 720 C (2 min)} and 72 C
(10 min) for the mitochondrial control region. PCR condi-
tions for the p2 -microglobulin intron 2 were the same used
by Canavez et al.: 950 C (30 sec); 30 cycles of {950 C (20
sec), 60 C (30 sec), 72 (30 sec)} and 72 C (5 min). PCR
reactions were on a 1% agarose gel, since the control region
amplifications were expected to be around 900 bp long, and
the p2-microglobulin amplifications not longer than 700 base
pairs, taking interspecific variation in length into account.

Upon successful amplification, PCR products were trans-
formed into pCR@4-TOPO plasmids which were subse-
quently transformed into One Shot Escherichia coli cells
using the TOPO TA Cloning@ Kit for Sequencing (Invitrogen
Corporation). Cells were plated on LB-agar plates 50 tg/ml
ampicillin, and left to grow at 370 C overnight. Any cells trans-
formed with the plasmid were conferred resistance to ampi-
cillin. Ten to fifteen colonies were picked for each species,
and cultured overnight in 2 ml of LB-amp both at 370 C.
Plasmids were then isolated using the QIAprep Spin
Miniprep Kit (Qiagen Inc.). EcoRI sites at both ends of the
insertion site of the plasmid allowed for digestion and subse-
quent screening for plasmids with inserts on a 1% agarose
gel. Plasmids containing inserts were selected for sequencing.
Primers M13 (forward and reverse), provided by Qiagen Inc.,
allowed for sequencing of the inserts from plasmids ends. The
cloning and transformation process allowed for separation of
desired sequences from any non-specific amplifications, or
"smears", that would otherwise muddle sequencing perfor-
mance. Amplifications were sequenced with an ABI Prism
310 Spectrum sequencer.

Sequences were aligned by eye, using the program Sequencer.
Additionally, the sequences obtained in this study were com-
bined with those obtained from GenBank using accession
numbers given in Tagliaro et al. (1997) for the mitochon-

drial control region, and Canavez et al (1999) for the nuclear
intron. For the control region, sequences of three Callithrix
argentata, three C. mauesi, two C. humeralifera, two C.
geoffroyi, two C penicillata, two C jacchus, five C kuhli, two
C. aurita, one Cebuella pygmaea, and one Leontopithecus
chrysomelas were obtained from GenBank. For the [32-
microglobulin intron 2, sequences were obtained for one
Callithrix humeralifera, one C emiliae, one C. kuhli, one C
penicillata, one Cebuella pygmaea, one Callimico goeldii, one
Saguinus midas, one S. imperator, and one Leontopithecus
chrysopygus. Upon alignment, sequences were entered into the
PAUP program (Swofford, 1994) for phylogenetic analysis.
A maximum parsimony (MP) algorithm was used to analyze
the data, and bootstrap analyses (100 replicates) were per-
formed on the resulting consensus trees. Pair-wise nucleotide
distances were also determined for the species under consid-


DNA extraction and PCR amplification proved to be equally
successful for tissues stored at either freezing conditions or
100% alcohol before deposition in RNA later. For the mito-
chondrial control region, an alignable 902 base pair (BP) seg-
ment including both hypervariable regions and the conserved
central domain was amplified and sequenced for all speci-
men, except Cebuella pygmaea. This sequence constitutes ap-
proximately 90% of the entire mitochondrial control region.
For Cebuella pygmaea, only a 532 bp segment including the
central conserved domain and the primate insertion sequence
was obtained. For the 12 microglobulin intron 2, a 607 bp
segment was obtained for Callithrix manicorensis alone. These
sequences were aligned with corresponding sequences from

Pair-wise divergences for Callithrix and Cebuella were calcu-
lated. For the mitochondrial control region, those among Ama-
zonian marmosets (Callithrix) ranged from approximately 3%
to 7% (2-4% within each subclade, 6-7% between the
argentata and mauesi subclades). Divergences between any
Amazonian marmoset and Cebuella were on the order of 11-
12%. Within-species divergences ranged from 0-3%. C
mauesi and C humeralifera showed the greatest within-spe-
cies divergence, possibly due to hybridization between these
two species. Their separation by the Rio Mau&s-Aoi, a minor
tributary of the Rio Amazonas and contact zone with C
humeralifera between the headwaters of the Rio Mauns and
the Rio Abacaxfs, may explain why hybridization still occurs
between these two species, having never become completely
isolated from one another. These findings call for a closer in-
vestigation into hybridization rates between populations of
these two species, and whether they actually represent dis-
tinct species.

Neotropical Primates 8(1), March 2000

Map of the Distributions of
Amazonian Marmosets,
Cebuella and Callithrix

Figure 3. A schematic representation of the distributions of the Amazonian marmosets, Callithrix, delimited by rivers.
Illustration by Stephen D. Nash.

Neotropical Primates 8(1), March 2000



C. nigriceps


C. aff. emiliae

C. marcai

C. humilis


Figure 4. The Rio Acarf marmoset, Callithrix acariensis and the Rio Manicor6 marmoset, Callithrix manicorensis compared with Callithrix
nigriceps Ferrari and Lopes, 1992, Callithrix aff. emiliae (to be described as a new species), Callithrix marcai Alperin, 1993 (possibly
parapatric in the south of the range of C. manicorensis) and Callithrix humilis Van Roosmalen, Van Roosmalen, Mittermeier and Fonseca,
1998 (sympatric with C. manicorensis). Drawings not to scale. Illustration by Stephen D. Nash.

Neotropical Primates 8(1), March 2000 11

Figure 5. Adult male Callithrix manicorensis (INPA 2512) from the Rio Mariepaui (left), and a young female (INPA 3930) (right).
Photographs by Marc G.M. van Roosmalen.

Figure 6. The adolescent male C manicorensis, the paratype which is still alive, displaying its genitals. Photograph
by Marc G.M. van Roosmalen.

12 Neotropical Primates 8(1), March 2000

Figure 7. Adult female Callithrix acariensis holotypee INPA 3931), showing the distinct thigh stripe (left), and the hypertrophied pudenda
(right). Photographs by Marc G.M. van Roosmalen.

Figure 8. Adult female Callithrix acariensis holotypee INPA 3931). Photograph by Marc G.M. van

Neotropical Primates 8(1), March 2000 13

Table 4. Cranial and dental measurements of the holotype (adult female) of Callithrix acariensis (INPA 3931) and the adult
female of Callithrix manicorensis (INPA 3930) (mm).

Skull and mandible [N PA 39 30 INPA 3931
Greatest Skull Length (SL) 45.7 47.6
Zygomaric Width (ZW) 31.3 30.8
Biorbital Width (BW) 26.2 27.3
Postorbital Width (PW) 22.4 23.3
Nasion-Basion 32.2 33.1
Basion-Prosthion 33.7 34.8
Biauricular Breadth 26.3 26.5
Height of Canine (CH) 4.3 5.2
Length of Mandible (ML) 28.8 31.1
Across First Molars (AM) 11.70 13.5
Across Canines 8.4 8.9
Dental Field (P2-M2) 11.10 10.8
Premaxilllary Height 8.3 8.3
Intradentale Superior to Premaxilla-Maxilla Junction at Alveolus (IS-P) 5.0 4.6
Intradentale Superior to Nasion (IS-NA) 15.1 15.8
Intradentale Superior to Posterior Nasal Spine (IS-PNS) 15.6 16.3
Bregma to Nasion (BR-NA) 23.6 23.3
Nasion to Fronto-Malar Junction at Orbit (NA-FM) 14.1 14.2
Fronto-Malar Junction to Pterion 6.7 8.3
Fronto-Malar Junction to Zygomaxillare Superior (FM-ZS) 4.9 6.8
Fronto-Malar Junction to Maxillary Tuberosity (FM-MT) 13.2 9.9
Zygomaxillare Inferior to Premaxilla-Maxilla Junction at Alveolus (ZI-PM) 12.0 12.1
Zygomaxillare Inferior to Zygomaxillare Superior (ZI-ZS) 7.5 6.8
Zygomaxillare Inferior to Maxillary Tuberosity (ZI-MT) 5.6 6.0
Anterior Teeth
I' Length 2.1
I' Breadth 1.6
I' Height 2.7
12 Length 1.9
12 Breadth 1.6
i, Length 1.4
i, Breadth 1.6
i, Height 3.0
i Length 1.3 1.4
i2 Breadth 2.2 2.2
C,Length 2.5 2.7
CBreadth 1.5 1.7
Cheek Teeth
P, Length 2.4
P, Breadth 1.7
P.Length 1.9
P4 Breadth 1.8
M, Length 2.5
M, Width 2.0
M2 Length 2.0
M, Width 1.7
o = missing

Neotropical Primates 8(1), March 2000

The pairwise divergences between the Atlantic forest
Callithrix (the Callithrix jacchus clade), or ouistiti's, and the
Amazonian Callithrix (the Callithrix argentata clade) range
from 12-13%, while divergence between Atlantic forest
Callithrix and Cebuella range from 14-15%. Divergence be-
tween all marmosets, pygmy marmoset, and dwarf marmoset
and the out-group, the lion tamarins Leontopithecus was over
20%. These genetic distances suggest that, contrary to sug-
gestions by previous molecular studies (e.g., Tagliaro et al.,
1997; Barroso et al., 1997) to subsume Cebuella within the
Callithrix genus, the Amazonian Callithrix species and the
Atlantic forest Callithrix species should be separated into dis-
tinct genera. Given a 3-7% divergence for species within ei-
ther clade, and a 10-11% divergence between clades, it seems
phylogenetically more parsimonious to group the two clades
into separate genera in recognition of their evolutionary dis-
tinctness. If we want to use taxonomic classifications to reflect
actual evolutionary distinctness, we should keep the Atlantic
forest Callithrix or ouistiti's in the genus Callithrix Erxleben,
1777, and place the Amazonian Callithrix or Amazonian mar-
mosets in the genus Mico Lesson, 1840. Groves (in press) lists
the Amazonian marmosets as belonging to the subgenus Mico
Lesson, 1840.

Using PAUP, a maximum parsimony analysis with 100 boot-
strap replications yielded the phylogenetic tree for the mito-
chondrial control region seen in Figure 11. This tree does
not include the shorter sequence of Cebuellapygmaea obtained
in this study. When this sequence is included in the phylog-
eny, curtailed to only 532 bp for all species, bootstrap values
are raised significantly for some nodes. The relationship of
the Atlantic forest marmosets, or ouistiti's (the Callithrixjacchus
clade), as diverging before the radiation of Amazonian mar-
mosets (also known as the Callithrix argentata clade) includ-
ing the pygmy marmosets (Cebuella pygmaea), remains in con-

cordance with the phylogenetic trees presented by Tagliaro
et al. (1997). Callithrix manicorensis joins most closely with
Callithrix argentata with significant bootstrap values.

The p32-microglobulin intron 2 phylogenetic tree yielded
much less percent variation (0.4-1.0 %) among taxa, and was
insufficient for clear resolution. However, it did support the
grouping of Callithrix manicorensis within the Amazonian
Callithrix clade with a significant bootstrap value.


The finding that Callithrix acariensis is more closely related
to the C. mauesi/humeralifera clade concurs with these spe-
cies' shared geographical location between the Rios Tapaj6s
and Madeira/Aripuana, an area that is largely contiguous and
undivided by major rivers. Given the morphological similar-
ity between C acariensis and C melanura and the recently
described C. saterei, and the fact that they all share the same
geographic region between the Rio Tapaj6s and the Aripuana/
lower Madeira, it seems plausible that marmoset species in
this region form a separate subclade within the Amazonian
marmoset subgenus. It, therefore, seems likely that starting
with the archetypic C. melanura with its huge distribution in
the headwaters of the Rios Madeira, Mamor6 and Guapord,
and Aripuana and Juruena, lacking major river barriers, this
subclade speciated toward the north via C. intermedia con-
fined to the Rios Roosevelt and Aripuana interfluvium, C
acariensis, confined to the interfluvium of the Rios Acarf and
Sucundurf, C humeralifera in the interfluvium of the Rios
Tapaj6s, Amazonas and Mauds-A d, finally speciating into
C. chrysoleuca, occupying the Rios Amazonas, Madeira,
Canumi, Aripuana and Acarf interfluvium from north to
south, while C saterei branched off into the area between
the Rios Canuma and Abacaxfs, and C mauesi between the

Figure 9. Skull of adult female Callithrix manicorensis
(INPA 3930). Photograph by Stephen D. Nash.

Figure 10. Skull of adult female Callitbrix acarie-is
(INPA 393 1). Photograph by Stephen D. Nash.

Neotropical Primates 8(1), March 2000 15

Parani Urarii and Rio Maues-Acq. In this context it is inter-
esting to note the strikingly white hip/thigh stripe consistent
in all these species (rather inconspicuous only in C mauesi),
and the partially hairy auricles in especially juvenile C.
intermedia and C acariensis, culminating in the tufted ear C
humeralifera, C mauesi, and C chrysoleuca. Both characteris-
tics are absent in all other existing Amazonian marmosets.

The grouping of C manicorensis with C. argentata seems to
be a paradox because the two species' geographic ranges are
on opposite ends of the Amazonian marmosets' distribution.
Neighboring species of C. manicorensis such as C. marcai, C.
nigriceps and C aff. emiliae should be analyzed before any
conclusions may be drawn from this result. However, these
results do indicate a division of the Amazonian marmosets
that includes C melanura, C. intermedia, C acariensis, C.
saterei, C. mauesi, C. humeralifera, and C chrysoleuca in one
clade, and C. aff. emiliae, C. nigriceps, C. manicorensis, C.
marcai, C emiliae, C. leucippe and C argentata in the other
dade. C marcai and C nigriceps are morphologically similar
to C manicorensis (Van Roosmalen et al., 1998). The status
of C emiliae is less easy to predict, although its geographical


Figure 11. Callitrichid gene tree based on 902 base pairs (bp) of
the mitochondrial control region and 635 bp of intron 2 of the
nuclear p2-microglobulin gene, analyzed using maximum parsimony.
Bootstrap values are given above branches. Callithrix manicorensis
dearly belongs to the C argentata clade, Callithrix acariensis is most
closely related to C.mauesi and C. humeralifera, the tufted ear
Callithrix clade.

distribution suggests it is more closely related to the C
argentata/manicorensis subclade. Genetic analysis of all spe-
cies should be done in the future to help resolve this ques-
tion. The present data indicate that one subclade (C
argentata/manicorensis) is divided geographically by the other
subclade (C acariensis/mauesi/chrysoleuca). A plausible scenario
would be the speciation from an ancestral C melanura-like
archetype going both northwest and filling the interfluvium
of the Rios Madeira, Aripuana and Roosevelt with C aff.
emiliae in the Rios Madeira and Ji-Parani interfluvium, C
nigriceps in the Rios Madeira and Marmelos interfluvium, C
marcai in the Rios Marmelos and Roosevelt interfluvium, and
finally C manicorensis 'trapped' in the interfluvium between
the unsurpassable Rio Madeira and the lower Rio Aripuana,
and northeast passing the upper Rios Juruena and Tapajds
filling the Rios Xing6 and Tapaj6s interfluvium with C
emiliae in the Rios CuruA and Irirf interfluvium, C argentata
in the Rios Xingd, Amazonas and Cuparf interfluvium, and
C leucippe in the Rios Tapaj6s and Cuparf interfluvium. It is
rather likely that new species will be found in the near future
east of the Rios Juruena and Tapaj6s.

Patterns of marmoset radiation emerging from the findings
here presented suggest that the river barrier hypothesis
(Wallace, 1852) could be used to account for much of the
primate diversity in the Amazon basin. Unlike the small ro-
dents in Da Silva and Patton's (1993) study, primates as a rule
cannot swim for very long distances and particularly marmo-
sets, tamarins and titi monkeys drown at the spot when fallen
into the water. Therefore, their only chance for substantial
cross-river migration involves passive migration, in which tracts
of land (with periodically flooded forest) 'jump' from one side
of the river to the other, in the process of ox-bow lake forma-
tion of meandering rivers. Although ox-bow lakes are com-
mon in the upper reaches of the Rios Purds and JuruA in
western Amazonia (Peres et al, 1997), the greater slope of
rivers coming from the Brazilian Shield in eastern Amazonia,
starting with the Rio Madeira and its upper course tributar-
ies and moving east, precludes such meandering and, with it,
passive cross-river migration of breeding primate populations.
Therefore, it is possible that rivers in central and eastern
Amazonia are effective barriers that have led to the genetic
diversity despite their relatively recent divergences, 1-3 mil-
lion years ago, according to molecular docks based on IRBP
intron sequence divergences (Barroso et al., 1997). Besides
the evidence for hybridization found in this study and Tagliaro
et al. (1997) between C humeralifera and C mauesi, there
have been no reports of naturally occurring hybridizations
between other Amazonian marmosets, suggesting that its vari-
ous taxa represent actual species, or distinct evolutionary en-
tities, a conclusion further corroborated by the extreme in-
tolerance of parapatric species toward each other (pers. obs.
in captivity). Tamarins, on the other hand, are not exclusively
allopatric, as they are separated by meandering rivers which
are less effective as reproductive barriers, and their popula-
tions are characterized by geographic gradations of pelage
color variations. The greater slope in the marmoset's distribu-
tion, from the Brazilian Shield in the south toward the Ama-

16 Neotropical Primates 8(1), March 2000

zon River, leading to south-northward, fast-flowing, straight
rivers may thus have contributed to the species diversity of
the marmosets, whereas many of the tamarin taxa continue
to interbreed at irregular intervals, and are thus unable to
fully differentiate into reproductively-isolated species. For this
reason, the current classification of many of the tamarin taxa
into subspecies (Rylands etal., 1993) and marmosets into dis-
tinct species is probably truly reflective of the evolutionary
history of these primates.

In summary, our findings suggest that the common ancestral
population of Callithrix, the dwarf marmoset (see Van
Roosmalen and Van Roosmalen, in prep.), and Cebuella was
initially separated into an Amazonian and Atlantic clade. The
Amazonian clade subsequently gave rise to the ancestral dwarf
marmoset and then to Cebuella. The isolation of the ancestral
Cebuella from other marmosets (after passing the upper Rio
Madeira) in the geographic range of the tamarins west of the
Rio Madeira, may explain its subsequent divergence from the
other marmosets. The dwarf marmoset, is, however, sympat-
ric with Callithrix manicorensis. There is no way of knowing
whether its initial divergence involved its isolation from the
ancestral Callithrix in its present location in the northern part
of the interfluvium of the Rios Madeira and Aripuanr, and
that subsequent invasion by Callithrix followed much more
recently. This possible scenario would explain most parsimo-
niously their present distributions. The geographic division
of the Callithrix argentata subclade from the Callithrix mauesi/
humeralifera subclade recognized in this paper, suggests that
eastern Amazonia was predominantly occupied by the
Callithrix argentata subclade, and that a subsequent radia-
tion from the south by the Callithrix mauesi/humeralifera
subclade drove the Callithrix argentata subclade species from
the region between the Rios Canuma and Aripuana, and the
Rio Tapaj6s. Alternately, the Callithrix mauesi/humeralifera
subclade may have predated the Callithrix argentata subclade
in eastern Amazonia, and a radiation of the argentata subclade
from the south into two directions, northeast and northwest,
enveloped the mauesi/humeralifera subclade. The lack of tol-
erance for marmoset sympatry, due to ecological niche simi-
larities, necessitated much of this excluding radiation and ex-
treme competition, leaving no room for sympatric existence
even between clearly distinct species such as those in the
argentata subclade and the mauesi/humeralifera subclade. It
follows that river isolation is a necessary prerequisite for ini-
tial isolation and diversification, and that rivers tend to act as
divisions between marmoset species that have evolved into
separate species, acting as buffer zones that prevent poten-
tially more successful species from invading neighboring ar-
eas and effectively out-competing other species. Rivers, there-
fore, do not merely give rise to new species by genetic isola-
tion, they also serve as boundaries that help maintain species
diversity throughout east central Amazonia.

Conservation Status of the New Species

There is no reason to suspect that Callithrix manicorensis is
threatened. However, the total area in which it has been con-

firmed to occur is c. 15,000 km2 (c. 90 x 170 km), a rather
small distribution. High densities of this monkey are found
close to human habitations, but it seems to be rather rare in
undisturbed terra firme rain forest far from the banks of riv-
ers and larger creeks, where human settlements are mainly
situated. The forests between the lower Rio Aripuana and
Rio Madeira are still in good, almost pristine condition, al-
though logging has taken place along the more navigable riv-
ers, occurring especially when the rivers are high. There are
no major towns or cities in the area, except for Manicord.
Novo Aripuana is located just north of the species' northern
limit, and is on the opposite side of the Rio Aripuana near its
mouth. No Indian tribes (which usually hunt even the small-
est birds and mammals) live in the area, and the local people
(called caboclos) are widely scattered and live only in small
settlements of one to several families along the major rivers,
the Rio Madeira and the Rio Aripuani, and along the lower
courses of a number of minor rivers, such as the Rios Marauri,
Urui, Maripaui, Araud, Atininga, and the right bank of the
Rio Manicord (Fig. 2). The interfluvial basins of these black-
and clear-water rivers are practically uninhabited.

The conservation status of Callithrix acariensis is unknown,
but the interfluvium of the Rios Acarf and Sucunduri, where
it occurs, though relatively small, is practically uninhabited.
The only way to reach the area is by means of a 40 km dirt
road which links the town of Novo Aripuana with the lower
Rio Araras, a tributary of the Rio Acarf, or by boat entering
the Rio Canumi via the Rio Madeira, all the way upriver to
the confluence with the Rio Sucundurf and Rio Acarf. Since
marmosets generally prefer edge habitats and secondary for-
est on high dry land (terra firme), Callithrix acariensis is as-
sumed to be relatively rare, considering the fact that there is
almost no human habitation in the interfluvium of the Rios
Acarf and Sucundurf. No major deforestation or logging has
been planned for the area, and therefore the monkey is not
considered threatened.


The authors thank Betty van Roosmalen-Blijenberg for tak-
ing care of the captive primate specimens, Stephen Nash for
producing the drawings that illustrate this paper, John Fleagle
for his help in describing the skulls, William Konstant for
assistance in editing and lay-out, and Ella Outlaw for sup-
port from the office in Washington, DC. We extend special
thanks to Valquemar Sousa de Aradijo, who assisted us in the
field. Support for fieldwork and for publication of this paper
was provided by the Margot Marsh Biodiversity Foundation
and Conservation International, Washington, DC, and Con-
servation International do Brasil, Belo Horizonte.


Alperin, R. 1993. Callithrix argentata (Linnaeus, 1771):
Consideraq6es taxon6micas e descriico de subespdcie nova.
Bol. Mus. Para. Emilio Goeldi, Sir. Zool. 9(2): 317- 328.

Neotropical Primates 8(1), March 2000 17

Avila-Pires, E D. de. 1986. On the validity and geographical
distribution of Callithrix argentata emiliae Thomas, 1920
(Primates, Callithricidae). In: A Primatologia no Brasil 2,
M. T. de Mello (ed.), pp.319-322. Sociedade Brasileira de
Primatologia, Brasilia.
Barroso, C. M. L. 1995. Filogenia molecular da subfamilia
Callitrichinae (sensu Rosenberger, 1981). Doctoral Thesis,
Universidade Federal do Pari, Belkm.
Barroso, C. M. L., Schneider, H., Schneider, M. P. C.,
Sampaio, I., Harada, M. L. Czelusniak, J. and Goodman,
M. 1997. Update on the phylogenetic systematics of New
World monkeys: Further DNA evidence for placing the
pygmy marmoset (Cebuella) within the genus Callithrix. Int.
J. Primatol. 18(4): 651-674.
Brown, A. D. and Rumiz, D. I. 1986. Distribuci6n de los
primates en Bolivia. In: A Primatologia no Brasil 2, M. T.
de Mello (ed.), pp.335-363. Sociedade Brasileira de
Primatologia, Brasilia.
Canavez, E C., Moreira, M. A. M., Ladasky, J. J., Pissinatti,
A., Parham, P and Seuinez, H. N. 1999. Molecular phy-
logeny of New World Primates (Platyrrhini) based on p2-
microglobulin DNA sequences. Molec. Phylogenet. Evol. 12
(1): 74-82.
Coimbra-Filho, A. E 1984. Situaqio atual dos calitriquideos
que ocorrem no Brasil (Callitrichidae-Primates). In: A
Primatologia no Brasil, M. T. de Mello (ed.), pp.15-33.
Sociedade Brasileira de Primatologia, Brasilia.
Coimbra-Filho, A. E 1985. Sagiii-de-Wied Callithrix kuhli
(Wied, 1826). FBCN/Inf., Rio de Janeiro 9(4): 5.
Coimbra-Filho, A. F. 1990. Sistemitica, distribuicio
geogrdfica e situacqo atual dos simios brasileiros (Platyrrhini
Primates). Rev. Brasil. Biol. 50: 1063-1079.
De Vivo, M. 1985. On some monkeys from Rond6nia, Brasil
(Primates: Callitrichidae, Cebidae). Pap. Avulsos Zool., Sao
Paulo 4: 1-31.
De Vivo, M. 1988. Sistemitica de Callithrix Erxleben, 1777
(Callitrichidae, Primates). Doctoral thesis, Universidade de
Sao Paulo, Sao Paulo.
De Vivo, M. 1991. Taxonomia de Callithrix Erxleben, 1777
(Callitrichidae, Primates). Fundacio Biodiversitas, Belo
Ferrari, S. E and Lopes, M. A. 1992. A new species of mar-
moset, genus Callithrix Erxleben 1777 (Callitrichidae, Pri-
mates) from western Brazilian Amazonia. Goeldiana Zoologia
(12): 1-3.
Groves, C. P 1993. Order Primates. In: Mammal Species of
the World: A Taxonomic and Geographic Reference, 2nd Edi-
tion, D. E. Wilson and D. M. Reeder (eds.), pp.2432-77.
Smithsonian Institution Press, Washington, DC.
Groves, C. P. In press. The Taxonomy of Primates. Smithsonian
Institution Press, Washington, DC.
Hershkovitz, P. 1977. Living New World Monkeys (Platyrrhini)
with an Introduction to Primates, Vol. 1. University of Chi-
cago Press, Chicago.
Kocher, T. D., Thomas, W. K., Meyer, A., Edwards, S. V.,
Paabo, S., Villablanca, E X. and Wilson, A. C. 1989. Dy-
namics of mitochondrial DNA evolution in animals: ampli-
fication and sequencing with conserved primers. Proc. Natl.
Acad. Sci. USA 86: 6196-6200.

Lonnberg, E. 1940. Notes on marmosets. Arkiv Zoologi,
Stockholm 32A (10): 12-21.
Mittermeier, R. A., Rylands, A. B. and Coimbra-Filho, A. E
1988. Systematics: species and subspecies an update. In:
Ecology and Behavior of Neotropical Primates, Vol. 2, R. A.
Mittermeier, A. B. Rylands, A. E Coimbra-Filho and G. A.
B. da Fonseca (eds.), pp.13-75. World Wildlife Fund, Wash-
ington, DC.
Mittermeier, R. A., Schwarz, M. and Ayres, J. M. 1992. A
new species of marmoset, genus Callithrix Erxleben 1777
(Callitrichidae, Primates), from the Rio Mauds region, state
of Amazonas, Central Brazilian Amazonia. Goeldiana
Zoologia (14): 1-17.
Nagamachi, C. Y., Pieczarka, J. C. and Souza Barros, R.M.
de. 1992. Karyotypic comparison among Cebuellapygmaea,
Callithrix jacchus and C. emiliae (Callitrichidae, Primates)
and its taxonomic implications. Genetica 85: 249-257.
Napier, P. H. 1976. Catalogue of Primates in the British Mu-
seum (Natural History). Part 1: Families Callitrichidae and
Cebidae. British Museum (Natural History), London.
Natori, M. 1990. Numerical analysis of the taxonomical sta-
tus of Callithrix kuhli based on the measurements of the
postcanine dentition. Primates 31(4): 555-562.
Peres, C. A., Patton, J. L. and da Silva, M. N. E 1997. River-
ine barriers and gene flow in Amazonian saddle-back tama-
rins. Folia PrimatoL 67: 113-124.
Rosenberger, A. L. 1981. Systematics: the higher taxa. In:
Ecology and Behavior of Neotropical Primates, VoL 1, A. E
Coimbra-Filho and R. A. Mittermeier (eds.), pp.92-7.
Academia Brasileira de Ciencias, Rio de Janeiro.
Rosenberger, A. L. and Coimbra-Filho, A. F. 1984. Morphol-
ogy, taxonomic status and affinities of the lion tamarins,
Leontopithecus (Callitrichinae, Cebidae). Folia Primatol 42:
Rosenberger, A. L., Setoguchi, T. and Shigehara, N. 1990.
The fossil record of callitrichine primates. J. Hum. Evol 19:
Rylands, A. B. 1989. Sympatric Brazilian callitrichids: the
black-tufted-ear marmoset, Callithrix kuhli, and the golden-
headed lion tamarin, Leontopithecus chrysomelas. J. Hum.
Evol. 18(7):679-695.
Rylands, A. B., Coimbra-Filho, A. E and Mittermeier, R. A.
1993. Systematics, distributions, and some notes on the con-
servation status of the Callitrichidae. In: Marmosets and
Tamarins: Systematics, Behaviour and Ecology, A. B. Rylands
(ed.), pp. 11-77. Oxford University Press, Oxford.
Schneider, H., Sampaio, I., Harada, M. L., Barroso, C. M.
L., Schneider, M. P. C., Czelusniak, J. and Goodman, M.
1996. Molecular phylogeny of the New World monkeys
(Platyrrhini, Primates) based on two unlinked nuclear genes:
IRBP Intron 1 and epsilon-globin sequences. Am. J Phys.
Anthropol. 100: 153-179.
Silva Jr., J. S. and Noronha, M. de A. 1998. On a new species
of bare-eared marmoset, genus Callithrix Erxleben, 1777,
from central Amazonia, Brazil (Primates: Callitrichidae).
Goeldiana Zoologia (21): 1-28.
Stallings, J. D. and Mittermeier, R. A. 1983. The black-tailed
marmoset (Callithrix argentata melanura) recorded from
Paraguay. Am. J. Primatol. 4: 159-163.

18 Neotropical Primates 8(1), March 2000

Swofford, D. L. 1993. PAUP: Phylogenetic analysis using par-
simony (version 3.1). Illinois Natural History Survey,
Champaign, Illinois.
Tagliaro, C. H. 1997. A filogenia molecular na subfamflia
Callitrichinae (Platyrrhini, Primates): As relaq6es
intergen6ricas e intragendricas de Callithrix (sensu Schneider
et al., 1996). Doctoral thesis, Universidade Federal do Pari,
Tagliaro, C. H., Schneider, M. P C., Schneider, H., Sampaio,
I. C. and Stanhope M. J. 1997. Marmoset phylogenetics,
conservation perspectives, and evolution of the mtDNA
control region. Molec. Biol. Evol. 14: 674-684.
Thomas, 0. 1920. On mammals from the lower Amazonas
in the Goeldi Museum, Para. Ann. Mag. Nat. Hist. 7(14):
Van Roosmalen, M. G. M. and Van Roosmalen, T. 1997. An
eastern extension of the geographical range of the pygmy
marmoset, Cebuella pygmaea. Neotrop. Primates 5(1): 3-6.
Van Roosmalen, M. G. M., Van Roosmalen, T., Mittermeier,
R. A. and Fonseca, G. A. B. da. 1998. A new and distinc-
tive species of marmoset (Callitrichidae, Primates) from the
lower Rio Aripuand, state of Amazonas, central Brazilian
Amazonia. Goeldiana Zoologia (22): 1-27.
Wallace, A. 1852. On the monkeys of the Amazon. Proc. Zool.
Soc. London 1852: 107-110.

Neotropical Primates 8(1), March 2000 19


Alejandro Estrada1, Rosamond Coates-Estrada1, Lucia Castellanos2, Andr6meda Rivera2, Hector Gonz1lez3, Ana
Ibarra3, Yasminda Garcia3, David Mufioz3 and Berenice Franco3

lEstacidn de Biologia "Los Tuxtlas", Instituto de Biologia UNAM, Apdo 176, San Andris Tuxdta, Veracruz, Mixico, e-mail:
2Depto. de Biologia, Facultad de Ciencias, Universidad Nacional Autdnoma de Mixico, Mixico City.
3Divisidn de Ciencias Bioldgicas, Universidad Judrez Autdnoma de Tabasco, Villahermosa, Tabasco, Mixico.


Entre Enero y Mayo del 2000 se llevaron a cabo tres reconocimientos de la poblaci6n del mono aullador negro existente en la
selva del sitio arqueol6gico de Palenque, Chiapas, Mexico. Detectamos la presencia de 99 individuos repartidos en 13 tropas,
con la possible existencia de 3-5 tropas adicionales. La densidad ecologica estimada fue de 33 individuos/km2. El tamafio
medio de las tropas fue 7.6 (rango 2-13). Todos los avistamientos de los monos aulladores fueron en selva alta perennifolia y
en arboles >20 m en altura. Las densidades reportadas para Palenque son mis altas que aquellas reportadas para la misma
especie en Belice y Guatemala y en Quintana Roo en Mexico y otro contrast es que, en Palenque, la mayorfa de las tropas
(70%) eran multimacho. La protecci6n de un perimetro amplio (c. 1700 ha) alrededor de la selva que cubre el sitio arqueol6gico
por el gobierno Mexicano asegura la conservaci6n de la selva y de la poblaci6n del mono aullador negro existente en el sitio.


A survey of the population of the black howler monkey (A. pigra) present at the Mayan site of Palenque was conducted during
three visits between January and May of 2000. We detected the presence of 99 individuals in 13 troops and the possible
existence of 3-5 additional groups. Ecological density was estimated at 33 individuals/km2. Mean troop size was 7.6, ranging
from 2-13. All sightings of howler monkeys were in evergreen rain forest and in trees >20 m in height. The reported densities
and mean troop size are higher than those reported for the species in Guatemala, Belize and in central Quintana Roo, Mexico,
and the majority (70%) of the troops in Palenque were multimale. Protection of a large perimeter area (c. 1700 ha) around the
archeological site by the Mexican government ensures the conservation of the forest and of the black howler monkey population
present at the site.

Key words: Howling monkeys, Alouatta pigra, population census, Mexico


Las selvas del sur de Mexico alojan las representaciones mis
septentrionales de primates silvestres en el Neotr6pico. Tres
species de primates existen en estos habitats: Alouattapalliata,
A. pigra y Ateles geoffroyi. La primera y la iltima presentan
una distribuci6n amplia en el sur de Mexico y en al area
Mesoamerica. La distribuci6n geografica de A. pigra sin
embargo es restringida y s6lo se le encuentra en algunas parties
de los estados de Tabasco y Chiapas y es el dnico representante
del ginero en la peninsula de Yucatan (Smith, 1970; Horwich
y Johnson, 1986).

La distribuci6n geografica restringida de A. pigra en
Mesoamerica y la ripida conversion (4.5-12.5% annual,
Masera, 1986) de su habitat natural a pastizales y campos
agrfcolas coloca a las poblaciones de esta especie, end6mica a
nivel regional, bajo riesgo (Rylands et al., 1995). Por ejemplo,
en Tabasco, M6xico, en donde se encuentran poblaciones de
ambas species de Alouatta en simpatria, cerca del 60% de la
superficie del estado (24,141 km2) originalmente presentaba
selva alta perennifolia. Como resultado de la actividad

humana, entire 1977 y 1991, mas del 60% de esta vegetaci6n
desapareci6 a un ritmo annual de 600 km2 (SEMARNAP,

En Mexico, el studio de la poblaci6n, ecologia y
comportamiento de A. palliata ha sido investigado durante
various afios en la region de Los Tuxtlas, Veracruz (Estrada y
Coates-Estrada, 1996). Sin embargo, informaci6n sobre estos
aspects paraA. pigra es inexistente. Hasta ahora s6lo contamos
con algunos reconocimientos breves (Horwich y Johnson,
1986; Watts y Rico-Gray, 1987; del Campo y Jorgenson,
1998) de los que s6lo uno, llevado a cabo en Muchunuk,
Quintana Roo, en la peninsula de Yucatan, podria ser
considerado como sistemAtico (Gonzales-Kirchner, 1998).

En el caso de Guatemala, studios breves se han efectuado en
Tikal (Coelho et al., 1976; Schlichte, 1978). Los unicos
studios detalladas sobre poblaci6n, ecologia y conduct de
A. pigra, acoplados a un program de conservaci6n sostenido
a travis de los afios, es aquel realizado en solo dos localidades
en todo el rango de distribuci6n geogrifica de A. pigra y ambas
localidades se ubican en Belice (Silver et al., 1998; Ostrum et
al., 1999).

20 Neotropical Primates 8(1), March 2000

Considerando la protecci6n brindada por el gobierno
Mexicano, federal y estatal, a las selvas que rodean muchos de
los sitios arqueol6gicos de la zona Maya en el sur de M6xico
con el objeto de proteger el patrimonio cultural que ahl se
resguarda, y utilizando como modelo los studios poblacionales
y ecol6gicos que hemos realizado en las selvas de Los Tuxtlas
con la especie A. palliata, hacia fines de 1999 iniciamos una
investigaci6n de las poblaciones del mono aullador negro,
Alouattapigra, existentes en la selva del Parque Nacional Zona
Arqueol6gica de Palenque, Chiapas.

Esta investigaci6n tiene como prop6sito recabar informaci6n
precisa acerca del tamafio de la poblaci6n del mono aullador
negro en la zona del Parque y Areas aledafias, recabar
informaci6n acerca de sus hAbitos alimenticios y necesidades
de espacio y diagnosticar la manera en que estos primates
participan en la dinimica del ecosistema selvAtico. Dicha
informaci6n enriquecerA los bancos de datos sobre A. pigra y
sera tambien fundamental para promover la conservaci6n de
la especie en los alrededores y en otras localidades y para
comprender las respuestas de sus poblaciones a los cambios
antropog6nicos en la distribuci6n de su habitat natural.

Entre Enero y Mayo del 2000 invertimos, en tres sesiones
diferentes, 30 dias efectivos de trabajo de campo en el sitio
con el objeto de determinar el ndmero de tropas de aulladores
que ahi existen y su distribuci6n en el espacio. Un nmmero de
dias adicionales fueron invertidos en la recopilaci6n de
informaci6n contextual y llevando a cabo observaciones sobre
el comportamiento de tropas selectas. En este trabajo
presentamos informaci6n derivada de los reconocimientos
demogrdficos de la poblaci6n de A. pigra que existe en la selva
de la zona arqueol6gica de Palenque.


El Parque Nacional Zona Arqueol6gica de Palenque se localiza
al noroeste del estado de Chiapas formando parte del

Figure 1. Localizaci6n del Parque Nacional Zona Arqueol6gica de
Palenque en el estado de Chiapas, M6xico. La zona sombreada
represent vegetaci6n selvitica (597 ha), areas con rayas horizontales
= a vegetaci6n secundaria (300 ha). Areas en blanco son pastizales
(874 ha). Linea punteada = terraceria. ZA = zona arequeol6gica.

municipio del mismo nombre y decretado presidencialmente
el 26 de Junio de 1981. El Parque cuenta con una extension
territorial de 1771 hectareas y se localiza entire los paralelos
17o27'05" y 17o30'05" de latitud Norte y a los 92001'30"
y 92004'42" de longitud Oeste.

El dima del area donde esti ubicado el sitio ha sido clasificado
como cilido-hdmedo (Garcia, 1981). La precipitaci6n media
annual es de 2156 mm con una distribuci6n estacional a travys
del afio. La precipitaci6n media mensual es significativamente
menor entire Enero y Abril (promedio = 62 18 mm)
comparada con la media para el resto de los meses del afio
(240 106 mm). El mes mis seco es Abril con 49 mm y el
mes mas lluvioso es Septiembre con 433 mm. La temperature
media annual es de 26C. El mes mis caluroso es Mayo con
una temperature de 29.7oC y el mes mis frfo es Enero con

La topograffa de la zona en donde esta distribuida la selva del
sitio es abrupta, con pendientes en algunas areas de hasta 50%,
pero es possible encontrar pendientes del 2% en los alrededores
de los templos principles. En la zona del Parque con cobertura
de vegetaci6n selvatica, el gradiente altitudinal va de los 150
a 500 m sobre el nivel del mar.

La vegetaci6n selvitica reportada para el Parque Nacional de
Palenque es conocida tdcnicamente como selva alta
perennifolia y cubre aproximadamente una superficie de 597
hectireas. Otras 300 hectireas presentan vegetaci6n
secundaria, representando distintas fases en la regeneraci6n
de la vegetaci6n selvitica y el resto de la superficie del Parque
consiste en pastizales inducidos (Dfaz Gallegos, 1996) (Fig. 1).

Studios floristicos de la vegetaci6n selvitica reportan la
existencia de 510 species de plants, distribuidas en 351
gdneros representando a 136 families botinicas (Diaz Gallegos
1996). De estas, las mejor representadas son Leguminosae
con 47 species y Rubiaceae con 27 species. La forma de
vida de crecimiento dominant en el parque son los Arboles,
representados por 241 species. La selva del Parque Nacional
sobresale por presentar una diversidad floristica alta en relaci6n
con otros sitios en el sur de M6xico y Sudamerica (Diaz
Gallegos, 1996). En relaci6n a la estructura de la vegetaci6n,
parece existir una distribuci6n regular de species arb6reas a
lo largo del estrato vertical, predominando en el estrato de los
0-10 m species como Rinorea guatemalensis y Astrocaryum
mexicanum, mientras que en los estratos superiores por arriba
de los 20 m predominan species como Vatirea lundelli,
Guatteria anomala, Manilkara zapota, Brosimum alicastrum,
Quararibeafunebris y Ficus spp. (Diaz Gallegos, 1996).

Censo de la poblacidn de monos aulladores: Registros aclisticos
A travys de sesiones matutinas (5-7 a.m.) y vespertinas (6-8
p.m.) equipos de 2-3 personas se ubicaron en puntos
estratigicos en las estructuras arqueol6gicas mas altas del sitio.
A partir de estos puntos se registry el intercambio de coros
entire tropas de monos aulladores, indicando la orientaci6n
de los aullidos en grados en relaci6n a los cuatro puntos
cardinals y se estim6 la distancia a la que los bramidos fueron

NeotropicalPrimates 8(1), March 2000

escuchados. La ubicaci6n de las tropas asi determinada se
transfiri6 a un mapa a escala de la zona de trabajo.

Censo de la poblacidn de monos aulladores: Registros visuals
Estos tuvieron como meta lograr una verificaci6n de la
ubicaci6n de las tropas de monos posterior al registro ac6stico
matutino, identificaci6n de las tropas y obtener conteos
precisos sobre su composici6n por edades (adulto, juvenile,
infante) y sexos. La localizaci6n de las tropas de saraguatos
fue facilitada por la accesibilidad permitida a diferentes parties
del area selvatica por el sistema de veredas existentes en el
Parque. Las tropas asi encontradas fueron confirmadas en
cuanto a su ubicacid6n relative en relaci6n a rasgos topogrificos
y/6 arqueol6gicos y respect a la ubicaci6n relative de otras
tropas. Confirmaci6n de la identidad de las tropas y de su
tamafio y composici6n por edades y sexos fue llevado a cabo a
trav6s de contacts repetidos con cada una en dias

Como parte complementaria a estos sondeos demogrificos,
tambien se realizaron observaciones preliminares sobre el
comportamiento alimenticio y actividades generals de los
individuos en tres de las tropas detectadas. En este caso
individuos representatives de cada clase de edad y de cada
sexo fueron observaron durante periods fijos de tiempo. En
las observaciones del comportamiento alimenticio se tom6 nota
de la parte de la plant consumida (hojas, frutos y/6 hojas
j6venes 6 maduras, frutos j6venes 6 maduros, flores, otros) y
se trat6 de identificar la especie.


Esfuerzo de muestreo
El esfuerzo de campo result en la acumulaci6n de 30 dfas de
trabajo y 672 horas hombre invertidas en el reconocimiento
demogrifico de los monos aulladores en el Parque. De estas,
el 40% fue invertido en la triangulaci6n de la ubicaci6n
relative de las tropas de aulladores a traves de registros actisdcos.
El 60% restante se invirti6 en los conteos de las tropas sobre
el terreno. Aproximadamente 60 horas hombre adicionales
fueron invertidas en observaciones del comportamiento y
actividades generals de tres tropas de aulladores.

La poblacidn de monos aulladores
Los reconocimientos resultaron en un conteo de 99 monos
aulladores repartidos en 13 tropas. Cerca de 3-5 tropas
adicionales fueron escuchadas a 1.5-2.0 km. de distancia de
los templos principles en direcci6n Oeste y Suroeste, pero
no fue possible su localizaci6n. En esta poblaci6n los individuos
adults contribuyeron al 62% de los conteos, 19% fueron
juveniles y 19% infants. Los machos adults contribuyeron
al 35% de los conteos y las hembras adults al 27%.

El tamafio medio de las tropas fue calculado en 7.6 individuos
(rango 2-11) y la composici6n promedio de las tropas fue de
2.7 machos adults, 2.0 hembras adults, 1.6 machos
juveniles, 1.3 hembras juveniles y 1.6 infants. La relaci6n de
machos a hembras entire los adults fue de 1: 0.74 y la relaci6n

Tabla 1. Tropas de mono aullador negro detectadas en el Parque
Nacional Zona Arqueol6gica de Palenque, Chiapas, Mdxico.

Troop AduJtos luvcniles Infanmes Total
Mi F M F

1 3 2 1 1
1 2 I i
A :i 2 I 1 2 9

I.A 2 2 2 2 2 0

I- \ 3 2 I I I
Total 35 26 1 S 19 99

Figura 2. Localizaci6n aproximada de las tropas de monos aulladores
detectadas en la zona selvitica del Parque. El area mostrada estA
cubierta, a excepci6n de las estructuras arqueol6gicas principles en
la vencindad del estacionamiento (E), por vegetaci6n selvitica. Lineas
punteadas = arroyos, lines negras delgadas = senderos. Area
representada aprox. 500 ha. El gradiente altitudinal de Norte a Sur
es de 100 500 m sobre el nivel del mar.

22 Neotropical Primates 8(1), March 2000

de adults a no adults fue de 1:0.62. La densidad ecol6gica
estimada para la poblaci6n de A. pigra en el Parque fue de 33
individuos por km2 y la poblaci6n total estimada fue de 200
monos aulladores repartidos en unos 15-18 grupos.

Aunque preliminares, los datos sobre actividades generals
indicaron que los aulladores invierten el 52% de su tiempo
en descansar, el 28% en la actividad alimenticia, el 13% en
interacciones sociales, el 5% en actividades locomotoras y el
1% en desplazamientos sincronizados de una zona a otra
dentro del Parque.

Todos los avistamientos de las tropas de aulladores fueron en
selva alta perennifolia y en arboles >1.3 m en d.a.p. y >20 m
de altura. Algunas de las species de las cuales fueron
observados alimentdndose fueron Brosimum alicastrum, Ficus
spp., Poulsenia armata y Spondias mombin entire otras. El 50%
de las parties consumidas fueron hojas j6venes y el restante
50% frutos maduros de species como las sefialadas


Los datos obtenidos sobre el tamafio de la poblaci6n de
Alouatta pigra en el Parque Nacional Palenque deben ser
considerados como preliminares. Continuidad en el trabajo
de campo permitira obtener informaci6n sobre la consistencia
y variabilidad en los rasgos demogrificos presentados en este
report para la poblaci6n del mono aullador negro present
en el Parque.

Tomando esto en cuenta, la densidad estimada de la poblaci6n
de monos aulladores en el Parque Nacional Palenque de 33
individuos km2 es mas alta que aquella que ha sido reportada
para la misma species en Belice (8-22 individuos/km2) (Bolin,
1981; Horwich y Gerhardt, 1983) y en Tikal en Guatemala
(5-9 individuals/km2) (Coelho et al., 1976; Schlichte, 1978).
El dinico otro sitio en Mexico en donde aspects demograficos
de A. pigra han sido investigados es la selva de Muchunuk en
el centro de Quintana Roo y en donde se reportan densidades
mis bajas (16.53 individuos/km2) (Gonzales-Kirchener, 1998)
que aquellas detectadas para la especie en Palenque. En
comparaci6n, en Los Tuxtlas, Veracruz, a 500 km al norte de
Palenque, las poblaciones de la otra especie de saraguato en el
pais, Alouatta palliata, se presentan en densidades de 23
individuos/km2 en selvas no perturbadas por la actividad
humana (Estrada, 1982; Estrada y Coates-Estrada, 1996).

La alta densidad de A. pigra detectada en Palenque parece
contradecir la aseveraci6n de que esta especie present las
densidades mis bajas reportadas para las species del genero
Alouatta (Crockett y Eisenberg, 1987; Gonzales-Kirchener,
1998). El tamafio miximo de las tropas del mono aullador
detectadas en Palenque (N = 11 individuos) y el tamafio
promedio de las tropas de 7.6 individuos tambien son mis
altos que aquellos valores reportados para la misma species
en Belice y Guatemala, donde el tamafio de las tropas varia
de 4.4 a 6.3 individuos Coelho et al., 1978; Bolin, 1981;

Horwich y Gerhardt, 1983; Ostro et al., 1999) y para el
centro de Quintana Roo (3.16 individuos) (Gonzales-
Kirchner, 1998).

Otro contrast con los datos publicados provenientes de
Guatemala y Belice es que la mayorfa (70%) de las tropas
detectadas en Palenque estaban compuestas por various machos
adults y un numero menor de hembras adults. En
Guatemala y Belice diferentes autores de manera consistent
reportan tropas unimacho (un solo macho adulto y varias
hembras adults) (Coelho et al., 1976; Schlichte, 1978;
Horwich y Johnson, 1986).
Consideramos que estos contrastes podrian ser el resultado
de la pobreza de datos que existe sobre A. pigra en la literature.
La informaci6n hasta ahora disponible proviene de studios
realizados en solo tres sitios (Tikal en Guatemala, Bermuda
Landing en Belice y Muchunuk en Quintana Roo, Mexico).
Obviamente se necesitan mas studios del mono aullador
negro en otros sitios dentro de su rango de distribuci6n
geografica para documentary el rango de variaci6n en
parametros demograficos como densidad ecol6gica y tamafio
y estructura de las tropas.

La selva de Palenque posee las caracterfsticas estructurales y
los conjuntos de species arb6reas adecuadas para el
sostenimiento de la poblaci6n de monos aulladores. Los arboles
del dosel superior de la selva son los principles contribuyentes
al area basal total registrada en los reconocimientos de la
vegetaci6n que se han llevados a cabo en el sitio (Diaz
Gonzales, 1996). Estos irboles pertenecen a species de las
families botinicas Moraceae (por ej., Ficus spp, Brosimum
alicastrum, Poulsenia armata), Sapotaceae (por ej., Pouteria
campechiana, Pouteria zapota), Anacardiaceae (por ej.,
Spondias mombin) y Lauraceae (por ej. Nectandra spp., Ocotea
spp.) entire otras, reportadas como fuente important de hojas
y de frutos en la dieta de los monos aulladores en el Neotrt6pico
(Estrada y Coates-Estrada, 1993).

Nuestros reconocimientos preliminares del area que rodea al
Parque Nacional Palenque indican que much de la vegetaci6n
selvitica ha desaparecido y ha sido convertida a pastizales. Por
consiguiente la selva present en el Parque Nacional Palenque
es de importancia estrategica ya que su conservaci6n ha
permitido la conservaci6n de una representaci6n de este
ecosistema y de representantes de Alouatta pigra en la porci6n
mis nororiental de su distribuci6n geografica en el estado de
Chiapas en el sureste de Mexico.


Agradecemos el permiso otorgado por el Director del Parque
Nacional Zona Arqueol6gica de Palenque, Lic. Juan Antonio
Ferrer Aguilar, para la realizaci6n de estos studios. Tambidn
agradecemos al Arque6logo Edwin Barnhardt el habernos
facilitado un mapa de referencia que fue fundamental para
lograr una orientaci6n expedita de nuestro grupo en las
distintas zonas del sitio. Agradecemos al Sr. John Scott su

Neotropical Primates 8(1), March 2000


Barnhardt, E. 1998. Palenque Mapping Project. Field Sea-
son Report. Technical report. Foundation for advancement
of Mesoamerican Studies, Inc. FAMSI Webpage.
Barnhardt, E. 1999. Palenque Mapping Project. Field Sea-
son report. Foundation for Avancement of Mesoamerican
Studies, Inc. FAMSI Webpage.
Bolin, I. 1981. Male parental behavior in black howler mon-
keys (Alouatta palliata pigra) in Belize and Guatemala. Pri-
mates 22: 349-360.
Coelho, A. M., Bramblett, C. A., Quick, L. B. y Bramblett,
S. 1976. Resource availability and population density in pri-
mates: A socio-bioenergetic analysis of the energy budgets
of Guatemalan howler and spider monkeys. Primates 17:
Crockett, C. M. y Eisenberg, J. E 1987. Howlers: Variation
in group size and demography. En: Primate Societies, B. B.
Smuts, D. L. Cheney, R_ M. Seyfarth, R. W. Wrangham y
T. T. Struhsaker (eds.), pp. 56-68. University of Chicago
Press, Chicago.
Del Campo P L. A. y Jorgenson, J. P 1998. Notes on the
distribution and conservation status of spider and howler
monkeys in the state of Quintana Roo, Mexico. Primate
Conserv. (18): 25-29.
Diaz Gallegos, J. R. 1996. Estructura y Composici6n
Floristica de la Vegetaci6n del Parque Nacional Zona
Arqueol6gica de Palenque, Chiapas, Mexico. Tesis,
Universidad Juairez Aut6noma de Tabasco, Villahermosa,
Tabasco, Mexico.
Estrada, A. 1982. Survey and census of howler monkeys
(Alouattapalliata) in the rainforest of Los Tuxtlas, Veracruz,
Mexico. Am. J. Primatol. 2: 363-372.
Estrada, A. 1999. Comportamiento Animal: el Caso de los
primates. Fondo de Cultura Econ6mica. Serie la Ciencia
desde Mixico. (65).
Estrada, A. y Coates-Estrada, R. 1988. Tropical rain forest
conversion and perspectives in the conservation of wild pri-
mates (Alouatta and Ateles) in Mexico. Am. J. Primatol. 14:
Estrada, A. y Coates-Estrada. R. 1993. Aspects of ecological
impact of howling monkeys (Alouatta palliata) on their
Habitat: A Review. En: Avances en Estudios Primatoldgicos
en Mixico, A. Estrada, E. Rodriguez Luna, R. Lopez- Wilchis
y R. Coates-Estrada (eds.), pp. 87-117. Asociaci6n Mexicana
de Primatologfa, A.C. y Patronato Pro-Universidad
Veracruzana, A. C. Xalapa, Veracruz, Mexico.
Estrada, A. y Coates-Estrada, R. 1996. Tropical rain forest
fragmentation and wild populations of primates at Los
Tuxtlas. Int. J. Primatol. 5: 759-783.
Estrada, A. y Coates-Estrada, R. 1995. Las Selvas Tropicales
de Mexico. Fondo de Cultura Econ6mica. Serie la Ciencia
desde Mixico. (132).
Gonzales-Kirchner, J. P. 1998. Group size and population
density of the black howler monkey (Alouatta pigra) in
Muchukux forest, Quintana Roo, Mexico. Folia Primatol.
69: 260-265.

Horwich, R. y Gerhardt, K. 1983. Roaring rhythms in black
howler monkeys (Alouattapigra) of Belize. Primates 24: 290-
Horwich, R. y Johnson, E. W 1986. Geographic distribu-
tion of the black howler monkeys (Alouatta pigra) in Cen-
tral America. Primates 27: 53-62.
Masera, 0. R. 1996. Deforestaci6n y degradaci6n forestal
en Mexico. Documento de Trabajo. Grupo
Interdisciplinario de Trabajo de Tecnologia Rural Apropiada
19: 1-15.
Ostrum, L. E. T., Silver, S. C., Koontz, E W., Young, T. P. y
Horwich, R. H. 1999. Ranging behavior of translocated
and established groups of black howler monkeys Alouatta
pigra in Belize, Central American. Biol. Conserv. 87: 181-
Rylands, A., Mittermeier, R. A. y Rodrfguez-Luna, E. 1995.
A species list for the New World primates (Platyrrhini): dis-
tribution by country, endemism, and conservation status ac-
cording to the Mace-Land system. Neotrop. Primates 3
(suppl.): 114-164.
Silver, S. C., Ostro, L. E. T., Yeager, C. P. y Horwich, R. 1988.
Feeding ecology of the black howler monkey (Alouattapigra)
in northern Belize. Am. J. Primatol. 45: 263-279.
Schlichte, H. 1978. A preliminary report on the habitat uti-
lization of a group of howler monkeys (Alouatta villosapigra)
in the national park ofTikal, Guatemala. En: The Ecology of
Arboreal Folivores, G. G. Montgomery, (ed.), pp.551-561.
Smithsonian Institution Press, Washington, DC.
Smith, J. D. 1970. The systematic status of the black howler
monkeys, Alouatta pigra Lawrence. J Mammal 51: 358-
Watts, E. y Rico-Gray, V. 1987. Los primates de la peninsula
de Yucatin, Mexico: studio preliminary sobre su
distribuci6n actual y estado de conservaci6n. Bidtica 12:

24 Neotropical Primates 8(1), March 2000


Robert B. Wallace', R. Lilian E. Painter, Damian I. Rumiz and Andrew B. Taber
Wildlife Conservation Society, 185th Street and Southern Boulevard, Bronx, New York, 10460, USA.
'Address correspondence to: Robert B. Wallace, Wildlife Conservation Society Bolivia, Casilla 3-35181, San Miguel, La Paz,
Bolivia, e-mail: .


We document primate diversity in the Rfos Blanco y Negro Wildlife Reserve and compare encounter rates between primate
species and the field sites examined. We also present an encounter rate for overall primate biomass at each field site. Primate
diversity encountered within the Reserve appears typical of other sites in eastern lowland Bolivia but interesting patterns in
species distribution and abundance are discussed.

Key Words: Primate diversity, Census, Amazon, Bolivia


Until recently, little information had been published on the
distribution of primate species in the Bolivian eastern low-
lands. Primate surveys have been restricted to two protected
areas (Braza and Garcia, 1988; Garcia and Tarifa, 1988;
Wallace et al, 1998), with other distributional information
coming from specimens collected at scattered locations (Brown
and Rumiz, 1985; Anderson, 1997). The Rios Blanco y Ne-
gro Wildlife Reserve (RBYNWR) covering 14,239 km2 (Fig-
ure 1), was created in 1990, and forms part of the Bajo Paragua
Forestry Reserve (BPFR) of 33,882 km2. The reserve is di-
vided into forestry concessions that reach areas of up to 3,113
km2 (FAN/PL480/WCS, 1994), and which have been selec-
tively logged for three high-value timber species; Swietenia
macrophylla, Amburana cearensis, and Cedrela odorata. Selec-
tive logging has intensified in lowland Bolivia in recent years
and is typically associated with technically illegal, heavy hunt-
ing of larger wildlife species.

In terms of biodiversity, the area was almost unexplored until
1992-1993 when multi-disciplinary surveys were carried out
as part of a project designed to produce a management plan
for the reserve (FAN/PL480/WCS, 1994). These surveys were
necessary to provide baseline data for future ecological moni-
toring, as well as to assess the impact of human activities on
biodiversity within the reserve. In this report we document
primate diversity in the Rios Blanco y Negro Wildlfe Reserve
and compare group and individual encounter rates between
primate species and the field sites examined. We also calcu-
late an "encounter rate" for overall primate biomass at each
field site.


The Rios Blanco y Negro Wildlife Reserve (RBYNWR) is situ-
ated in the north-west of Santa Cruz Department, and in-
cludes two distinct biogeographical regions: the Beni alluvial

plain which lies west of the Rio Blanco, and the Brazilian Shield
which forms the eastern part of the Reserve (Fig. 1). The for-
ests found within the reserve are broadly classified as either
Humid Forest of the Brazilian Shield or Lowland Humid For-
est (Killeen et al., 1993). The Wildlife Reserve was sparsely
populated during successive rubber booms but has been
largely uninhabited since the 1950's. Eight survey sites were
selected within the reserve. Five sites were directly accessible
through existing logging roads; Arroyo Chuchui, San Mar-
tin, San Luis, Oquiriquia, and El Tutumo. Three others were
accessible by river and/or plane; Perseverancia, Pajaral, and
Rio Negro de Caimanes (Figure 1). Further information re-
garding survey sites is provided in Table 1.


Each field site was visited for approximately three weeks. Line
transects were employed to survey the areas using standard
techniques (Burnham et al., 1980; Brockelman and Ali,
1987; Buckland et al., 1993). With sufficient transect en-
counters, density estimates can be calculated using this meth-
odology. However, since at least 40 encounters of each spe-
cies are required for reliable estimates, many researchers have
been restricted to presenting relative abundances using en-
counter rates (Branch, 1983; Mate and Colell, 1995). Nev-
ertheless, encounter rates are a useful long-term monitoring
tool and can also allow site comparisons, providing the area
censused along a standard transect length is similar (Wallace
et al., 1998).

Diurnal transects were run by two observers in fair weather
conditions between 06:00 to 11:30 and 15:00 to 18:00, along
existing logging trails or roads and/or newly established trails
cut by the survey team. Transect speed ranged from 1-2 km
per hour and depended on trail conditions and associated
noise levels. Periods of walking were regularly interspersed
with brief "listening stops" in order to increase the probabil-
ity of detecting more cryptic species. Nocturnal censuses were
conducted over shorter distances between 19:00 and 23:00

Neotropical Primates 8(1), March 2000 25

Figure 1. Location of the eight survey sites within the Rios Blanco y Negro Wildlife Reserve, Santa Cruz Department, Bolivia.

and were generally walked at a slower speed. Details regard-
ing the diurnal transect and nocturnal census effort at each
site are presented in Table 1. The following information was
recorded for all groups of primates encountered on transects:
species, group size (and where possible age/sex composition),
date and time detected, observation duration, transect posi-
tion, habitat type, and the perpendicular distance from the
transect trail to the estimated geometric centroid of the group.
Incidental non-transect observations and primate vocalizations
also provided information on primate diversity.

Results are expressed as relative abundances, calculated as the
number of group encounters for each primate species per 10
km censused (groups/10 kin). For Ateles, Cebus, and Saimiri
the transect width was truncated at 40 m from the trail for all
sites considered. Alouatta, Aotus and Callithrix sightings were
truncated at 20 m from the trail due to the more cryptic na-
ture of these species. These analytical norms were established
to ensure that the transect width, and hence the area censused
at each site, was similar. Individual encounter rates are ex-
pressed as the number of observed individuals of each species
encountered per 10 km censused (inds/10 km). Primate bio-
mass encounter rates for each site are expressed as the ob-
served primate biomass per 10 km censused (kg/10 km). Pri-
mate body weights were taken from Ford (1994). Since data

on group composition was difficult to obtain during censuses,
and weights for other age/sex classes unknown, for the pur-
poses of analysis all animals were assumed adult, and adult
sex ratios were assumed 1:1 for all species at all sites. Statisti-
cal procedures follow Siegel and Castellan (1988), and sig-
nificance was set at the 0.05 level.


A total of seven non-human primate species were registered
within the reserve: black-tailed marmoset (Callithrix
melanura), owl monkey (Aotus azarae), Bolivian squirrel mon-
key (Saimiri boliviensis), brown capuchin monkey (Cebus
apella), black howler monkey (Alouatta caraya), red howler
monkey (Alouatta seniculus), and black spider monkey (Ateles
chamek). In addition, at three sites within the reserve
(Perseverancia, Rio Negro de Caimanes, Oquiriquia) groups
of Cebus were observed which included only very pale and
more slender individuals. These three sightings may repre-
sent white-fronted capuchin monkeys (Cebus albifrons), how-
ever, Cebus apella is renowned for within population varia-
tion in pelage colour (Emmons and Feer, 1990). Thus, until
specimens and/or further observations are available we pre-
fer to treat these sightings as Cebus apella.

Table 1. Transect kilometres and location of the eight field sites within the Rios Blanco Y Negro Wildlife Reserve, Santa
Cruz Department, Bolivia.
Surney Site Diurnal Nocturnal Biophysical Location Surves Period
ikmi ILkman Region'
Pir i':.eo.- i] Y SIs [u_ B5 1.3 'S 2 -" \B lune ld'. 19C7
P'iiri It, ibS/BAP I -. 1,; .' I '.X August 1902
.krrmo Chuhuit S 8 13 BS I ,. 35 u2 -.5 '.Y October 19"-2
,Jr. 1 lr-in t30 ;;.I. B 14..7',0 ': '. ',' Nuovember IVQ2
RJ, N ,) Camrn.tr.,: 161.6 uBA0 -. -. S -1 '6 'X lunch 1903
Sjr Luis lOS '1; 2B Bc Fiure I Augut.r 1)93
O-quiriquia '3..2 B_ ) 6.i.3 -o:l '.A Sepc.rmbcr 19'73
El Turumo 9'.' 21 5 bS/BA' I.0 .. c.. hi '. October 19',3
iBS = Brazilian Shield, BAP = Beni Alluvial Plain, BS/BAP = Ecotone between Brazilian Shield and Beni Alluvial Plain.

26 Neotropical Primates 8(1), March 2000

Four species were encountered at all of the sites visited within
the reserve: Callithrix melanura, Aotus azarae, Cebus apella,
and Ateles chamek. Howler monkeys appear to have a more
limited distribution: Alouatta caraya was only observed in the
semideciduous forests at Pajaral, and Alouatta seniculus was ob-
served in seasonally inundated forest at Perseverancia, Pajaral,
Rio Negro de Caimanes, and Arroyo Chuchui. Squirrel mon-
keys (Saimiri boliviensis) were registered at four sites:
Perseverancia, Pajaral, Rio Negro de Caimanes and El Tutumo.
The most diverse sites were Pajaral with seven primate species,
and Perseverancia and Rio Negro de Caimanes with six spe-
cies (see Table 2).

Group and individual encounter rates are presented in Table
2, and details for each species are as follows:

Callithrix argentata: Individual encounter rates were signifi-
cantly different between sites (Chi-squared = 65.99, d.f.7,
p<0.001), and encounter rates were low for all sites consid-
ered, both in terms of observed groups (0.1-0.5 grps/10 km)
and individuals (0.1-2.8 inds/10 km).

Aotus azarae- Insufficient encounters prevented a statistical test,
but group and individual encounter rates for owl monkeys
varied among sites (0.4-3 grps/10 km, 0.8-7.4 inds/10 km).
Arroyo Chuchui had a group encounter rate over twice that
of any other site, although the individual encounter rate for
El Tutumo (7.4 inds/10 km) was higher than that of Arroyo
Chuchui (5.2 inds/10 km) due to site differences in mean
group size.
Saimiri boliviensis: Only observed on transects at two sites,
individual encounter rates varied from 4.9-9.8 inds/10 km.
Alouatta seniculus: Only observed on transects at three sites,
individual encounter rates varied from 0.3-1.8 inds/10 km.
Alouatta caraya: This species was only registered during non-
transect observations at Pajaral.
Cebus apella: In terms of individuals, capuchin monkeys were
consistently the most frequently encountered diurnal primate,
and individual encounter rates (4.4-14.7 inds/10km) were sig-
nificantly different between sites (Chi-squared = 97.01, d.f.7,
p<0.001). Group encounter rates varied from 0.6-1.6 grps/
10 km censused. The three highest sites for both group and
individual encounter rates were Pajaral, Rio Negro de

Caimanes and Arroyo Chuchui.
Ateles chamek: Individual encounter rates were also signifi-
cantly different between sites for spider monkeys (Chi-squared
= 357.95, d.f.7, p<0.001). Both group (0.1 0.3 grps/10km)
and individual (0.3-2.3 inds/lOkm) encounter rates were low
for all sites considered except Arroyo Chuchui, where indi-
vidual encounter rates reached 11.5 inds/10km censused.

Primate biomass encounter rates varied between 15.3-148.2
kg/10 km censused (mean = 49.2, SD 41.8). Notably, at
Arroyo Chuchui (148.2 kg/10 km) we encountered over
double the primate biomass of any other site, the nearest be-
ing Pajaral (57.2 kg/10 km). Four sites (Perseverancia, San
Martin, Rio Negro de Caimanes and El Tutumo) had similar
primate encounter rates (36.5-38.2 kg/10 km). Finally, the
primate biomass encounter rates at San Luis (15.3 kg/10 km)
and Oquiriquia (23.5 kg/10 km) were relatively low.


Primate diversity within the Rios Blanco y Negro Wildlife Re-
serve, at seven species, appears typical of other protected ar-
eas in eastern lowland Bolivia (Braza and Garcia, 1988; Garcia
and Tarifa, 1988; Wallace et al., 1998). A notable absence
was the dusky titi monkey (Callicebus donacophilus) which is
frequently observed in forests bordering the city of Santa Cruz
to the south of the reserve, but appears absent in northern
Santa Cruz Department (Braza and Garcia, 1988; Wallace et
al., 1998; this study). Nevertheless, in the Lago Caiman re-
gion C. brunneus is present on the Brazilian side of the Itefiez
River (Wallace et al., 1996).

To our knowledge the Bolivian squirrel monkey (Saimiri
boliviensis) has as yet not been registered east of Perseverancia
in Santa Cruz Department, although anecdotal reports sug-
gest it may reach as far east as the upper San Martin River.
Where registered, squirrel monkey troops were frequently
observed in close association with Cebus apella troops. This
association has been noted in previous studies in the Neotropics
(Terborgh, 1983).
This study also underlines the apparent rarity of howler mon-
keys in the expansive seasonally evergreen forests of the Bra-
zilian Shield. Alouatta seniculus appears to be confined to flood-
plain forest where it is found at relatively low densities (Wallace

Table 2. Primate diversity, group (and individual) encounter rates (per 10km), and primate biomass encounter rates at eight sites within
the Rios Blanco y Negro Wildlife Reserve, Santa Cruz Department, Bolivia.

Spec ies Perse~erancia Pajaral Arroyo San Rio Negro cie San Luis Oquiriqtua El
Cbucbui Martin Caimanes Tun.mio

0i i'l 8.31 3 1'7 1 3u -ifi 9910i, 1.5 (90)i 1-).ui..1 12. 1-t o. w31

"131 u.-,il1.8i 1 (1311.21

TouL Species 5 5 -1 6 -1
['EE~g~ I''k,' ~ 252 1-18.2 1. 65 -~ 35 4; 3
*Spe~i. rcgli.cued in -A-Ernin~ccr codun 'BE R P~ronale Bsomn.,' Encounere R.i,c

Neotropical Primates 8(1), March 2000 27

etal, 1998). The black howler monkey (Alouatta caraya) also
appears to be restricted to certain habitats, mainly
semideciduous forests. The restrictive use of habitats by this
genus may be due to the floristic composition of the forests,
and/or poor nutrient background of the soils, which may limit
folivory as a dietary niche (Wallace et al., 1998). These fac-
tors are certainly important in explaining variations inAlouatta
abundance in lowland Brazilian Amazonia (Peres, 1997).

Apart from the presence or absence of Saimiri and Alouatta
spp., primate community structure also varied in the relative
abundances of the more widespread species: Callithrix, Aotus,
Cebus and Ateles. Although encounter rates varied for
Callithrix, they were rare at all sites, indicating low popula-
tion densities for this region. The variations in Aotus encoun-
ter rates should be treated with some caution, as censuses were
run during different phases of the moon at different sites.
Aotus are more conspicuous around the full moon due to in-
creased calling behavior (pers. obs.). Cebus encounter rates
also varied and capuchins were the most commonly encoun-
tered diurnal primate at all visited sites. Previous studies have
demonstrated a considerable adaptability in habitat use by
Cebus apella and have suggested that this is linked to their
generalist diet (Terborgh, 1983). This adaptability may ac-
count for their numerical dominance at the sites considered

Due to their superior body weight, spider monkey density
variations account for much of the variation in overall pri-
mate biomass encounter rates. Ateles were infrequently en-
countered at most sites, suggesting that typically spider mon-
key population densities are relatively low for this region. Ar-
royo Chuchui features markedly high Ateles encounter rates
and is characterized by the presence of rocky escarpments
that result in a local diversity of forest habitats. Surveys con-
ducted at Lago Caiman in Noel Kempff Mercado National
Park (Wallace et al, 1998) revealed a high primate biomass
encounter rate (109.4 kg per 10 km censused) which was
also due to increased Ateles encounter rates (see Table 3). Lago
Caiman features a prominent quarzite escarpment, again re-
sulting in local habitat diversity. We suggest that local habitat
diversity provides an abundance of fruit resources at differ-
ing times of the year, resulting in increased Ateles densities.
Phenology results and ranging patterns from a recent long-
term autoecological study on Ateles at Lago Caiman support
this hypothesis (Wallace, 1998).

Aside from these anomalies, it appears that primate biomass
encounter rates are greater in the westernmost sites, which
are either on or bordering the Beni alluvial plain. Surveys
from other protected areas in lowland Bolivia support this
generalization. The Beni Biosphere Reserve, situated c.190
km west of the western border of the Wildlife Reserve, in the
Beni Department, has a high primate encounter rate of 72.8
kg per 10 km censused (see Table 3). Similarly, surveys in the
nearby Chimane Forest revealed an even higher primate en-
counter rate (146.9 kg per 10 km censused), even though
the larger primate species have suffered long-term hunting
pressure. The Beni alluvial plain is also associated with higher
densities, relative to the Brazilian Shield, of many terrestrial
frugivores. This biogeographical variation is thought to be
linked to the relative densities of important fruit resources
such as figs and palms (Painter et al., in prep.), which are
particularly abundant on the Beni alluvial plain (FAN/PL480/
WCS, 1994).

The most immediate threat to the Rios Blanco y Negro Wild-
life Reserve is the logging industry. However, the survey re-
sults reported here are insufficient to draw conclusions about
how much logging disturbance levels are affecting primate
populations in this region. Comparisons between sites are
problematic since pre-logging differences in primate abun-
dance are unknown. In addition, accurate details regarding
the history of logging activities at each site are difficult to
obtain, and any differences could prove important when in-
terpreting results. In any case, although spider monkeys,
which have been identified as being particularly prone to habi-
tat disturbance (McFarland Symington, 1988), were rarely
encountered at many of the sites where logging activities were
occurring (e.g., San Luis, Oquiriquia), this was not always the
case (e.g., Arroyo Chuchui). Nevertheless, high-value timber
populations have been greatly reduced and attention is switch-
ing to a number of potential secondary timber species. This
will not only result in increasing extraction intensities and as-
sociated damage, but in addition many of these secondary
species are important fruit trees for forest frugivores, for ex-
ample, Hymenaea courbaril, Spondias mombin, Ficus sp.,
Ampelocera ruizi (Wallace and Painter, unpubl.). In the long
term, significant reductions in the densities of important
frugivore resources could prove limiting to wildlife popula-

Table 3. Diurnal primate encounter rates (per 10 km) for the Beni Biosphere Reserve, Bosque Chimanes, and Noel Kempff Mercado
National Park.
Beni Biosphere Reserve' Bosque Chimane-' Noel Kem iff Nercado'
Specie, Group ER Individual ER Group ER Indjvidual ER Group ER Individual ER
C t. f'. t,. l 1., -.. _,_,'.'__ ___1 0 _____2
S.;., .r 1 'J, _,__",_,_,).O 21i I i(,_, -_
2 h.,, .i I 2 I-i I 3,- 31A. 1.1 ..<
;,:,. ,..,. t," 0.3 3.4: ''2 ',"I 010.6
FBER 2 S I-6 010).4
PBER = Primate Biomass Encounter Rate.
Source: Adapted from 'Painter etal., 1995, 2Rumiz and Herrera, 1994 and 3Wallace etal, 1998

28 Neotropical Primates 8(1), March 2000

The surveys did uncover considerable evidence of hunting
by logging crews and other employees. Logging-associated
hunting threatens several wildlife species, most notably un-
gulates and cracids (Townsend et al., in prep.). Although pri-
mates are rarely hunted by logging crews in this region (So-
lar, 1996), the most frequently taken species, Ateles chamek,
is particularly sensitive to this activity due to its prolonged
inter-birth interval and a low intrinsic rate of increase
(McFarland Symington, 1988). Spider monkey juveniles also
appear to be popular as camp pets, later to be sold in the
markets of Santa Cruz (Wallace, pers. obs.). This results in a
preference to hunt adult females, which further decreases the
population's ability to recover. In terms of the logging indus-
try, at present extraction intensities hunting probably has the
most negative effect on larger wildlife species in lowland Bo-
livia (Rumiz et al., in press; Townsend et al., in prep.). Every
effort must be made to enforce this aspect of the new law in
the future.


The study was funded as part of a grant to Fundaci6n Amigos
de la Naturaleza (FAN) and Wildlife Conservation Society
(WCS) from the Secretaria Ejecutiva PL-480 Titulo III-
USAID/B. This grant was designated for a biodiversity sur-
vey and management plan of the Rios Blanco y Negro Wild-
life Reserve (RBYNWR). We thank Rudy Guzman and the
late Pavlik Nikitin who coordinated the RBYNWR project
activities and field visits. Our field assistants Jose Chuvifia,
Nicolas Tagua and Manuel Wan were invaluable during surveys.


Anderson, S. 1997. Mammals of Bolivia taxonomy and
distribution. Bull. Amer. Mus. Nat. Hist. 231:1-652.
Anonymous. 1996. Reglamento General de la Ley Forestal. La
Paz, Bolivia.
Branch, L. C. 1983. Seasonal and habitat differences in the
abundances of primates in the Amazon (Tapaj6s) National
Park, Brazil. Primates 24: 424-431.
Braza, E and Garcia, J. E. 1988. Rapport preliminaire sur les
singes de la region montagneuse de Huanchaca, Bolivie. Fo-
lia Primatol. 49: 182-186.
Brockelman, W. Y. and Ali, R. 1987. Methods of surveying
and sampling forest primate populations. In: Primate Conser-
vation in the Tropical Rain Forest, C. W. Marsh and R. A.
Mittermeier (eds.), pp. 23-62. Alan R. Liss, NewYork.
Brown, A. D. and Rumiz, D. I. 1985. Distribuci6n y
conservaci6n de los primates en Bolivia. Estado actual de su
conocimiento. Unpublished report, NewYork Zoological So-
ciety, Bronx, NY.
Buckland, S. T., Anderson, D. R., Burnham, K. P. and Laake,
J. L. 1993. Distance Sampling Estimating Abundance of
Biological Populations. Chapman and Hall, London.
Burnham, K. E, Anderson, D. R. and Laake, J. L. 1980.
Estimation of density from line transect sampling biologi-
cal populations. Wildlife Monographs 72. 202 Pp.
Emmons, L. H. and Feer, E 1990. Neotropical Rainforest

Mammals: A Field Guide, University of Chicago Press, Chicago.
FAN/PL480/WCS. 1994. Plan de Manejo. Reserva de Vida
Silvestre de Rios Blanco y Negro. Santa Cruz, Bolivia.
Ford, S. M., 1994. Evolution of sexual dimorphism in body
weight in platyrrhines. Amer. J. Primatol. 34: 221-244.
Garcia, J. E. and Tarifa, T. 1988. Primate survey of the
Estaci6n Biol6gica Beni, Bolivia. Primate Conserv. (9): 97-100.
Killeen, T. J., Garcia, E. and Beck S. G. (eds.). 1993. Guia de
Arboles de Bolivia. Herbario Nacional de Bolivia & Mis-
souri Botanical Garden.
Mate, C. and Colell, M. 1995. Relative abundances of forest
cercopithecines in Ariha, Bioko Island, Republic of Equa-
torial Guinea. Folia Primatol. 64: 49-54.
McFarland Symington, M., 1988. Environmental determi-
nants of population densities in Ateles. Primate Conserv. (9):
Painter, R L. E., Wallace, R. B. and Pickford, D. 1995. Rela-
tive abundances of peccaries in areas of different human
pressures within the Beni Biosphere Reserve, Bolivia. Ibex,
J. Mountain Ecol. 3: 49-52.
Peres, C. A. 1997. Effects of habitat quality and hunting pres-
sure on arboreal folivore densities in Neotropical forests: A
case study of howler monkeys (Alouatta spp.). Folia Primatol.
68: 199-222.
Rumiz, D. I. and Herrera, J. C. 1994. Mammal and large
bird surveys in Bosque Chimanes. Unpublished Report,
Wildlife Conservation Society, New York.
Rumiz, D. I., Giunart, D, Solar, L. and Herrera, J. C. In press.
Logging and hunting in community forests and logging con-
cessions: Two contrasting case studies in Bolivia. In: Con-
serving Wildlife in Managed Tropical Forests: An Assessment
of Logging Impacts and Options for Conserving Wildlife and
their Habitat in Production Forests, R. A. Fimbel, A. Grajal
and J. G. Robinson (eds.). Columbia University Press, New
Siegel, S. and Castellan, N. J. Jr. 1988. Nonparametric Statis-
tics for the Behavioral Sciences. McGraw-Hill International
Solar, R. L. 1996. Aprovechamiento de la fauna silvestre, y
actividades de busqueda maderera en el Bajo Paragua. Un-
published thesis, Universidad Autonoma Gabriel Rene
Moreno, Santa Cruz, Bolivia.
Terborgh, J., 1983. Five New World Primates: A Study in Com-
parative Ecology. Princeton University Press, Princeton, NJ.
Townsend, W. R., Solar L. and Rumiz, D. I. In prep. The
impact on wildlife populations made by logging personnel
in Bolivian tropical dry forests.
Wallace, R. B. 1998. The behavioral ecology of black spi-
der monkeys in north-eastern Bolivia. Doctoral thesis. Uni-
versity of Liverpool, Liverpool, U.K.
Wallace, R. B., Painter, R. L. E., Taber, A. B. and Ayres, J.
M. 1996. Notes on a distributional river boundary and south-
ern range extension for two species of Amazonian primates.
Neotrop. Primates 4(4):149-151.
Wallace, R. B., Painter, R. L. E. and Taber, A. B. 1998. Pri-
mate diversity, habitat preferences and population density
estimates in Noel Kempff Mercado National Park, Santa
Cruz, Bolivia. Am. J Primatol. 46: 197-211.

NeotropicalPrimates 8(1), March 2000


Patricia G. Guedes
Diva M. Borges-Nojosa
Juliana A. G. da Silva
Leandro 0. Sales


O genero Alouatta Lacepede, 1799 encontra-se distribuido
amplamente pela America do Sul, ocorrendo nas mais variadas
formag6es vegetais desde o sul da Amedrica Central at6 a Ar-
gentina. t o genero mais bem documentado na America do
Sul no que diz respeito a aspects ecol6gicos. A dieta desses
animals e folivora-frugivora e podem formal grupos com 4 a
15 individuos. Por6m, a taxonomia e distribuigco do complex
de especies desse genero continue a ser uma das menos
conhecidas, principalmente para as formas do nordeste
brasileiro. Este fato agrava-se corn a alarmante situacao de
conservagao dos biomas dessa region.

Segundo Gregorin (1996), slo reconhecidas nove especies
brasileiras para o genero entire as quais destaca-se A. b. ululata
Elliot, 1912, que tern distribuico geogrifica aparentemente
disjunta abrangendo desde o norte do estado do Maranhio
ao Ceari. Coimbra-Filho e Camara (1996) e Coimbra-Filho
et al. (1995) remarcam a significativa presenqa de guaribas
no nordeste brasileiro, apesar de serem poucos os dados sobre
a sua distribuicao nesta regiao. Segundo os autores, estes
primatas deveriam ter uma distribuigco muito mais ampla do
que se admite, provavelmente. ocorrendo em todos os estados
da regiao.

Este trabalho tern como objetivo: 1) Registrar duas novas lo-
calidades, no estado do Ceari, complementando a distribuicio
do g&nero Alouatta conhecida para este estado; 2) divulgar o
aumento do nimero de exemplares provenientes do Ceara
disponfveis em coleq6es cientificas; e 3) contribuir de forma
indireta para a elaboraqao de propostas biogeogrificas
associadas ao status taxon6mico dos guaribas dessa regiao.


0 material referente aos guaribas do estado do Ceara e escasso
nas coleq6es mastozool6gicas brasileiras. Existem registros so-
mente para os municipios de Granja (localidade Goiabeira -
MNRJ 23140) e Sao Benedito (localidades Bom Jardim -
MNRJ 23141, MNRJ 23142; e Cinta Sulidon MNRJ
21096). Os exemplares estio depositados na Colegdo de
Mamiferos do Museu Nacional e sao datados dos anos de 1953
(MNRJ 21096) e 1973 (demais).

Neste trabalho sio apresentados registros recentes provenientes
de duas novas localidades desta regiao: Ladeira do Mucambo
(UFC M018) e Cinta da Boa Vista (UFC M019), no Muni-
cfpio de Ibiapina, vizinho ao municfpio de Slo Benedito. Estio
localizados na Chapada da Ibiapaba, uma regiao caracterizada
como um brejo-nordestino com floresta de mata-timida
relictual devido a altitude e posiqao geogrifica pr6xima ao
litoral. A region e considerada como um interessante
remanescente de Mata AdIntica no entanto, vArias areas
vem sendo, ao long destas duas ultimas ddcadas, intensamente
desmatadas. Em um levantamento de mamiferos do estado
do Ceara, Paiva (1973) refere-se aos guaribas como
praticamente extintos e jA naquela 6poca sugeria medidas
urgentes para protecao destes primatas, que podem ser
considerados os mais ameacados dentre os primatas
neotropicais. Porem, esses novos registros confirmam que
ainda existem populac6es isoladas no estado, aparentemente
apenas na regiio da Chapada da Ibiapaba, que foram
comentadas em trabalhos anteriores (Bonvicino et al., 1984;
Coimbra-Filho et al., 1995; Coimbra-Filho e Camara, 1996;
Hirsch et al., 1991; Langguth et al., 1987) e principalmente
pela populacao local durante uma visit i regiao.

Identificaoo do Material

0 material, que encontra-se depositado na Colegio de Ma-
miferos da Universidade Federal do Ceari (UFC), foi doado
por moradores da regiao e consta de dois crAnios, suas
respectivas mandfbulas e um hi6ide, sendo um individuo
adulto (UFC M018) e um jovem (UFC M019). Os
exemplares apresentam os caracteres diagn6sticos e medidas
cranianas semelhantes aos listados por Gregorin (1996) para
a especie A. b. ululata, principalmente no que diz respeito a
morfologia do aparato hi6ide. Este autor apresenta uma
proposta que eleva a subespecie A. belzebul ululata ao status

Y ,



410': 40'30'
Figura 1. Mapa da region da Chapada da Ibiapaba, ilustrando os
novos registros de Alouatta: Ladeira do Mucambo (UFC M018) e
Cinta da Boa Vista (UFC M019), Municipio de Ibiapina (Ceari).

Neotropical Primates 8(1), March 2000

de especie (A. ululata), embora ate o moment seja
correntemente aceita a primeira hip6tese. A proposta teve
como base uma revisao das especies brasileiras do genero,
incluindo vdrios caracteres morfol6gicos, principalmente
relatives a pelagem, que os distinguem da forma A. belzebul
amaz6nica, al6m de outras observag6es mais gerais, expresses
da seguinte maneira:

"Aldm do dicromatismo, da coloraqdo e do tamanho menor
em algumas estruturas cranianas, como evidenciado por
Dollman (1910) e Elliot (1912), A. ululata habitat um tipo
de vegetacio particular. Enquanto que A. belzebul e A.
discolor habitam a floresta equatorial amazonica, A. ululata
habitat uma vegetacao aberta de transicao corn uma alta
freqi!encia de babaqu." (Gregorin, 1996; pp.79-80).

Ainda existed muitas divergencias sobre a validade dos tixons
subespecificos, nio somente para Alouatta como tambem para
outros primatas. Embora nao seja objetivo deste trabalho, cabe
ressaltar que o reduzido nmmero de exemplares reference k A.
ululata pode dificultar qualquer elaboragio de propostas
taxon6micas para esta forma.

Estes registros do Nordeste do Brasil sgo particularmente
interessantes porque constituem mais um exemplo de
distribuigio faunisrica que corrobora a hip6tese da existencia
preterita, pela regiao nordestina, de uma extensa e continue
liga;io entire a Floresta Amaz6nica e a Floresta Atldntica, o
que deve ter permitido que essas populac6es originalmente
amaz6nicas prolongassem sua distribuicao para estas outras
areas e sofressem posteriores especiag6es (Borges, 1991;
Coimbra-Filho e Camara, 1996). Ficaram assim as populac6es
isoladas nos remanescentes, testemunhos desta provivel
conexao paleoambiental.


Esta nova amostragem nio somente dobra o numero de loca-
lidades conhecidas para a especie como tambem dobra o
nimero de exemplares para o Estado. Este incremento de
material disponfvel em coleq6es cientificas poderi contribuir
em futures estudos taxon6micos e/ou biogeogrificos para as
formas de Alouatta. Alim disso, confirmam a sobrevivencia
destes grupos isolados na region quase quinze anos ap6s o
iltimo registro, atraindo a atencqo para a alarmante escassez
de informac6es sobre estes primatas e seu status de


Esse estudo foi apoiado pela CAPES Fundaco Coordenagqo
de Aperfeicoamento de Pessoal de Nivel Superior (PGG) e
PIBIC Programa Institucional de Bolsas de Iniciagao
Cientffica, Universidade Federal de Ceari (JAGS).
Patricia G. Guedes, Laborat6rio de Sistemitica e Evoluglo
de Mamiferos, Museu Nacional, Universidade Federal do Rio
de Janeiro, 20940-040 Rio de Janeiro, Rio de Janeiro, Brasil,
E-mail: , Diva M. Borges-Nojosa,
Caixa Postal 52.856, Universidade Federal do Ceari, 60151-

970 Fortaleza, Ceari, Brasil, Juliana A. G. da Silva, Caixa
Postal 52.856, Universidade Federal do Ceard, 60.151-970
Fortaleza, Ceari, Brasil, e Leandro 0. Salles, Laborat6rio de
Sistemitica e Evoluqdo de Mamiferos, Museu Nacional,
Universidade Federal do Rio de Janeiro, 20.940-040 Rio de
Janeiro, Rio de Janeiro, Brasil.


Bonvicino, C. R., Langguth, A. e Mittermeier, R. A. 1984. A
study of pelage color and geographic distribution in Alouatta
belzebul (Primates:Cebidae). Rev. Nordestina Biol. 6(2): 139-
Borges, D.M. 1991. Herpetofauna do Macico de Baturit6,
estado do Ceari: composigio, ecologia e considerag6es
zoogeograficas. Dissertacgo de Mestrado, Universidade Fed-
eral da Paraiba (UFPB), Joao Pessoa. 91pp.
Coimbra-Filho, A. E e Camara. I. de G. 1996. Os Limites
Originais do Bioma Mata Atltntica na Regido Nordeste do
Brasil. Fundacqo Brasileira para a Conservacqo da Natureza
(FBCN), Rio de Janeiro. 86pp.
Coimbra-Filho, A. E, Camara, I. de G. e Rylands, A. B. 1995.
On the geographic distribution of the red-handed howler
monkey, Alouatta belzebul, in Northeastern Brasil. Neotrop.
Primates 3(4): 176-179.
Gregorin, R. 1996. Variaqao geogrifica e taxonomia das
especies brasileiras do genero Alouatta Lacepede, 1799 (Pri-
mates, Atelidae). Dissertagqo de Mestrado, Instituto de
Biociencias, Universidade de Sao Paulo, Sio Paulo. 181pp.
Hirsch, A., Landau, E. C., Tedeschi, A. C. de M. e Menegheti,
J. 0. 1991. Estudo comparative das espdcies do genero
Alouatta Lacdpede, 1799 (Platyrrhini, Atelidae) e sua
distribuicqo geogrnfica na America do Sul. In: A
Primatologia no Brasil 3, A. B. Rylands e A. T. Bernardes
(eds.), pp.239-262. Fundaqao Biodiversitas, Sociedade
Brasileira de Primatologia, Belo Horizonte.
Langguth, A., Teixeira, D. M., Mittermeier, R. A. e Bonvicino,
C. R. 1987. The red-handed howler monkey in northeast-
ern Brazil. Primate Conserv. (8): 36-39.
Paiva, M. P. 1973. Distribuicao e abundancia de alguns
mamfferos selvagens no Estado do Ceari. Ciancia e Cultura
25(5): 442-450.


Paulo Auricchio
Marco Aurdlio Ferreira da Silva

0 mono-carvoeiro, ou muriqui, tern tido sua presenqa assinala-
da em virias localidades da Mata Atlintica em pequenos grupos
esparsos (Antonietto et al., 1994; Martuscelli et aL, 1994; Oli-
veira et al, 1996; Auricchio, 1997). Em duas ocasi6es em julho
de 1999 pudemos observer um grupo de mono-carvoeiros, a
apenas 300 metros da movimentada primeira pista da rodovia

Neotropical Primates 8(1), March 2000

dos Imigrantes, no Ndcleo Cubato, Parque Estadual da Serra
do Mar. Os avistamentos foram feitos na Subestaco n.53 (esta-
qo de energia) em umra estrada vicinal desta rodovia, chamada
pelos moradores locais como "Estrada de Servico" a 250-300m
de altitude.

0 Parque Estadual da Serra do Mar estende-se do litoral norte
do Estado de Sao Paulo, a partir da divisa corn o Rio de Janeiro,
ate os munidpios de Peruibe e Pedro de Toledo, no sul. Corn
315.390 ha, e o maior parque estadual paulista, e possui a maior
area de florestas do dominio da Mata Adtntica, al6m de various
ecossistemas a ela associados (Raimundo e Lorejan, 1998). 0
Nticleo Cubatio, que abrange as regi6es de planalto, e litoral do
Parque Estadual, abrange uma area de aproximadamente
139.000 ha. E uma regiao montanhosa, corn escarpas de flores-
tas primirias de dificil acesso, que e entrecortada pelas rodovias
que interligam as Regi6es Metropolitanas de Sio Paulo e da
Baixada Santista Caminho do Mar, Anchieta e Imigrantes -
bem como linhas de alta tensIo, oleodutos, gasodutos e ferrovias.

A localidade das avistagens fica aproximadamente a 70 km em
linha reta de MongaguA, ponto mais pr6ximo em que o muriqui
ji foi avistado (Martuscelli, 1994). Para o norte, a localidade de
ocorrencia mais pr6xima desta especie e Poruba, em Ubatuba,
SIo Paulo (Auricchio, 1997) que dista aproximadamente 150
km a nordeste. No dia 16 de julho um grupo de cerca de 10
individuos, foi observado entire 09:00 h e 10:00 h. Uma nova
observagio foi feita no dia 26 de julho. Possivelmente era o mesmo
grupo, pois tambem continha cerca de 10 indivfduos. Nesta
ocasiIo, foi-nos possivel fotografar alguns animals.

Os tr8s funcionirios que trabalham nesta Subestagio disseramr
que v&em os muriquis regularmente nesta 6poca do ano. 0 fun-
cionirio com mais tempo de servico diz que os v& ali desde que
comecou a trabalhar naquele local, ha onze anos. Diz aparece-
rem sempre quando ha sol, em intervalos de dez ou quinze dias
na parte da manha (no horirio entire nove e dez horas) ou a
tarde (mais freqilentemente, segundo o funcionirio da tarde)
por volta das 17:00 h. Os monos deste grupo passam vocalizando
corn gritos e roncos altos, o que torna ficil aos funcionirios da
subestago perceberem sua presenga. Ali, permitem set observa-
dos a 25 m de distincia, alimentando-se e parecendo nio teme-
rem a presence humana.

No grupo havia uma femea que carregava um filhote ainda nas
costas. Um funcionirio diz que as vezes chegam a ficar mais de
uma hora por ali pela manha e afirma tamb6m que ja assistiu os
adults segurarem-se uns nos outros para former uma esp6cie de
"ponte" entire uma irvore e outra mais afastada, para que os
filhotes possam atravessar sobre eles. Um outro morador do lo-
cal, diz que ve freqUientemente os muriquis, nas redondezas do
"Cachoeirio", uma queda d'gua formada pela descida do Rio
Passaritiva, a cerca de 2,8 km da subestacao. Segundo os entre-
vistados, nao hi relatos de cagadores na regiio.

A presenga desta esp&cie na regiio era esperada visto que, como
mencionado anteriormente, e present tambim tanto ao note
quanto ao sul. Como ji indicado em Mendes (1994), Auricchio
(1997) e Olmos etal. (1997), sao necessdrias medidas que possi-

bilitem a protegio efetiva de areas onde haja a ocorrencia desta
espdcie ou a translocaqao destes animals para outras dreas onde ji
existam grupos. Outra media indicada por Auricchio (1997) e
a criacao em instituic6es que fagam seu manejo em cativeiro, aldm
do Centro de Primatologia do Rio de Janeiro, aumentando o
conhecimento biol6gico desta esp&cie, podendo futuramente
tratar de sua reintroduoo. Acreditamos que a ago mais indicada
neste caso seja implementar um program de educacao ambiental
que tenha como foco os trabalhadores da drea, moradores e, num
plano mais amplo, as escolas da regiao.

De qualquer forma, considerando-se que a populacio total desta
esp6cie a 11 anos atrrs foi estimada em 350-400 animals
(Nishimura et al., 1988), a descoberta de um novo grupo de
muriquis em uma regiIo em que eram considerados extintos, e
um alento para aqueles preocupados corn a preservaiao desta

Paulo Auricchio, Instituto Pau Brasil de Hist6ria Natural, Caixa
Postal 282, 07400-970 Aruji, SIo Paulo, Brasil, e-
mail: e Marco Aurelio
Ferreira da Silva, Rua Cons. JoIo Alfredo 342, ap. 73, Estuirio
de Santos, 11015-220 Santos, Sao Paulo, Brasil.


Antonietto, L. A. e Mendes, E D. C. 1994. Sao Francisco Xavier.
A new site for primatological research and conservation in the
Brazilian Atlantic forest. Neotrop. Primates 2(3): 3-4.
Auricchio, R 1997. A new locality for Brachyteles arachnoides
and the urgency of finding new directions for muriqui conser-
vation. Neotrop. Primates 5(3): 78-80.
Fonseca, G. A- B. da, Rylands, A. B., Costa, C. M. R., Machado,
R. B., Leite, Y. L. R. e Furlani, C. (eds). 1994. Livro Vermelho
dos Mamiferos Brasileiros AmeaCados de Extinao. Fundacio
Biodiversitas, Belo Horizonte.
Martuscelli, P, Petroni, L. e Olmos, E 1994. Fourteen new lo-
calities for the muriqui (Brachyteles arachnoides). Neotrop. Pri-
mates 2(2): 12-15.
Mendes, E D. C. 1994. Muriqui conservation: the urgent need
for an integrated management plan. Neotrop. Primates_2(2):
Nishimura, A, Fonseca, G. A. B., Mittermeier, R. A., Young, A.
L., Strier, K. B. e Valle, C. 1988. The muriqui, genus Brachyteles.
In: Ecology and Behavior of Neotropical Primates. Vol. 2., R. A
Mittermeier, A. B. Rylands, A. E Coimbra-Filho e G. A. B. da
Fonseca, pp.577-610. World Wildlife Fund-US, Washington,
Oliveira, M. E e Manzarti, L. 1996. New locality for the muriqui
(Brachyteles arachnoides) in the state of SIo Paulo, Brasil. Neotrop.
Primates 4(3)84-85.
Olmos, E, Franco, A D. C. and Auricchio, P. 1997. A new lo-
cality for Brachyteles arachnoides and the urgency of finding
new directions for muriqui conservation. Neotrop. Primates 5(2):
Raimundo, S. e Lorejan, S. F. 1998. Planos de Manejo das
Unidades de Conservafdo Parque Estadual da Serra do Mar -
Nicleo Cubatao. Plano de Gestho Ambiental Fase 1. Secretaria
do Meio Ambiente, Sio Paulo. 177pp.

32 Neotropical Primates 8(1), March 2000


Robin C Brockett
Robert Horwich
Clara B. Jones

The purpose of this note is to document female dispersal in
the Belizean black howling monkey (Alouatta pigra). Female
dispersal has been reported in three species of Alouatta (the
mantled howling monkey [A. palliata] by Jones, 1978, 1980a,
1980b; Glander, 1992; the red howling monkey [A. seniculus]
by Sekulik, 1982; Agoramoorthy and Rudran, 1995; Pope,
1992; and the black-and-gold howling monkey [A. caraya]
by Calegaro-Marques and Bicca-Marques, 1996; Giudice,
1997; Agoramoorthy and Lohmann, 1999), and is thought
to be related to food of poor quality (e.g., a folivorous diet,
see Wrangham, 1980; Jones, 1999) and low within-group
competition for food (Wrangham, 1980; Sterck et al., 1997).
Although the causes of female dispersal in primates and other
mammals are still debated, it is assumed that females leave
their resident groups when the costs of remaining outweigh
the costs of dispersal (Jones, 1999).

Our research is conducted at the Community Baboon
Sanctuary (CBS) (1733' N, 8835' W), Belize District, Belize,
C.A. Details of the study site and howler population can be
found elsewhere (Horwich and Lyon, 1990; Silver et al.,
1998; Ostro etal., 1999). Systematic studies of these "arboreal
folivores" have taken place since 1985 in secondary moist tro-
pical forest, including riparian habitat along the Belize River
(Horwich and Lyon, 1990). Black howlers may be locally
endangered or extinct in Belize (Horwich et al., 1993), and
studies of their behavior, social organization, ecology, and
genetics are in their early stages. Preliminary studies describe
a primarily polygynous breeding structure (Horwich et al.,
in prep.) with relaxed or avoidant social relations among
females (Brockett et al., in press). Our research suggests that
habitat is saturated for black howlers at the CBS (Horwich et
al., in prep.) and that frequent male takeovers have significant
consequences for the behavior and fitness of females (Brockett
et al., 1999; Brockett et al., 2000 ; Horwich et al., in press).

Our ad libitum. behavioral observations of black howlers at
the CBS lead us to the conclusion that juvenile and adult
females disperse from their resident groups. Female behavior
is seminal to an understanding of primate social organization
because the decisions that females make in relation to limiting
resources are thought to determine a population's structure
and productivity under density-dependent conditions since,
all other things being equal, female dispersion in time and
space will map onto resources while male dispersion will map
onto females (Emlen and Oring, 1977; Wrangham, 1980).
Monthly surveys of 19 groups (N approximately 100
individuals) at the CBS between 1995 and 1997 show that
two adult females and four juvenile females have emigrated
from groups while no adult females and one juvenile female

have immigrated into groups. Thus, female group
membership appears relatively stable because immigration is
rare, similar to reports for polygynous A. seniculus (Sekulik,
1982; Agoramoorthy, 1994; Agoramoorthy and Rudran,
1995). Secondary dispersal (transfer of adults from one group
to another) also appears to be rare, similar to reports for
polygynandrous A. palliata, (Glander, 1992) and polygynous
A. caraya (Agoramoorthy and Lohmann, 1999) and A.
seniculus (Sekulik, 1982; Agoramoorthy and Rudran, 1995;
Pope, 1992).

Our observations suggest that female-female aggression,
including "targeting" behavior (see Sterck et al., 1997) may
be the proximate cause of patterns of female dispersal. For
example, females at the CBS have been observed to
aggressively expel other females from groups, although
coalitions between two females to expel a third, reported for
polygynandrous A. palliata (see Jones, 1980a) have not been
observed in black howlers. Further, female-female aggression
appears to increase with male takeovers. After a male takeover,
one of us (RCB) observed an adult female immigrant to the
Baizar/Joseph group aggressively expel a resident adult female
and a resident juvenile female, both of whom subsequently
emigrated. The juvenile female suffered severe injuries and
was never seen again. These and other observations suggest
that competition for group membership is intense in black
howlers, as suggested for A. palliata (see Jones, 1980a) and
that female-female aggression, possibly for limiting resources,
preceded female dispersal in howlers.

What is the fate of emigrating female A. pigra? Our
observations suggest that colonization is a major reproductive
strategy for emigrating females, as reported for A. seniculus
(e.g., Pope, 1992; Crockett, 1996). In one instance, for
example, an adult male was observed to establish a new group
>1 km from his group of origin with solitary females of
unknown origin (see Horwich et al., in press, Fig. 1). New
groups have been observed to form in uninhabited patches
of forest as well as in areas of home range overlap between
existing groups. Since female dispersal is a necessary condition
for colonization (Horn and Rubenstein, 1984, p.289), female
dispersal can be assumed to be ancestral to colonization as a
reproductive strategy. Alternatively, female dispersal and
colonization may both be a function of a third factor (e.g.,
female-female aggression over limiting resources or
infanticide). Tracing these evolutionary routes has significant
implications for Alouatta sociobiology since (polygynandrous)
A. palliata females at Hacienda la Pacifica are more likely to
immigrate into existing groups than to colonize open habitat
(Glander, 1992).

Although present evidence indicates that black howlers, like
other species of Alouatta, should be classified as "dispersal-
egalitarian" according to the system employed by Sterck et al.
(1997), the behaviors reported in- this note and additional
observations suggest that black howlers may violate certain
assumptions of Sterck et al.'s ecological model. For example,
our observations suggest that "targeting" behavior by females
represents intense within-group competition that may be

Neotropical Primates 8(1), March 2000

increased by male takeovers. Additionally, we have observed
females of different groups behave aggressively towards one
another, suggesting that between-group competition may
influence patterns of social behavior among female A. pigra.
Wrangham (1980) argued that female philopatry was favored
by between-group competition, a conclusion that may require
modification as a result of research on Alouatta. Related to
this, Koenig et al. (1999), studying Semnopithecus entellus,
demonstrated that competition for food may be intense among
folivores (see Fedigan et al., 1998; Jones, 1980a). Although
Hanuman langur females generally exhibit philopatry, the
findings of Koenig et al. (1999) may indicate that female
dispersal is not strictly a function of food type or degree and
type of competition per se but of food dispersion and,
especially, quality (Jones, 1999). Supporting this
interpretation is Koenig et al.'s report that Hanuman langurs
eat a broader range of food types than howlers, in particular,
bark and non-herbivorous foods (e.g., insects). Thus, female
dispersal may in part be a function of some threshold of food
quality relative to the biology of a species, differential
competitive regimes, and other factors (e.g., food abundance
or avoidance of inbreeding). Female dispersal is a primitive
trait in Alouatta, and its further investigation will provide
insights into the costs and benefits of "non-female bonded"
groups, in particular, and female social relations in general.

Acknowledgments: We are grateful to Dr. K. E. Glander for
his assistance in marking the animals. These studies were
carried out with the support of the National Geographic
Society (Grants #5352-94 and #5653-96).

Robin C. Brockett, Zoo Atlanta, Atlanta, Georgia, 30315
USA and Community Conservation, Inc., Robert Horwich,
Community Conservation, Inc., R.D. 1, Box 96, Gays Mills,
WI 54631, USA and Clara B. Jones, Livingstone College,
Salisbury, NC 28144, USA, and Community Conservation,
Inc. Address for correspondence: Clara B. Jones, Livingstone
College, Division of Social Sciences, 701 W. Monroe Street,
Salisbury, North Carolina 28144, U.S.A, e-mail: < cjones
@livingstone .edu>.


Agoramoorthy, G. 1994. An update on the long-term field
research on red howler monkeys, Alouatta seniculus, at Hato
Masaguaral, Venezuela. Neotrop. Primates 2: 7-9.
Agoramoorthy, G. and Lohmann, R. 1999. Population and
conservation status of the black-and-gold howler monkey,
Alouatta caraya, along the Rio Riachuelo, Argentina.
Neotrop. Primates 7:43-44.
Agoramoorthy, G. and Rudran, R. 1995. Infanticide by adult
and subadult males in free-ranging red howler monkeys of
Venezuela. Ethology 99: 75-88.
Brockett, R_ C., Horwich, R. H. and Jones, C. B. 1999. Dis-
appearance of infants following male takeovers in the
Belizean black howler monkey (Alouatta pigra). Neotrop.
Primates 7: 86-88.

Brockett, R. C., Horwich, R. H. and Jones, C. B. 2000. A
model for the interpretation of grooming patterns applied
to the Belizean black howling monkey (Alouatta pigra). Pri-
mate Report 56:23-32.
Calegaro-Marques, C. and Bicca-Marques, J. C. 1996. Emi-
gration in a black howling monkey group. Int. J. Primatol.
17: 229-235.
Crockett, C. M. 1996. The relation between red howler mon-
key (Alouatta seniculus) troop size and population growth
in two habitats. In: Adaptive Radiations of Neotropical Pri-
mates, M. A. Norconk, A. L. Rosenberger, and P. A. Garber
(eds.), pp.489-510. Plenum Press, New York.
Emlen, S. T. and Oring, L. W. 1977. Ecology, sexual selec-
tion and the evolution of mating systems. Science 197: 215-
Fedigan, L. M., Rose, L. M. and Avila, R. M. 1998. Growth
of mantled howler groups in a regenerating Costa Rican
dry forest. Int. J. Primatol. 19: 405-432.
Giudice, A. M. 1997. Comportamiento social en aulladores:
El caso de la emigraci6n de una hembra sabadulta en
Alouatta caraya. Neotrop. Primates 5: 39-43.
Glander, K. E. 1992. Dispersal patterns in Costa Rican
mantled howling monkeys. Int. J. Primatol 13: 415-436.
Horn, H.S. and Rubenstein, D.L. 1984. Behavioural adap-
tations and life history. In: Behavioural Ecology: An Evolu-
tionary Approach, J. R. Krebs and N. B. Davies (eds.),
pp.279-300. Sinauer Associates, Sunderland, MA.
Horwich, R. H. and Lyon, J. 1990. A Belizean Rainforest.
Orang-Utan Press, Gays Mills, WI.
Horwich, R. H., Brockett, R. C. and Jones, C. B. In press.
Alternative male reproductive behaviors in the Belizean
black howler monkey (Alouatta pigra). Neotrop. Primates 8.
Horwich, R. H., Koontz, F., Saqui, E., Saqui, H. and Glander,
K. 1993. A reintroduction program for the conservation of
the black howler monkey in Belize. Endangered Species UP-
DATE 10: 1-6.
Jones, C. B. 1978. Aspects of reproduction in the mantled
howler monkey (Alouattapalliata Gray). Unpublished Ph.D.
Dissertation, Cornell University, Ithaca, NY
Jones, C. B. 1980a. The functions of status in the mantled
howler monkey, Alouatta palliata Gray: Intraspecific com-
petition for group membership in a folivorous Neotropical
primate. Primates 21: 389-405.
Jones, C. B. 1980b. Seasonal parturition, mortality and dis-
persal in the mantled howler monkey, Alouattapalliata Gray.
Brenesia 1: 1-10.
Jones, C. B. 1999. Why both sexes leave: Effects of habitat
fragmentation on dispersal behavior. Endangered Species UP-
DATE 16: 70-73.
Koenig, A., Beise, J., and Chalise, M. K. 1999. Gentle leaf-
eaters? Food, competition and female social relationships of
Hanuman langur monkeys (Presbytis entellus) from south-
ern Nepal. J. Nepal Res. Centre 11: 37-50.
Ostro, L. E. T, Silver, S. C., Koontz, E W., Young, T. P., and
Howich, R. H. 1999. Ranging behavior of translocated and
established groups of black howler monkeys Alouatta pigra
in Belize, Central America. Biol. Conserve. 87: 181-190.

34 Neotropical Primates 8(1), March 2000

Pope, T. R. 1992. The influence of dispersal patterns and
mating system on genetic differentiation within and between
populations of the red howler monkey (Alouatta seniculus).
Evolution 46: 1112-1128.
Sekulik, R. 1982. Behavior and ranging patterns of a solitary
female red howler (Alouatta seniculus). Folia Primatol. 38:
Sterck, E. H. M., Watts, D. P., Van Schaik, C. P. 1997. The
evolution of female social relationships in nonhuman pri-
mates. Behav. Ecol Sociobiol. 41: 291-309
Silver, S. C., Ostro, L. E. T., Yeager, C. P., Koontz, EW. and
Horwich, R. 1998. The feeding ecology of the black howler
monkey (Alouattapigra) in northern Belize. Am. J. Primatol.
45: 263-279.
Wrangham, R. W. 1980. An ecological model of female-
bonded primate groups. Behaviour 75: 262-299.


Romari A. Martinez
Rosa A. Moscarella
Marisol Aguilera
Eladio Mdrquez

Among Venezuela's endemic mammal species, the capuchin
monkey from Margarita Island, Cebus apella margaritae
Hollister, 1914, deserves special attention. It is the only
primate inhabiting the island, which is off the north coast of
Venezuela. These capuchins are isolated, with the nearest
Cebus apella populations occurring in the Amazon region,
approximately 800 km to the south. This gap is enigmatic
and remains unexplained (Bodini and P&rez-Hernindez,
1988). The most recent information on wild populations of
C. apella margaritae comes from Mirquez and Sanz (1992)
and Sanz and Marquez (1994). According to these authors
C. a. margaritae populations on Margarita Island are
threatened due to habitat degradation, illegal hunting and
commerce. Farmers consider the monkeys crop pests and have
sold individuals as pets for prices up to US$41 (Ottocento et
al., 1989). Over the past 15 years, an increasing human
population has pushed farmers to the borders of the island's
reserves. Settlers have already invaded the upper regions of
the Cerro El Copey National Park (7,130 ha, created in
1974), at altitudes up to 500 m. The more humid soils found
in the mountain ranges of Copey, Tragaplata and Matasiete
(a Natural Monument of 1,672 ha, created in 1974) have
also encouraged timber cutting for small slash-and-burn
agricultural plots. These combined factors are creating
significant concern for the capuchins of Margarita Island.

Currently, we are carrying out a study of the genetics and
conservation status of the Margarita Island capuchin. The
aims of our project are to obtain peripheral blood of pet ca-
puchins in order to investigate the origin of this disjunct is-
land population and to carry out a survey to better under-

stand how illegal hunting and commerce effect the status of
the species. In February 1997, we visited homes and small
businesses in the eastern part of the island. The selection of
places to visit was based on information obtained from set-
tlers (Fig. 1), as well as an interview with the local office of the
Ministry of Environmental Resources in Margarita. We in-
terviewed people who admitted having or to have had a pet
monkey. When we found a pet we recorded its age, sex,
precedence and habits and carried out a physical examina-
tion. We also asked for information on prices paid for mon-
keys, resellers, veterinary care, and general knowledge of Ven-
ezuelan laws on the possession of wild animals as pets.

Fourteen pet capuchins were found. To our surprise, only
five were Cebus apella margarita: two juvenile males, one
adult male, one juvenile female, and one adult female
(reported by its owner as a male). The others were weeper
capuchins Cebus olivaceus, and two were reported to have been
captured from Cerro Matasiete. The diet for most of the
monkeys consisted of human foods, especially bread and milk
(>80% of the diet), and fruits. Only one owner admitted to
seeking local veterinary care. Prices for monkeys were as high
as US$270. Some of the people claimed to have shot monkeys
either invading their crops or in the vicinity of their fields.
Most of those interviewed (especially near to the Cerro El
Copey National Park) admitted knowing that Venezuelan laws
deam it illegal to own a wild animal as a pet, and that punitive
measures could be taken against those found selling or taking
monkeys from the park.

We found a eukaryote parasite in the total of four blood
samples of C. a. margarita. This microorganism is similar to
Trypanosoma sp., but a definitive identification has yet to be
made. None of the interviewees knew of the taxonomic simi-



Margarita Island
--. BRAZIL .....

Figure 1. Margarita Island, Venezuela. Locations where Cebus
monkeys were being kept as pets. Map by Stephen D. Nash.

Neotropical Primates 8(1), March 2000 35

larity of the margaritae monkey to C. olivaceus and the pres-
ence of the unknown blood parasite in all the samples re-
quires further studies to evaluate the risks of zoonosis, not
only for pet owners, but for the entire human population
inhabiting the island.

The occurrence of C. olivaceus as a pet is evidence of
considerable illegal traffic of capuchin monkeys into Margarita
Island. This was not mentioned as a threat to C. a. margaritae
during our interview with the local officer from the Ministry
of Environmental Resources, but if feral populations of C
olivaceus exist on the island (mentioned by two interviewees),
this may be an additional threat to the endemic population
of monkeys. The possible occurrence of free ranging C.
olivaceus may be a recent phenomenon. Several capuchin
owners admitted to having released pets into the mountains
without knowing the species. Although Cebus species are
sympatric on the continent (Terborgh, 1981), the amount of
deforestation and loss of natural habitat due to farming and
human settlement could result in competition between the
species on the island.

Surveys of the natural populations of Ca. margaritae, and
for the possible presence of C olivaceus, are urgently needed,
as is an environmental education program to make people
aware of the significance of the island's monkey species. The
success of similar projects with other endangered species on
the island, such as the Margarita parrot (yellow-shouldered
parrot, Amazona barbadensis), has shown that the people are
sensitive towards wildlife in their area and are willing to modify
their habits to favor the conservation of species and habitats.

We still do not know whether the Margarita capuchins were
isolated prior to the arrival of human settlers or whether they
were introduced. The two populations on the island are
disjunct; occurring between two widely separated mountain
ranges. The increase in human settlements, especially near to
the park, small-scale farming and burgeoning tourism and
recreation, coupled with the generally poor economy of the
island, have resulted in widespread habitat fragmentation, the
consequences of which have yet to be assessed (Ottocento et
al., 1989; Marquez and Sanz, 1992). There is an urgent need
for effective protection and management of the Cerro El
Copey and Matasiete reserves. Genetic studies will hopefully
give us an idea as to the origin of these monkeys and their
unexplained isolation from their closest subspecies, and will
allow for the design of a management plan. It is possible that
C a. margaritae is suffering from some degree of inbreeding
depression, or even that it may be recovering from a
population bottleneck (founder effect). The assessment of the
genetic status of this endemic and little known monkey is
extremely important to answer such questions and for an
effective management plan to be put into place.

Romari A. Martinez, GIBE, Lab. 96, 4 Piso, FCEN-UBA,
Pabell6n II, Ciudad Universitaria (1428), Capital Federal,
Argentina, e-mail: , Rosa
Moscarella, Marisol Aguilera and Eladio Marquez,

BIOEVO, Departamento de Estudos Ambientales, Pab. II,
Universidad Sim6n Bolivar, Caracas 1080-A, Venezuela.


Bodini, R. and Pdrez-Hernindez, R. 1988. Distribution of
species and subspecies of cebids in Venezuela. Fieldiana
Zoology 39: 231-244.
Bisbal, E 1983. Inventario preliminary de fauna de la Isla de
Margarita. Serie Informe Tecnico DGSIIA/IT/138,
Ministerio del Ambiente, Caracas, Venezuela. 24pp.
Cabrera, A. 1958. Catalogo de mamfferos de America del
Sur". Rev. Mus. Argentino Cienc. Nat. "Bernardino
Rivadavia" 4: 1-307.
Hill, W. C. 0. 1960. Primates. Comparative Anatomy and
Taxonomy, IV Cebidae, Part A. Edinburgh University Press,
Marquez, L. and Sanz, V. 1992 Evaluaci6n de la presencia de
Cebus apella margaritae en la Isla de Margarita, Venezuela.
T.E.G., U.C.V., Fac. de Cs., Caracas, Venezuela. 68pp.
Ottocento, R., Marquez, L., Bodini, R. and Cordero, G.
1989. On the presence of Cebus apella margaritae on
Margarita Island, Venezuela. Primate Conservation (10):19-21.
Rodriguez, J. P. and Rojas-Suarez, E 1995. El Libro Rojo de
la Fauna Venezolana. Provita Fundaci6n Polar, Caracas.
Sanz, V. and Marquez, L. 1994. Conservaci6n del mono
capuchino de Margarita (Cebus apella margaritae) en la Isla
de Margarita, Venezuela. Neotrop. Primates 2(2): 5-8.
Terborgh, J. 1981. Five New World Primates. Princeton Uni-
versity Press, Princeton, NJ.


Adrian A. Barnett
Becca Shapley
Shawn Lehman, Mireya Mayor
Everton Henry, Paul Benjamin
Michael McGarrill and Ruford Nagala


Guyana is one of the few Neotropical countries where
substantial tracts of forest remain (Groombridge, 1992,
Bowles et al., 1998). Even with recent work (Phillips-Conroy
and Sussman, 1995; Sussman and Phillips-Conroy, 1995;
Lehman, 1999; Lehman et al., 1995), much of the country
remains unsurveyed for primates.

Here we report on the primate fauna of the Potaro Plateau,
Guyana, a 1,165 km2 outlier of the Pakaraima Mountains,
and the eastern-most extension of the Guayana Highlands
(see Fig.1), with a basal altitude of some 500 m and higher
peaks (2042 m, Mt. Ayanganna; 1594 m, Mt. Kopinang;
1470 m, Mt. Wokamung and others). Like much of the

36 Neotropical Primates 8(1), March 2000

Guyana Highlands, the Plateau consists of pink Proterozoic
Roraima Formation sandstone (1600-1800 m.y.b.p.), capped
with younger volcanic dolerites (Haffer, 1974). Soils are old
and nutrient-poor (Maguire, 1970; Haffer, 1974). A portion
of the Plateau constitutes the Kaieteur National Park (this is
currently being expanded from 11.65 km2 to 580.16 km2;
see The World Bank, 1998) (see Fig. 2).

The vegetation of the National Park has been described by
Henkel (1994) and by Kelloff and Funk (1998). The
vegetation of some other parts of the Plateau has been
described by Maguire et al. (1948). The floristic communities
vary from white sand scrub to flooded riparian forest,
basimontane forest, montane forest, upper montane forest,
high tepui forest and high tepui scrub (Huber et al, 1995).
Though the region has received attention from botanists,
zoological data is sparse for the Plateau.

This report is based on two surveys. One was conducted by
SL and MM in March 1995 (dry season). The other,
conducted by AB, BS, EH and PB, was carried out between
June and August 1998 (wet season) and formed part of the
work of the Potaro Plateau Expedition (PPE). Data were
collected during field observations and through interviews
with local residents.

Upon contact with primate groups, habitat type was recorded
and location was determined with hand-held GPS units
(Garmin 12XL, AB & BS; Magellan NAV 5000D, SL). Ad
libitum notes on behavior, vocalizations and obvious individual
physical characteristics were also collected. Altitudes were
taken from 1:50,000 topographic maps (Survey Department
of Guyana 1975: Kurukabaru Sheet 49 NE, Kurukabaru
Sheet 49 NW and Ayanganna Sheet 42 SE).


Survey methods differed. SL and MM conducted repeat
surveys (N = 22) of a 5.5 km transect, plus 8 km of once-only
surveys of an established trail, along the western edge of the
post-falls river canyon and from the edge of the Kaieteur Falls
through the National Park and into the surrounding habitat.
The PPE recorded primates as they were encountered during
fieldwork surveys for other mammal groups (bats, otters, small
mammals and birds). Survey sites were Muri-muri and
Amamuri Creeks (mouth located at 05010 N, 59030 W and
05007 N, 59032 W, respectively), Chenapou village (04059
N, 59035 W), Tiger Bay village (04059N, 59034W), Tappa
airstrip (0454 N, 59045 W), Conrad's mining camp
(0459N, 59039W), Menzie's Landing (04.51'.N, 59.41'W)
and Mount Kowa (04051 N, 59042 W). Locations of all survey
sites are given in Figure 2.

For the Potaro Plateau Expedition, AB interviewed the
inhabitants of the villages of Tiger Bay and Chenapou and
the park ranger (Mike Phang), using the methods of Barnett

Figure 1. Location of the Potaro Plateau and other places
mentioned in the text.

.---Mt. Ayanganna PB

MnzivsLanding ^Potato River

1o Miii-Mm Creek -
Kaieteur Falls N
CoZrnd's Anamui Creek T
P ienapou
STiger Bay

A pp 4 Chenapou River -

('> Mt. Kowa

Figure 2. Location of study sites on the Potato Plateau.

(1996) and Cunha and Barnett (1989). Using the methods
of Lehman (1994) and Sussman and Phillips-Conroy (1995),
SL and MM conducted interviews near Kaieteur Falls with
the previous park ranger (Lawrence Gibbons), three resident
miners and three Amerindians from Paramakatoi village who
were traveling through the National Park., Great care was
taken to avoid leading questions. Interviews consisted of: (i)
having people identify local monkey species from laminated
colour photocopies of selected illustrations in Eisenberg
(1989) and Reid (1998) (PPE), illustrations in Emmons

Neotropical Primates 8(1), March 2000

(1989) and laminated colour print photographs (SL and
MM); (ii) asking about primates of the immediate area versus
those that occur anywhere in Guyana; (iii) asking for physical
descriptions of each species; (iv) asking about the relative
abundance of primates in the area (i.e., not found, rare,
sometimes seen, abundant, very abundant); and (v) requesting
information on hunting techniques (if any).

In addition to interviews, the PPE looked for examples of
hunting and of pet keeping during visits at the following
locations: Chenapou, Tiger Bay, Tappa Airstrip, Conrad's
mining camp and Menzie's Landing.


Primate locations are given in Figure 2. SL and MM censused
a total of four groups of primates during repeat surveys in the
park; three groups of wedge-capped capuchins (Cebus o.
olivaceus) (N = 15) and one male-female pair of red howlers
(Alouatta seniculus). All encounters were outside the
boundaries of the park, with SL observing no primates during
his single surveys at Kaieteur Falls National Park. The total
sighting rate for primate groups was low, only 0.38 groups/
10 km.

The PPE achieved nine primate encounters of five species. C.
o. olivaceus were observed four times on Mt. Kowa. Group
size ranged from four to eight animals. Over a 10-day period,
groups consisting of four spider monkeys (Ateles paniscus) were
seen three times on Mt. Kowa at distances of between 20 and
60m, and assumed to be repeat sightings of the same group.
A red howler was seen in waterside vegetation at Muri-muri
Creek. Pre-dawn vocalizations from several troops of A.
seniculus were heard every morning at Chenapou and again
in the forests surrounding the airstrip at Tappa. None were
heard on Mt. Kowa, though the species is known to inhabit
montane forest areas (Husson, 1957; Mittermeier, 1977;
Ochoa et al., 1993). Since hunting pressure on Mt. Kowa is
low to non-existent, the reason for the unexpected absence of
A. seniculus there is obscure.

On 1 August 1998, a group of four Cebus albifrons was seen
byAB, BS, EH and PB at a distance of 30 m, some 7 m up in
flooded forest bordering Amamuri Creek, a left bank
tributary of the Potaro River. The light was good and the
animals were clearly visible with both binoculars and the naked
eye. The animals displayed all the field characters of the species
(see Hershkovitz, 1949; Eisenberg, 1989; Kinzey, 1997): body
colour was much browner than the C. o. olivaceus seen on
Mt. Kowa; there was no black line down the middle of the
forehead; the facial fur was white; the forearms were not gray
and the shoulders were white. Two of the four observers also
reported that the animals had whitetipped tails. This is the
first record of C. albifrons for Guyana and appears to be the
first time this species has been recorded east of the Rio Branco,
Brazil. A single Pithecia p. pithecia was observed with the C.

A combined total of 18 interviews were conducted, with
information from 34 people (12 interviews with 27 people
by AB, SL conducted 6 interviews with 7 people). The
residents of Chenapou confirmed and identified all the species
encountered by the PPE. The interviewees specifically denied
that Saguinus midas, Aotus trivirgatus, Cebus apella, Chiropotes
satanas and Saimiri sciureus occurred on the Plateau. Most
informants knew of Saimiri, but considered it to be a coastal
species. Aotus was said to occur in the north of the country.
Local reports of a 'night monkey' were found to refer to the
kinkajou (Potus flavus) (see Roussillhon, 1988, for a similar
occurrence in French Guiana). In the opinion of local people,
overall rank order of abundance for the region's primates was:
Alouatta, Cebus spp., Ateles, Pithecia. Ateles was considered to
be locally abundant in remoter areas.

SL and MM's informants reported that Saimiri, Cebus apella,
C olivaceus and Alouatta were to be found in or near the
park. Ateles was considered very rare in the Kaieteur region,
the result of intensive hunting by miners. According to two
inhabitants of Paramakatoi village, Ateles did still occur 2-3
days walk SE of the falls (i.e. the Mt. Kowa region). The
existence of Chiropotes was denied by all informants. There
was disagreement on the presence of Pithecia in the Kaieteur
region, some interviewees affirmed its presence and some
denied it. No data was obtained on the presence of Aotus.
That these interview results are not entirely consistent with
those of the PPE may be attributable to the PPE's near-
exclusive interviewing of Indigenous Patamona people for
information. Only three of SL and MM's informants were
long-term residents in the Plateau, though not of the region
in which the interviews were being conducted. In addition,
the "area around the park" included the lowland forests at
the base of the Falls and beyond, and thus is not neccasarily
relevant to the Plateau, of which the National Park is a small

In interviews for the PPE, AB found that no-one reported
hunting monkeys regularly. Paca (Agouti paca), peccaries
(Tayassu pecari and T tajacu) and tapir (Tapirus terrestris)
reportedly formed the hunters' mainstay. Monkeys were
hunted only rarely. The most common reasons given were i)
that they looked too human, either when alive (seven
interviewees), or when skinned (five interviewees) or ii) that
the difficulty of hunting them outweighed the cartridge costs
compared to the ease of hunting the abundant paca (four

No primates were observed being kept as pets in either
Chenapou, Tiger Bay or Menzie's Landing, though keeping
wild animals as pets appeared common in these villages. Unlike
the situation in eastern Bolivar (Kinzey et al., 1988), C.
olivaceus was not reported to attack crops and, as a result, was
not punitively hunted.

Neotropical Primates 8(1), March 2000

Table 1. Primate species recorded in areas close or adjacent to the
Potaro Plateau.
Species Mt. Canaima S E Boliiar
Roraima National Park i Norconk
ITate 1932, Ochoa etal/., etal..
19391 19931 19961

S ,. i . ,
(. 'th. <.'/f*\r.l* .
I'.Cl,,'" a tJp ,'',, l._

.lel. l l,- ,, l /.,,_ !

1 not seen, but considered very probable.


The abundance ranking from local interviews broadly agrees
with the data of Norconk et al. (1996) who also reported that
P. pithecia is everywhere rare in Guyana, and with the data in
Kinzey et al. (1988) from neighboring eastern Bolfvar state,

The primate records are of great interest as the distribution
of several species still requires clarification (Sussman and
Phillips-Conroy 1995; Phillips-Conroy and Sussman 1995).
These authors reported the distribution of Saimiri to be
patchy. The reported absence on the Plateau of Chiropotes
satanas and Saguinas midas is in agreement with Sussman and
Phillips-Conroy's belief that neither of these species occur west
of the Essequibo River nor north or west of the Rupununi
River (Sussman and Phillips-Conroy 1995).

Ateles paniscus is very sensitive to hunting (Robinson and
Ramirez, 1986), and both its presence and the group
members' reactions to human observers supported local
opinion that Mt. Kowa is very rarely visited by hunters.
Muckenhirn et al. (1975) considered that Ateles did not occur
north of the Potaro River. Our records at Mt. Kowa neither
confirm nor deny this, since Kowa lies on the southern bank
of that river. It does, however, show that the species is still
present in this remote region.

The north-west extent of the distribution of C. apella is
uncertain according to Sussman and Phillips-Conroy (1995)
and Phillips-Conroy and Sussman (1995). Its apparent
absence from the Potaro Plateau is in line with the belief of
Lehman (1999) that the species does not occur further north
than the riparian forests along the Rupununi River; and with
that of Muckenhirn et al. (1975) who believed the species to
be absent from the north-west of the country.
The taxonomy of the genus Cebus, particularly of C albifrons,
is in need of revision (Rylands et al., 1995). As noted by
Phillips-Conroy and Sussman (1995), further confusion to
the situation is brought about by the presence of hybrids (e.g.
Moonen 1987). However, the possibility that the animals seen
at Amamuri Creek represent a visually distinct subspecies of
C. olivaceus is unlikely given the possibilities (see Hershkovitz,

The record of C albifrons is significant as it appears to be the
first time it has been recorded in Guyana (see Sussman and
Phillips-Conroy, 1995; Norconk etal., 1996; Lehman, 1999),
and the first time it has been recorded east of the Rio Branco
in Brazil (see maps in Emmons, 1997; Kinzey, 1997). This is
a range extension of some 200 km. The species was not
recorded in adjacent parts of Venezuela by Bodini and Perez-
Hernindez (1987); their apparent record for Mount Roraima
on Map 6 (p.241) was refuted by their statements in the text
(see also Norconk et al., 1996). Primate species of adjacent
parts of Venezuela have been reported by Tate (1932, 1939),
Norconk et aL (1996) and Ochoa et aL (1993) (see Table 1)
and do not indicate the presence of C albifrons. However, it
appears that no primate surveys have been conducted in the
critical region between the eastern bank of the Rio Branco
and the Guyanese border with Brazil (John E Eisenberg, pers.
comm.; A. L. Gardner, pers. comm.; Anthony Rylands, pers.
comm.; Richard Thorington, pers. comm.; and see
Hershkovitz, 1977, p.901). It is therefore uncertain if this
record represents the documentation of a true long-
established historical range of the species, or a very recent
range extension (possibly as a result of the very severe forest
fires in the region in early 1998, see Hammond and ter Steege,
1998; Schulze, 1998). Further fieldwork is needed to resolve
this, but it should be noted that the presence of the isolated
C. albifrons subspecies on Trinidad (C. a. trinitatis) argues
for the species historically having a much wider range than
currently thought.

Importantly, the field characteristics of the animals seen at
Amamuri Creek are not consistent with those described for
any of the visually distinct subspecies of C olivaceus (see
Hershkovitz, 1949).


We thank the staff of Guyana's Environmental Protection
Agency the EPA committee and the Minister of Interior Affairs
for their help and kindness in granting permits for our study.
We would also like to thank. the people of Chenapou village,
Mike Tammassar (University of Guyana), Mike Phang
(Guyanese National Parks Service), A. L. Gardner
(Smithsonian Institution), Carol Kelloff (Smithsonian
Institution), Ann McLarnon (Roehampton Institute London),
Caroline Ross (Roehampton Institute London), Anthony
Rylands (Center for Applied Biodiversity Science,
Conservation International, Washington, DC), Richard
Thorington (Smithsonian Institution), the Librarians at the
Zoological Society of London (especially Ann Sylph) and the
staff of the Library at the Academy of Natural Sciences,
Philadelphia. SL and MM are grateful to the University of
Guyana, Office of the President, Ministry of Amerindian
Affairs, and the Smithsonian Institution for permission to
conduct their research. SL and MM's work was supported in
part by the Lincoln Park Zoo Scott Neotropic Fund, The
Biological Diversity of the Guianas Program of the
Smithsonian Institution, NSF, USAID and The World Bank.

Neotropical Primates 8(1), March 2000 39

Adrian A. Barnett, School of Life Sciences, Roehampton
Institute, London, SW15 3SW, UK, and Akodon Ecological
Consulting, Bryn Mawr, PA, USA, Rebecca Shapley, Akodon
Ecological Consulting, Walnut Creek, CA 94518 USA,
Shawn Lehman, Department of Anthropology, State
University of New York, Stony Brook, NY 11794, USA,
Mireya Mayor, Department of Anthropology, State
University of New York, Stony Brook, NY 11794, USA,
Everton Henry, Everton's Ecological Tours, Georgetown,
Guyana, Paul Benjamin, Chenapou Primary School,
Chenapou village, Potaro Plateau, Section 8, Guyana, Michael
McGarrill, Chenapou Village, Potaro Plateau, Section 8,
Guyana, and Ruford Nagala, Chenapou Village, Potaro
Plateau, Section 8, Guyana.


Barnett, A. A. 1996. Primates. Expedition Field Techniques
Series No. 7. Royal Geographical Society and Expedition
Advisory Centre, London.
Bodini R. and PNrez-Hernindez R. 1987. Distribution of the
species and subspecies of cebids in Venezuela. Fieldiana,
ZooL ns. 39: 231-244.
Bowles I., Rosenfeld A. B., Sugal C. A. and Mittermeier, R.
A. 1998. Natural resource extraction in Latin American
tropics: a recent wave of investment poses new challenges
for biodiversity conservation. CI Policy Brief (1): 1-12.
Cunha, A. C. and Barnett A. A. 1989. Project Uacari: First
Report: The Preliminary Survey-, Part One Zoology. Un-
published report to WWF-Netherlands and Royal Geo-
graphical Society, London.
Eisenberg, J. E 1989. Mammals of the Neotropics. Volume One:
The Northern Neotropics Panama, Colombia, Venezuela,
Guyana, Suriname, French Guiana. The University of Chi-
cago Press, Chicago.
Emmons, L. H. and Feet, E 1989. Neotropical Rainforest
Mammals: A Field Guide. The University of Chicago Press,
Chicago. 1st edition.
Emmons, L. H. and Feet, E 1997. Neotropical Rainforest
Mammals: a Field Guide. The University of Chicago Press,
Chicago. 2nd edition.
Groombridge, B. (ed.) 1992. Global Biodiversity: Status of
the Earth's Living Resources. World Conservation Monitor-
ing Centre, Cambridge.
Hammond, D. S. and ter Steege, H. 1998. Propensity for
fire in Guianan rainforests. Conserv. Biol. 12: 944-947.
Henkel, T. 1994. Expedition Reports. Smithsonian Institution,
Department of Botany, Washington, DC.
Hershkovitz, P. 1949. Mammals of northern Colombia. Pre-
liminary report No. 4: Monkeys (Primates), with taxonomic
revisions of some forms. Proc. U. S. Nat. Mus. 98: 323-427.
Hershkovitz, P. 1977. Living New World Monkeys
(Platyrrhini), With an Introduction to Primates. Volume 1.
The University of Chicago Press, Chicago.
Huber, 0., Gharbarran, G. and Funk V. 1995. Vegetation
Map of Guyana. Centre for the Study of Biological Diver-
sity, Georgetown.

Kelloff, C. and Funk, V. In press. Preliminary Checklist of the
Plants of Kaieteur National Park, Guyana. Biological Diver-
sity of the Guianas Publication Series, Smithsonian Institu-
tion, Washington, DC.
Kinzey, W. G. 1997. Synopsis of the New World Primates
(16 genera): # 20, Cebus. In: New World Primates: Ecology
Evolution and Behavior, W. G. Kinzey (ed.), pp.248-257.
Aldine de Gruyter, New York.
Kinzey, W G., Norconk M. A. and Alvarez-Cordero, E. 1988.
Primate survey of eastern Bolivar, Venezuela. Primate
Conserve. (9): 66-70.
Lehman, S. M. 1994. Preliminary survey of Cacajao
melanocephalus in southern Venezuela. Int. J. Primatol. 15:
Lehman, S. M. 1999. The Biogeography of the Primates of
Guyana. PhD Dissertation, Washington University, St.
Louis, MO.
Lehman, S. M., Prince, W. and Taylor L. L. 1995. Habitat
disturbance, hunting pressures and primate distribution in
N.E. Guyana. Am. J. Primatol. 36: 137.
Maguire, B. and collaborators. 1948. Plant explorations in
Guiana in 1944, chiefly to the Tofelberg and the Kaieteur
Plateau. Bull. Torrey Bot. Club 75: 56-115, 189-230, 286-
323, 374-438, 523-580, 633-671.
Maguire, B. 1970. On the flora of the Guayana Highland.
Biotropica 2: 85-100.
Mittermeier, R. A. 1977. Distribution, Synecology and Con-
servation of Surinam Monkeys. PhD Dissertation, Harvard
University, Cambridge, MA.
Moonen, J. M. 1987. Hybrid capuchin in Paramaribo Zoo.
Primate Conserv. (8): 57.
Rylands, A. B., Mittermeier, R. A. and Rodrfguez-Luna, E.
1995. A species list for the New World primates
(Platyrrhini): distribution by country, endemism and con-
servation status according to the Mace-Lande System. Neo-
tropical Primates 3 (suppl.): 113-160.
Muckenhim, 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 Orga-
nization, Washington, DC.
Norconk, M. A., Sussman, R.W. and Phillips-Conroy, J. 1996.
Primates of the Guayana Shield Forests: Venezuela and the
Guianas. In: Adaptive Radiations of Neotropical Primates,
M. A. Norconk, A. L. Rosenberger and P. A. Garber (eds.),
pp. 69-83. Plenum Press, New York.
NRC, 1981. Techniques for the Study of Primate Population
Ecology. National Research Council (NRC), National Acad-
emy Press, Washington, DC.
Ochoa, G., J., Molina, C. and Giner, S. 1993. Inventario y
studio comunitario de los mamfferos del Parque Nacional
Canaima, con una lista de las species registradas para la
Guayana Venezolana. Acta Cienc. Venezolana 44: 245-262.
Phillips-Conroy, J. and Sussman, R. W. 1995. A study of the
primates of Guyana: Survey and 20-year comparison. J.
Archaeol. Anthropol. 10: 3-18.
Reid, E 1998. A Field Guide to the Mammals of Central
America and South-east Mexico. Oxford University Press,

Neotropical Primates 8(1), March 2000

Robinson J. G. and Ramirez C. J. 1986. Conservation biol-
ogy of Neotropical primates. In: Mammalian Biology in
South America, M. A. Mares and H. H. Genoways (eds.),
pp. 329-344. Special Publication Series No. 6, Pymatuning
Laboratory, University of Pittsburgh, Linesville. PA.
Roussillhon C., 1988. The general status of monkeys in French
Guiana. Primate Conserv. (9): 70-74.
Schiirholz, G. 1991. Proposal for Expansion and Manage-
ment of Kaieteur National Park, Guyana. Unpublished re-
port to Guyana National Parks Commission, Guyana Agency
for Health and Science Education, Environment and Food
Policy, the Ministry of Trade and World Wildlife Fund (US).
Schulze, M. D., 1998. Forest fires in the Brazilian Amazon.
Conserv. Biol. 12: 948-950.
Sussman, R. W and Phillips-Conroy, J. 1995. A survey of the
distribution and diversity of the primates of Guyana. Int. f
Primatol. 16: 761-792.
Tate, G. H. H. 1932. Life zones at Mount Roraima. Ecology
13: 235-257.
Tate, G. H. H. 1939. The mammals of the Guiana region.
Bull Am. Mus. Nat. Hist. 76: 151-229.
World Bank. 1998. Guyana: National Protected Areas System
Project. The World Bank, Washington, D. C.


Sylvain Charlat
Oliver R. Thatcher
Nils Hartmann, Yogen G. Patel
Marjorie Saillan and Elsbeth Vooren


Howler monkeys (Alouatta) are among the larger New World
primates. They are found living in a wide range of forest habi-
tats over most of South and Central America from Mexico to
Argentina, with a total distribution of 483,000 km2 (Marsh
and Mittermeier, 1987). The mantled howler monkey
Alouatta palliata, can be found from southern Mexico,
through Central America, western Colombia and Ecuador
to the northwest of Peru (Eisenberg, 1989). Throughout this
geographical range, A. palliata occupies several different
forest types: dense primary forest in many places, but also
coastal mangrove forest in Panama, dry, deciduous forest in
Costa Rica, secondary forest in Mexico and mountain forest
in Guatemala (Wolfheim, 1983). Strong variations in density
have been observed between these different habitats (Peres,
1997; Chapman and Balcomb, 1998). Thus, an important
issue for A. palliata conservation is a better understanding of
the links between habitat characteristics and population
densities. In this paper, we report the results of a population
survey of A. palliata at the Bilsa Biological Station of Ecua-
dor. To our knowledge, this species has not previously been
studied in Ecuador; nothing is known about their local
ecology or abundance.


Alouatta palliata
Adult males of A. palliata weigh around 7.3 kg and females
around 5.8 kg (Martin, 1990). The basic color of both sexes
is black, sometimes brown, with a gold, white or brown fringe
along the flanks (the mantle). The colour and shape of the
mantle varies greatly within the species. A. palliata
aequatorialis, the subspecies we have been working on, is listed
as "Lower Risk" by Crockett (1998). The same status is given
for A. palliata palliata. However, the subspecies A. palliata
mexicana is listed as "Vulnerable" by Crockett (1998) as well
as in the 1996 IUCN Red List of Threatened Animals.

Study Site
The Bilsa Biological Station (00'20.8' N, 79042.7' W) (Fig.
1) is situated in the foothills of the Mache-Chindul Mountains
in the Province of Esmereldas, at an altitude of 300-750m, in
the western coastal region of Ecuador. The reserve covers 3000
hectares of some of the last remaining tropical pre-montane
wet and humid coastal forest in Ecuador. A small fraction of
the area is composed of very recent secondary forest, now
submitted to reforestation. Altitude variations and moisture
from the Pacific Ocean create microclimates that encourage
local species endemism (Brame, 1995). Robin Foster (in Parker
III and Carr, 1992) describes the area as pristine. The
surrounding area has only been disturbed by colonisation and
logging in the last 30 years. The temperature is fairly stable
year round at Bilsa, and the climate is described as uniform
temperate wet (Parker III and Carr, 1992). Average rainfall
is often more than 3 m of rain per year. In the wet season,
from January to June, the temperature range is usually
between 24C and 25C. In the dry season, from July to
December, the range is between 21C and 22C. During
the time of our study (4th August to 10th September), the
average temperature was 21.60C. The maximum tempera-
ture recorded during that time was 27.5C and the minimum

Bilsa station N

a ,.- -- .

---. Res ........ Re
boundaries ..,* sc e
......... M ajor riers 60 /
600 m
Approximate boundaries Totalarea
ofthe area covered by Lnangulaborn 3000 hectares
Observed hov.ler troops. troop size

Figure 1. Map of the Bilsa Biological reserve, from Carlos Aulestia.

Neotropical Primates 8(1), March 2000

was 18.20C. The average precipitation was 1.6 mm per day
and the total precipitation for 30 days was 48.7 mm.

The Survey: Troop Density Estimation
We took advantage of the fact that howlers make very loud
calls at dawn, which allowed us to use a triangulation method
(Milton, 1982; Stoner, 1994; Brockelman and Ali, 1987).
The idea of this method is to place as many groups of people
into the field as possible in order to record simultaneously
the time and the direction of calling troops. The groups of
listeners are placed at known, predetermined locations on the
map. Calls from one monkey troop at a specific time were
recorded by different observers, so that the point where the
direction lines crossed indicated the position of a monkey
troop. This method provided an estimation of the minimum
density since not all troops may not howl on a given morning.

Data were collected on 20 August, 1998 between 05.00 a.m.
to 08.00 a.m. with the help of 34 volunteers (17 groups of
two). Volunteers included staff from the station, park guards,
European university students and North American peace-
corps volunteers. Group sites were determined the day before
and marked with GPS. Sites on hills allowing the best listening
were preferred. The positions of the listeners and the
topography allowed us to estimate the area covered (positioned
on Fig. 2) as 4 km2. The precise times and directions of the
first howls of each troop were noted. As only seven compasses
were available, north was clearly marked at all sites.

A. A

....... ........ ......

"'" .. .SO.nOm

Figure 2. Results of the triangulation. Triangles represent
listener pairs. Circles represent howler troops. Doted lines
represent directions allowing to locate howler troops. As
an example, the troop located at the top of this map as been
heard by six listener pairs at the same time.

The Survey: Troop Size Estimation
Troop sizes were determined by direct observations. We
looked for howler troops in different randomly chosen areas.
We formed three groups of two and worked in shift. One
team worked from 5 a.m. to 2 p.m., one from 5 a.m. to 5
p.m., and the third from 2 p.m. to 5 p.m. As a consequence
there was at least one group of observers in the field at any
time of the day, and two for most of the day. Each area was
surveyed for three days. The researchers walked throughout
the day, with occasional stops to listen for howler calls and
cracking branches that indicated monkey troops. Using troop
size and composition as well as GPS position allowed
identification of the different troops encountered. All
observations using this method between 4 August and 10
September,1998, were recorded. Troops were studied until
all observers agreed on their size. Two major observations by
park guards were added to the data.
Mapping of the area and the positions of the troops used a
Global Positioning System (Garmin 12). Field notes were taken
on AquaScribe waterproof notebooks.


Troop Size
The location and size of the troops encountered is shown in
Figure 1. Of a total of 12 encounters,two were solitary indi-
viduals. Troop sizes ranged from two to nine animals.
Including solitary individuals, mean troop size is estimated at
N = 4.8 individuals per troop (SD 3.1).

Troop Density
Thirteen troops were detected in an area of 4 km2 (Fig. 2).
Troop density is therefore estimated to be 3.25 troops/km2 in
the covered area. Using N = 4.8 (SD 3.1), we estimate the
density to be 15.6 individuals/km2 ( 9.3). Given that the
area surveyed covers 1/8 of the whole reserve, and that it
roughly presents the different habitat types of Bilsa, and in
similar proportions, extrapolation of the density estimate to
the whole reserve seems justified.


Ecological factors affecting howler density at Bilsa
Our survey indicates a density of 15.6 individuals/km2 ( 9.3).
This density falls below the mean observed for the genus, es-
timated by Peres (1997) as 29.5 (estimation based on 106
densities for different habitat types). However, it is important
to notice that many sites reported in this study show low
densities while only a few show extremely high ones. Hence,
even though the density observed in Bilsa is inferior to the
mean for the genus, it is superior to the median (mean of
ranks: 12.8). The same kind of conclusion can be drawn from
literature data concerning A. palliata only (Table 1). Indeed,
we record that even though some sites harbour very high
densities (some of them (Baldwin and Baldwin, 1976) being
probably very unstable and due to recent disturbance in
surrounding areas), densities around 15 individuals / km2 are

Neotropical Primates 8(1), March 2000

Table 1. Alouatta palliata population densities from various sites.

Site Densides i/kln ) Source
Barro Colorado Island Pan -.a 16 2 C.'lli and Souith' id.. I Cl2,
Barro Colorado I:and PanT,,'a 2 -52 Carpenr i l'3 3- 19o2
Burro Colorado lland. Panama 62 Chi.ceri I 196'
B1rro Colorado land. Pan ima 62 6 MitcermeNer I I-3
Barro Lolorado .land Par,'-na -r S Sm',,h i 11 i
barro Colorid blind Par, ama 82.2-91.- MKlion I 152
Chirique P'anama IrO4 Bald," n and Bid,, in i '-l
La ['iFcica, (-oC ii R,, -4.3 H lint t,-.d I'l'i

La Pict'iLa. Co',ii R.,:

La ['scd i C ,: RJcaj
Lu. Tu.-Jr Ic'.lco
Prak \Verde Co. Ea RI c
Rio Jesus, Costa Rica 1
Rio Jesus, Costa Rica 2
Santa Rosa, Costa Rica
Santa Rosa, Costa Rica
Santa Rosa, Costa Rica

$9 -
10 1

Thus, it seems that the howler density in Bilsa is low compared
to what is seen in some habitats, but not compared to what is
ordinary observed. We will now try to explain this pattern by
considering the different ecological factors known to affect
howler density: hunting pressure, interspecific competition,
and food quality and availability. These factors are ultimately
determined by abiotic factors such as soil fertility, altitude,
seasonality and rainfall.

Hunting pressure has been shown to be a very significant fac-
tor affecting the population structure of howler monkeys
(Peres, 1997; Marsh and Mittermeier, 1987). Howlers are
hunted for food, medicine and fur, which has resulted in their
extinction from many areas, including parts of Ecuador (M.
Dilger, pers. comm.). However, discussion with park guards
as well as personal observations suggest that hunting pressure
is very low or non-existent at Bilsa.

Inter-specific competition for resources can affect howler
population densities in communities rich in primate species
(Eisenberg, 1979). Cebus albifrom (white-fronted capuchin)
is the only other monkey species present at Bilsa. However,
capuchins seem rare and are found mainly in secondary for-
est. Therefore the resource overlap is small, suggesting that
inter-specific competition is not likely to be a factor affecting
the observed howler troop size and population density at Bilsa.

The floristic composition can have a profound influence on
the population structure of howler monkeys. The flora of Bilsa
is remarkable by the fact that it contains low numbers of
Leguminosae and Ficus species (R. B. Foster in Parker III and
Carr, 1992), which have been shown to be very important
components of howler diet. Therefore, it may be that howler
density and troop size is limited in Bilsa because of floral com-

(Inddr I I"-' i
ldark, ind Zu ktr i I"1,
Sr-iner ( -iI
FihLmjrd and Su.,rmar1 ]"oi,
E:itrjdja I1S2
Rodricuc. l I0551
Sanchez Porrin I 1"c,l I
Sanchez P,;,rr. I I"l I
Fr.:ee (1976)
(-hapmair. ,; 1NlS
FIdigar : '9181

position. However, howler diet should be precisely ascertained
for this hypothesis to be confirmed.

Based on a wide comparison of population densities at different
sites, Peres (1997) concluded that once hunting pressure dif-
ferences are controlled for, two remaining factors are the main
determinants of Alouatta densities: (i) primary productivity
(which is increased by soil fertility and forest heterogeneity)
and (ii) toxin concentrations in leaves (which is decreased by
seasonality and soil fertility). Bilsa is composed mainly of pri-
mary forest, probably harbouring high concentrations of toxic
secondary compounds in leaves with limited primary produc-
tivity. Seasonality is significant but not as important as in sites
showing the highest howler densities. Furthermore, because
the area is mountainous, leaching of nutrients from the soil is
to be expected.

Thus, it seems that hunting pressure and interspecific com-
petition are not limiting howler density at Bilsa. However,
low abundance of food sources commonly consumed by
howlers, together with relatively low primary productivity and
high toxin concentrations may limit the carrying capacity of
this habitat.

Conservation Implications
The original distribution of the tropical rain forest and the
populations of A.palliata has been reduced by at least 90 per-
cent in the last 40 years as a result of the conversion of natural
habitat to pasture and agricultural fields (Estrada and Coates-
Estrada, 1988). Conservation initiatives are needed to save
this species from extinction. Habitat fragmentation and de-
struction, hunting, and pet trade are the factors responsible
in most cases for the decline of the species. In the Mache-
Chindul Mountains, hunting and pet trade do not seem to

Neotropical Primates 8(1), March 2000 43

play a role, however deforestation has been incredibly intense
during the last 30 years.

Howler conservation requires preservation of primary forest
habitat, as this is the type of habitat that is suitable for these
animals. Our results show that the Bilsa population is not
currently at a critical stage. The surrounding areas of Bilsa
are still covered by primary forest at the present time. These
areas must be protected from destruction by including them
in the reserve as well as continuing the education activities of
Fundaci6n Jatun Sacha to increase the awareness of the local
people. Furthermore, the effects of overall population
reduction due to the destruction of the outlying forest may
not yet have begun to show the long- term deleterious effects
of genetic diversity loss and genetic drift.

Howler conservation at Bilsa is an excellent way of protecting
the whole habitat. Howlers are probably among the largest
animals in the area and their presence is undoubtedly necessary
for the stability of the ecosystem, especially in the role of seed
dispersal (Estrada and Coates-Estrada, 1984). Given the vast
number of plant species consumed by howlers their continued
presence requires the protection of the entire ecosystem. The
mantled howler monkey is therefore an effective umbrella

Surveys are indispensable for conservation planning. They al-
low estimations of population status and provide material for
inter-site comparisons. Surveys must be performed several
times to provide information concerning population dynamics
which allows the recognition of declining primate populations
in areas where conservation efforts are most needed. More
detailed information must be obtained, particularly on how
the A. palliata diet in Bilsa differs from other sites. Local
alimentary habits must be known for conservation actions to
be conducted through reforestation. As a peculiar and unique
habitat, Bilsa may reveal many new and interesting aspects of
howler ecology. Sufficient data are already available in the
literature to allow fruitful comparisons. Hopefully such
projects will be realized at Bilsa in the near future.


We are deeply grateful to Prof. M. Hollingdale and the Uni-
versity of Leeds School of Biology, the charity Equafor, the
Expedition Committee, Travel Bursaries, Roger Stevens,Travel
Bursary of the University of Leeds, the Leeds Philosophical
and Literary Society, the Gilchrist Educational Trust, the staff
of the School of Biology and especially Prof. R. McNeill
Alexander, for funding this study. The following companies
contributed equipment, for which we are grateful: Johnson-
Progress Ltd, Hawkins & Manwaring Ltd, Chefaro Propri-
etaries Ltd, Stafford Miller Ltd, Johnson & Johnson Ltd, Oasis
Nets Ltd, and Care for the Wild International. Thanks to A.
Davis for advice during the preparation. Thanks to Frangoise
Bayart for advice and comments on the manuscript. And last
but not least, many thanks to the staff of the Fundaci6n Jatun
Sacha and the Bilsa Biological Station for help conducting
the triangulation.

Sylvain Charlat, Institut Jacques Monod, Laboratoire
Dynamique du Genome et Evolution, Tour 42 41me 6tage, 2
Place Jussieu, 75251 Paris Cedex 05, France, e-mail:
, Oliver R. Thatcher, Department
of Entomology, Natural History Museum, Cromwell Road,
London, SW7 5BD, UK, Nils Hartmann, Molekulare
Parasitologie, Institut fuer Biologie, Humboldt-Universitaet
Berlin, Philippstr. 13, 10115 Berlin, Germany, Yogen G.
Patel, Department of Biological Anthropology, University
College London, Gower Street, London WC1E 6BT, UK,
Marjorie Saillan, Institut Jacques Monod, Laboratoire
Dynamique du Genome et Evolution, Tour 42 4m" 6tage, 2
Place Jussieu, 75251 Paris Cedex 05, France, and Elsbeth
Vooren, Institut Jacques Monod, Laboratoire Dynamique du
Genome et Evolution, Tour 42 4am" 6tage, 2 Place Jussieu,
75251 Paris Cedex 05, France.


Baldwin, J. D. and Baldwin, J. I. 1976. Primate populations
in Chiriqui, Panama. In: Neotropical Primates. Field Studies
and Conservation, R. W Thorington and P G. Heltne (eds.),
pp.20-31. National Academy of Science, Washington, DC.
Barnett, A. 1995. Expedition Field Techniques: Primates. The
Expedition Advisory Centre, Royal Geographical Society,
Brame, C. W. 1995. The Bilsa Reserve Biological Station,
north-west Ecuador, Management plan. Fundaci6n Jatun
Sacha, Quito, Ecuador (Unpublished).
Brockelman, W. Y., and Ali, R. 1987. Methods of surveying
and sampling forest primate populations In: Primate Con-
servation in the Tropical Rain Forest, C. W. Marsh, and R.
A. Mittermeier (eds), pp.23-62. Alan R. Liss, New York.
Carpenter, C. R., 1934. A field study of the behaviour and
social relations of howling monkeys. Comp. Psychol. Monogr.
(10): 1-168.
Carpenter, C. R., 1962. Field studies of a primate popula-
tion. In: Roots of Behavior, E. L. Bliss (ed.), pp.286-294.
Harper and Row, New York.
Chapman, C. A., Chapman, L. J. and Glander, K. E. 1989.
Primate populations in northwestern Costa Rica: Potential
for recovery. Primate Conservation (10):37-44.
Chapman, C. A. and Balcomb, S. R. 1998. Population char-
acteristics of howlers: ecological conditions or group his-
tory. Int. J. Primatol. 19: 385-403.
Chivers, D. J.1969. On the daily behaviour and spacing of
howling monkey groups. Folia Primatol. 10:48-102.
Clarke, M. R. and Zucker, E. L. 1992. Survey of the howling
monkey population at La Pacifica: as seven-year follow-up.
Int. J. Primatol., 15: 61-73.
Collias, N and Southwick, C. 1952. A field study of popula-
tion density and social organization in howling monkeys.
Proc. Am. Phil. Soc. 96: 143-156.
Crockett, C. M. 1998. Conservation biology of the genus
Alouatta. Int. J. Primatol. 19:549-578.
Eisenberg, J. E 1979. Habitat, economy and society: some
correlation and hypotheses for the Neotropical primates.
In: Primate Ecology and Human Origins: Ecological Influ-
ences on Social Organization, I. S. Bernstein and E. 0. Smith
(eds), pp.215-262. Garland Press, New York,
Eisenberg, J. F. 1989. Mammals of the Neotropics. Vol. 1.
Panama, Colombia, Venezuela, Guyana, Suriname, French

Neotropical Primates 8(1), March 2000

Guiana. The University of Chicago Press, Chicago.
Eisenberg, J. E and Thorington, R. W. 1973. A preliminary
analysis of Neotropical mammal fauna. Biotropica 5: 150-
Estrada, A. 1982. Survey and census of howler monkeys
(Alouattapalliata) in the rain forest of "Los Tuxtlas" Veracruz,
Mexico. Am. J. PrimatoL 2: 363-372.
Estrada, A. and Coates-Estrada, R. 1984. Fruit eating and
seed dispersal by howling monkeys (Alouatta palliata) in
the tropical rain forest of Los Tuxtlas, Mexico. Am. J.
Primatol. 6: 77-91.
Estrada, A. and Coates-Estrada, R. 1988. Tropical rain forest
conversion and perspectives in the conservation of wild pri-
mates (Alouatta and Ateles) in Mexico. Am. J. Primatol. 14:
Fedigan, L. M., Rose, L. M. and Morera Avila, R. 1998.
Growth of mantled howler groups in a regenerating Costa
Rican dry forest. Int. J. Primatol. 19: 405-432.
Fiskind, A. S. and Sussman, R. W 1988. Preliminary survey
of the primates of the Zona Protectora and La Selva Bio-
logical Station, northeast Costa Rica. Primate Conserv. (8):
Freese, C. 1976. Censusing Alouatta palliata, Ateles geoffroyi
and Cebus capucinus in the Costa Rican dry forest. In: Neo-
tropical Primates. Field Studies and Conservation, R. W.
Thorington Jr. and P. G. Heltne (eds.), pp.4-9, National
Academy of Science, Washington, DC.
Glander, K. E., 1978. Howling monkey feeding behaviour
and plant secondary compounds: A study of strategies. In:
The Ecology ofArboreal Folivores, G. G. Montgomery (ed.),
pp.561-573. Smithsonian Institution Press, Washington,
Heltne, P G., Turner, D. C. and Scott, N. J. 1976. Compari-
son of census data on Alouatta palliata from Costa Rica and
Panama. In: Neotropical Primates. Field Studies and Conser-
vation, R. W Thorington Jr. and P. G. Heltne (eds.), pp. 10-
19. National Academy of Sciences, Washington, DC.
IUCN Red List of Endangered Animals. Website:http://
www.wcmc.org.uk/species/ animals/animal_redlist.html
Marsh, C. W. and Mittermeier, R. A. 1987. Primate Conser-
vation in the Tropical Rain Forest. Alan R. Liss, New York.
Martin, R. D. 1990. Primate Origins and Evolution.
Chapman and Hall, London
Milton, K. 1980. The Foraging Strategy of Howler Monkeys: A
Study in Primate Economics. Columbia University Press, New
Milton, K. 1982. Dietary quality and demographic regula-
tion in a howler monkey population. In: The Ecology of a
Tropical Forest: Seasonal Rhythms and Long-term Changes,
E. G. Leigh Jr, A. S. Rand, and D. M. Windsor (eds.),
pp.273-290. Smithsonian Institution Press, Washington,
Mittermeier, R. A. 1973. Group activity and population dy-
namics of the howler monkey on Barro Colorado Island. Pri-
mates 14: 1-19.
Myers, N. 1988. Threatened biotas: Hotspots in tropical for-
ests. The Environmentalist 8: 187-208.
Parker, T. A., III and Carr, J. L. 1992. Status of Forest Rem-

nants in the Cordillera de la Costa andAdjacentAreas ofSouth-
western Ecuador. RAP Working Papers 2. Conservation In-
ternational, Washington, DC.
Peres, C. 1997. Effects of habitat quality and hunting pres-
sure on arboreal folivore densities in Neotropical forests: A
case study of howler monkeys (Alouatta spp.). Folia Primatol.
68: 199-222.
Rodriguez, M. A. R. 1985. Algunos aspects sobre
compartamiento, alimentaci6n y nivel de problemaci6n de
los monos (Primates:Cebidae) en el Refugio de Fauna
Silvestre Palo Verde (Guanacaste, Costa Rica). In:
Investigaciones Sobre Fauna Silvestre de Costa-Rica, Edito-
rial Universidad a Distancia, San Jose, Costa Rica.
Sanchez Porras, R. E. 1991. Utilizaci6n del habitat,
comportamiento, y dieta del Mono Congo (Alouatta
palliata) en un bosque premontano humido, Costa Rica.
Master's thesis, Universidad Nacional, Costa Rica.
Smith, C. C. 1977. Feeding behavior and social organization
in howling monkeys. In: Primate Ecology, T. H. Clutton-
Brock, (ed.), pp.97-126. Academic Press, London.
Stoner, K. E. 1994. Population density of the mantled howler
monkey (Alouatta palliata) at la Selva Biological Reserve,
Costa Rica: A new technique to analyze census Data.
Biotropica 26:332-340.
Wolfheim, J. H. 1983. Primates of the World: Distribution,
Abundance and Conservation. Harwood Academic Publish-
ers, Seattle.


Jessica Ward Lynch
Jose Rimoli


Demographic variables play an important role in
understanding primate behavioral ecology (Dunbar, 1987;
Strier 1991, 1999a) and are crucial components of
conservation biology strategies for species' management
(Dobson and Lyles, 1989). While attention has focused on
collecting demographic data on endangered species, it is also
important to monitor demography of abundant and
widespread species, both for comparison to those which are
threatened (Hubbell and Foster, 1986) and for understanding
dynamics among species in primate communities (Waser,
1987; Strier, 1999b). Long-term studies in the Amazon, the
Ilanos of Venezuela, and Central America have yielded
demographic information on groups of Cebus apella (Izawa,
1988, 1990, 1992, 1994a, 1994b, 1997), C. olivaceus
(Robinson, 1988a, 1988b), and C capucinus (Fedigan et al.,
1996), but, until recently, there was no data available for the
capuchin monkeys of the Atlantic forest in Brazil. Here we
present information on group composition and membership

Neotropical Primates 8(1), March 2000 45

Table 1. Age-sex composition of a group of Cebus apella
nigritus at the Estacgo Biol6gica de Caratinga, Minas

July Jan July Jan July Jan
95 96 96 9" 9- 98

A1 (1 -
SAM 1 3 2 2 I I
SAF 0 A 1 2 4
JUT \ 10 I1) il i" C6
INF 2 i i 2 2 ,

AM=adult males, AF=adult females, SAM=subadult males,
SAF=subadult females, JUV=juveniles, INF=infants. Age
categories follow Izawa (1980).

dynamics of a group of tufted capuchin monkeys (Cebus apella
nigritus) in a fragment of Atlantic forest at the Estacao
Biol6gica de Caratinga (EBC), in Minas Gerais, Brazil.


The Estaqao Biol6gica de Caratinga is a semi-deciduous forest
fragment of 890 ha (see Strier, 1987, 1992; Ferrari, 1988,
and Rimoli and Ades, 1997, for more information on the
site). The study was carried out from June 1995 through
August 1997. When possible, the study group was followed
from daybreak to nightfall, and visual contact was made on
more than 340 days. Group sizes of the capuchins at EBC
ranged from approximately five to 30 individuals, although
the smaller "groups" may have been subgroups of the larger
ones (Lynch, 1999a, 1999b). Individuals were recognizable
through pelage patterns and facial characteristics. Group
membership was recorded daily. Data were collected ad
libitum on births, emigrations, disappearances and changes
in social structure. Follow-up censuses were conducted in
November 1997 and January 1998.


The study group ranged in size from 26-29 individuals,
including 3-5 adult males, 6-7 adult females, 3-7 subadults,
6-11 juveniles, and 2-6 infants (Table 1). The mean sex ratio
of adult males to females for the EBC group was 0.61 0.135
(range 0.43-0.83), or one adult male per 1.65 adult females.

Group size and age-sex composition were relatively stable.
However, over the 30-month study period, membership
changed due to emigrations, births, and disappearances
(Rfmoli and Lynch, 1999). The alpha male was expelled
following a take-over by the group's beta male. One adult
female and two subadult males left the group and were later
seen in other groups; two subadult females disappeared from
the group and were not seen again. Thirteen infants were
born during the study, five of which disappeared along with
one juvenile. An analysis of infant mortality restricted to infants
born during the study and followed for one year or until
disappearance, indicated survivorship to one year as being

~. ~"j

lun 1d au spt oc no5 dec Ja f mar a"t m

Figure 1. Capuchin births by month, compared to mean rainfall
by month at EBC, from June 1995 to December 1997.

only 45% (four of nine infants). The disappearance of two of
the infants occurred at the same time as the take-over of the
group by the beta male. All births occurred during the rainy
season, and there was a birth peak between October and
March (Rimoli and Lynch, 1999). The total number of births
by month (1995-1997) is compared to mean rainfall per
calendar month for the same time period in Figure 1.

The interbirth interval (IBI) was 25 months for the one female
with a surviving infant who gave birth a second time during
the study. No second birth was recorded for another female
with a surviving infant over 26 months. For the five IBIs in
which the first infant died or disappeared, the mean IBI was
11.2 +1.79 months (range 9-14 months).


Group size in the genus Cebus ranges from solitary individuals
up to 50 or more (Figure 2). The Atlantic forest studies of
Cebus apella nigritus extend the upper range of Cebus apella
group size. C. a. nigritus grouping patterns appear more
similar to those of C. olivaceus and C capucinus than to some
Amazonian C. apella populations (Janson, 1985), or to the
heavily hunted populations of C. apella margaritae (see Sanz
and Mirquez, 1994) on Margarita Island, Venezuela, and C.
xanthosternos (see Pinto and Tavares, 1993) in southern Bahia,

Interbirth intervals at EBC were similar to those reported for
capuchins at other sites (Table 2). The range for IBIs are
overlapping for all capuchin species, and as with other
capuchins, the loss of an infant allows EBC females to decrease
their IBI by about one year.

Capuchin births at EBC were concentrated in the wet season.
This suggests that infants are born during a period of relative
food abundance, since the rainy season is characterized by a
high number of fruiting and flowering trees (Lopes and
Andrade, 1986; Ferrari, 1988; Strier, 1991; Rimoli and Ades,
1997), as well as an increase in the number of available insects
(Ferrari, 1988). C. apella nigritus at Iguazi, Argentina also
have a birth season from October to February, during the

46 Neotropical Primates 8(1), March 2000

Table 2. Interbirth intervals for the genus Cebus.

Species Mean IBI in months after infain Mean IBI in months after Source
survival In of IBis: range) infant loss In of IBis; rangel
C .:p.'!. 25 in= IL 11.2 = I S In=5' '- 141 Present studv
C. ape/i 25.6 1 3.8&1n=10; 21-35) I' -- =5 9 In--1. 9-211 [Lawa, 1988. 1g90, 1992.
199-a, 199-4b. 1997
C..i.et. 22 Nor rcporiced Robinson & lanson, 19)8S
C. ,doa:b n, 18 No, rcponrejd Robinson & lanson, 1987
C .,:,".:., 26.9 In=23- 16 0 n=2' Fedigan & Roie. 1995
C .i ,:, 22 in=-; 12.5- q1 i10 rl=3.9S- I 31 FPerrs. 199'
C oheace., 26 Noi rcponed Robinson & [anson. 19S-
Note: IBIs reported in Robinson and Janson, 1987.

peak availability of fruits and insects (di Bitetti, 1997). Late
dry and early rainy season births are most frequent in Cebus
apella in Colombia (Izawa, 1988, 1990, 1992, 1994a,
1994b, 1997) and Peru (Janson, 1985). In C olivaceus, births
are most common at the end of the dry season and the
beginning of the wet season (Robinson, 1988a). By contrast,
in Panama and Costa Rica, C. capucinus births occur
throughout the year and may be more common in the dry
season (Oppenheimer, 1982; Robinson and Janson, 1987;
Fedigan and Rose, 1995).


Fifty-five percent of capuchin infants died in their first year
in the EBC group. This is the highest rate of infant mortality
reported for capuchins. In comparison Cebus apella in
Colombia has 19% infant mortality in the first year (Izawa,
1988, 1990, 1992, 1994a, 1994b, 1997), C. capucinus has
29% (Fedigan et al., 1996) and C. olivaceus, 19.4%
(Robinson, 1988a). In fact, a wide comparison of infant
mortality across primate taxa suggests that the rate at EBC is
unusually high (see Robinson, 1988a for summary). However,
Costa Rican squirrel monkeys (Saimiri oerstedt) have shown a
similarly high rate of infant mortality, due to avian predation
(Boinski, 1987).

EBC. BR t)
Sac Paulo BR 12)
Iguan Falls, AR (3)
La Macarena, CO (4-9)
isla de Margarita, VE (10)
Cahuana Isjand, PE (11i

Coshtu Cashu. PE (12)
Southern Bahia. BR 113)
C. CAPUCUJUS Santa Rosa, CR (14)
Santa Rosa CR(15)
Barro Colorado, PN (16)
C. OLIVACEU Hato Pmero. VE (17)

Hato Masaguaral. VE (18)
C. ALBIFRONS: El Tuparro, CO (1)j)
Coshu Cashu, PE (12)
caouona Island, PE (11)


0 5 10 15 20 25 30

35 4 45

Figure 2. Group size within the genus Cebus.

BR=Brazil, AR=Argentina, CO=Colombia, VE=Venezuela, PE=Peru, CR=Costa Rica, PN=Panama
All bars are ranges in group size except 2, 14, and 19 which are mean group size. 14 understimates variation in
group size as it is the range of MEAN group size for the population. (1) present study (2) Izar, 1999 (3) Janson,
1996 (4-9) Izawa, 1988, 1990, 1992, 1994a, 1994b, 1997 (10) Sanz and Marquez, 1994 (11) Soini, 1986 (12)
Janson, 1985 (13) Pinto and Tavares, 1993 (14) Chapman, 1990 (15) Fedigan etal., 1996 (16) Phillips, 1994 (17)
Miller, 1996 (18) Robinson, 1998a (19) Defler, 1979. Early reports of capuchin group size are summarized in
Freese and Oppenheimer, 1981.

Neotropical Primates 8(1), March 2000

Infant mortality in the EBC group may be the result of a
combination of factors. Predators at EBC include tayras (Eira
barbara), coatimundis (Nasua nasua), ocelots (Felis pardalis),
hawks (Buteo spp.), caracards (Polyborus plancus) and vultures
(Sarcoramphus papa and Coragyps atratus), as well as semi-feral
dogs. Capuchins give strenuous alarms and threats and will
change travel patterns in response to tayras, hawks and dogs.
When these predators are present, capuchin females will often
carry large infants or juveniles that are otherwise independent
(Rimoli and Lynch, unpubl. data). While there may be fewer
predator species at EBC than at other field sites in more
continuous forest, capuchins at EBC may be more restricted
in their ability to escape from predators because it is a forest
fragment. Capuchins at EBC use edge habitat and venture
out into corn and sugar cane fields (Rimoli and Ferrari, 1997),
which may put them at higher risk to both aerial and terrestrial

Intragroup dynamics may also contribute to infant mortality.
The instability of the group during the take-over period in
May 1996 coincided with the disappearance of two infants.
Although infanticide was not witnessed in this study,
infanticide has occurred in the same context in other Cebus
species (C. olivaceus, Valderrama et al., 1990; C. capucinus,
Fedigan et al., 1996). One difference at EBC was that the
take-over occurred from within the group; the alpha male
lost his position to a beta male who had been with the group
for at least one year. In C. capucinus (Fedigan et al., 1996),
and in fact in the vast majority of observed infanticides among
primates in general (see review in Struhsaker and Leland,
1987), it is usually new males entering the group that are
associated with male take-overs and infanticide. Calculating
conception dates (using a gestation length of 149-158 days,
from Robinson and Janson, 1987) for the individuals that
disappeared in the year after the new male was already in
power, one infant and one young juvenile would have already
been conceived before the take-over had occurred. This may
be more evidence for an infanticide hypothesis. However,
another two infants, both lost from the group in March 1997,
had been conceived after the new alpha male was in place.
These two infants were last seen two months before the mating
peak in May 1997. The females that lost these infants did
mate and conceive again during that period. Interestingly,
these were the two females that showed the most mating
fidelity to the new alpha male during 1997. One, TE, mated
exclusively with the alpha male, and the other, PT, was
interrupted by the alpha male in her one witnessed copulation
attempt with a subordinate male; all other sexually active adult
females in the group had 3-4 mating partners (Lynch, 1998).

Subgrouping may be another potential risk factor for
capuchin infants at EBC. The study group broke up into
smaller subgroups on nearly half the observation days in the
1996-1997 season (Lynch, 1999a, 1999b). On at least one
occasion during that time, a dependent infant was stranded
for several hours in a subgroup different from his mother.
Capuchin infants may be carried by group members other

than their mothers even in the first months of life; a young
infant primarily dependent on mother's milk might be at
serious risk of dehydration or starvation if separated from her
in this way for one or more days.
Despite the fact that deforestation around EBC has restricted
these primates from contact with a larger population for over
50 years (Strier, 1991), the high infant mortality within the
group does not seem to be associated with inbreeding effects
(Lande and Barrowclough, 1987). The capuchin monkeys
here have never been reported to have obvious congenital
defects. Further study of this and other groups at EBC will
be needed to test whether the high infant mortality found in
this study is an ongoing cost to groups of large size that split
up into subgroups, a short-term phenomenon related to a
change in the position of alpha male, a result of increased
predator pressure in edge habitat, or some combination of
these various social and ecological factors.


Research was funded by the Brazil Higher Education
Authority (CAPES), Liz Clairborne and Art Ortenberg
Foundation (Dr. Karen Strier), the Fulbright Foundation and
Wenner-Gren #6068. Special thanks to Dr. Karen Strier for
supporting this study and for useful comments on the

Jessica Ward Lynch, Department of Anthropology,
University of Wisconsin-Madison, 1180 Observatory Drive,
Madison, WI 53706, USA, and Jose Rfmoli, Universidade
Federal do Pari, Centro de Ciencias Biol6gicas, Caixa Postal
8607, 66075-150, Belkm, Para, Brasil.


Di Bitetti, M. 1997. Evidence for an important social role of
allogrooming in a platyrrhine primate. Anim. Behav. 54:
Boinski, S. 1987. Birth synchrony in squirrel monkeys (Saimiri
oerstedi): a strategy to reduce neonatal predation. Behav. Ecol.
Sociobiol. 21: 393-400.
Chapman, C. A. and Fedigan, L. M. 1990. Dietary differ-
ences between neighboring Cebus capucinus groups: local
traditions, food availability or responses to food profitabil-
ity? Folia Primatol. 54: 177-186.
Defler, T. 1979. On the ecology and behavior of Cebus
albifrons in eastern Colombia: I. Ecology. Primates 20(4):
Dobson, A. P., and Lyles, A. M. 1989. The population dy-
namics and conservation of primate populations. Conserv.
Biol. 3: 362-380.
Dunbar, R. I. M. 1987. Demography and reproduction. In:
Primate Societies. B. Smuts, D. Cheney, R. Seyfarth, R.
Wrangham and T. T. Struhsaker (eds.), pp.69-82. The Uni-
versity of Chicago Press, Chicago.
Fedigan, L. M. and Rose, L. M. 1995. Interbirth interval
variation in three sympatric species of neotropical monkey.
Am. J. Primatol. 37: 9-24.

Neotropical Primates 8(1), March 2000

Fedigan, L. M., Rose, L. M. and Avila, R. M. 1996. See how
they grow: Tracking capuchin monkey (Cebus capucinus)
populations in a regenerating Costa Rican dry forest. In:
Adaptive Radiations ofNeotropical Primates, M. A. Norconk,
A. L. Rosenberger and P. A. Garber (eds.), pp.289-307.
Plenum Press, New York.
Ferrari, S. E 1988. The Behaviour and Ecology of the Buffy-
headed Marmoset, Callithrixflaviceps (0. Thomas, 1903).
PhD Thesis, University College, London.
Freese, C. H., and Oppenheimer, J. R. 1981. The capuchin
monkeys, genus Cebus. In: Ecology and Behavior of Neotro-
pical Primates, Vol. 1, A. F. Coimbra-Filho and R. A.
Mittermeier (eds.), pp.331-390. Academia Brasileira de
Ciencias, Rio de Janeiro.
Hubbell, S. P. and Foster, R. B. 1986. Commonness and rar-
ity in a neotropical forest: implications for tropical tree con-
servation. In: Conservation Biology: the Science of Scarcity
and Diversity, M. E. Soule (ed.), pp.205-231. Sinauer As-
sociates, Inc., Massachusetts.
Izar, P. 1999. Aspectos de ecologia e comportamento de um
grupo de macacos-pregos (Cebus apella) em area de Mata
Atlntica, Sao Paulo. Tese de Doutoramento, Instituto de
Psicologia, Universidade de SIo Paulo, Slo Paulo. 144pp.
Izawa, K. 1980. Social behavior of the wild black-capped ca-
puchin (Cebus apella). Primates 21(4): 443-467.
Izawa, K. 1988. Preliminary report on social changes of black-
capped capuchins (Cebus apella). Field Studies of New World
Monkeys, La Macarena, Colombia 1: 13-18.
Izawa, K. 1990. Social changes within a group of wild black-
capped capuchins (Cebus apella) in Colombia (II). Field
Studies of New World Monkeys, La Macarena, Colombia 3:
Izawa, K. 1992. Social changes within a group of wild black-
capped capuchins (Cebus apella) (III). Field Studies of New
World Monkeys, La Macarena, Colombia 7: 9-14.
Izawa, K. 1994a. Group division of wild black-capped capu-
chins. Field studies of New World Monkeys, La Macarena,
Colombia 9: 5-14.
Izawa, K. 1994b. Social changes within a group of wild black-
capped capuchins (IV). Field studies of New World Monkeys,
La Macarena, Colombia 9: 15-21.
Izawa, K 1997. Social changes within a group of wild black-
capped capuchins (V). Field studies of New World Monkeys,
La Macarena, Colombia 11: 1-10.
Janson, C. H. 1985. Ecological and social consequences of
food competition in brown capuchin monkeys. Doctoral
dissertation, University of Washington, Seattle.
Janson, C. H. 1996. Toward an experimental socioecology of
primates. Examples from Argentine brown capuchin mon-
keys (Cebus apella nigritus). In: Adaptive Radiations of Neo-
tropical Primates, M. A. Norconk, A. L. Rosenberger and P.
A. Garber (eds.), pp.309-325. Plenum Press, New York.
Lande, R. and Barrowclough, E 1987. Effective population
size, genetic variation, and their use in population manage-
ment. In Viable Populations for Conservation, M. E. Sould
(ed.), pp.87-124. Cambridge University Press, Cambridge.

Lopes, M. A. and Andrade, P. 1986. Inventirio das espdcies
vegetais da Fazenda Montes Claros. Relat6rio nIo publicado.
Departamento de Botinica, Universidade Federal de Minas
Gerais, Belo Horizonte.
Lynch, J. W. 1998. Mating behavior in wild tufted capuchins
(Cebus apella) in Brazil's Atlantic forest. Am. J. Phys.
Anthropol. suppl. 26: 153.
Lynch, J. W 1999a. Hierarquia social, coaliz6es, e formacIo
de subgrupos em macacos-prego (Cebus apella nigritus) de
Minas Gerais, Brasil. Resumos do IX Congresso Brasileiro de
Primatologia, Santa Teresa, Espirito Santo, p.43.
Lynch, J. W 1999b. Dominance, coalitions, and sub-group-
ing patterns of Brazilian tufted capuchin monkeys (Cebus
apella). Am. J Primatol. 49: 76-77.
Miller, L. E. 1996. The behavioral ecology of wedge-capped
capuchin monkeys (Cebus olivaceus). In: Adaptive Radia-
tions of Neotropical Primates, M. A., Norconk, A. L.
Rosenberger and P A. Garber (eds.), pp.271-287. Plenum
Press, New York.
Oppenheimer, J. R. 1982. Cebus capucinus: Home range,
population dynamics, and interspecific relationships. In: The
Ecology of a Tropical Forest: Seasonal Rhythms and Long- Term
Changes, E. G. Leigh, A. S. Rand and D. M. Windsor, D.
M (eds.), pp.253-270. Smithsonian Institution, Washing-
ton, DC.
Perry, S. E. 1995. Social relationships in wild white-faced ca-
puchin monkeys, Cebus capucinus. Doctoral dissertation,
University of Michigan, Ann Arbor.
Pinto, L. P. S. and Tavares, L. I. 1993. DistribuigIo e
conservaqlo dos primates do sul da Bahia. Unpublished re-
port, Universidade Federal de Minas Gerais, Belo Horizonte.
Phillips, K A. 1994. Resource distribution and sociality in
white-faced capuchins, Cebus capucinus. Doctoral disserta-
tion, University of Georgia, Athens.
Rimoli, J. and Ades, C. 1997. Estratdgia de forrageamento
de um grupo de muriquis (Brachyteles arachnoides, Primates,
Cebidae) da Estaqao Biol6gica de Caratinga-MG. In A
Primatologia no Brasil 5, S. E Ferrari and H. Schneider
(eds.), pp.39-57. Universidade Federal do Pari. Sociedade
Brasileira de Primatologia, Beldm.
Rfmoli, J. and Ferrari, S. E 1997. Comportamento e ecologia
de macacos-prego (Cebus apella nigritus, Goldfuss, 1809)
na EstacIo Biol6gica de Caratinga (MG). Program e
Resumos, VIII Congresso Brasileiro de Primatologia. Jolo
Pessoa, Paraiba, p.231.
Rfmoli, J. and Lynch, J. W. 1999. Demografia de um grupo
de macacos pregos (Cebus apella nigritus, Primates: Cebidae)
na Estagco Biol6gica de Caratinga, Minas Gerais. Resumos
do IX Congresso Brasileiro de Primatologia, Santa Teresa,
Espirito Santo, p.71.
Robinson, J. G. 1988a. Demography and group structure in
wedge-capped capuchin monkeys, Cebus olivaceus.
Behaviour, 104(3-4): 202-232.
Robinson, J. G. 1988b. Group size in wedge-capped capu-
chin monkeys Cebus olivaceus and the reproductive success
of males and females. Behav. Ecol. SociobioL 23: 187-189.

Neotropical Primates 8(1), March 2000 49

Robinson, J. G., and Janson. C. H. 1987. Capuchins, Squir-
rel Monkeys, and Atelines: Socioecological Convergence
with Old World Primates. In: Primate Societies, B. Smuts,
D. Cheney, R. Seyfarth, R. Wrangham and T. Struhsaker
(eds.), pp. 69-82. University of Chicago Press, Chicago.
Sanz, V. and Mirquez, L. 1994. Conservaci6n del mono
capuchino de Margarita (Cebus apella margaritae) en la Isla
de Margarita, Venezuela. Neotrop. Primates 2(2): 5-8.
Soini, P. 1986. A gynecological study of a primate commu-
nity in the Pacaya-Samiria National Reserve, Peru. Primate
Conserv. (7): 63-71.
Strier, K. B. 1987. Demographic patterns in one group of
muriquis. Primate Conserv. (8): 73-74.
Strier, K. B. 1991. Demography and conservation of an en-
dangered primate, Brachyteles arachnoides. Conserv. Biol.
5(2): 214-218.
Strier, K. B. 1992. Faces in the Forest: The Endangered Muriqui
Monkeys of Brazil. Oxford University Press, New York.
Strier, K. B., 1999a. Predicting primate responses to "stochas-
tic" demographic events. Primates. 40(1): 131-142.
Strier, K. B., 1999b. Primate Behavioral Ecology. Allyn and
Bacon, Boston.
Struhsaker, T. and Leland, L. 1987. Colobines: Infanticide
by Adult Males. In: Primate Societies, B. Smuts, D. Cheney,
R. Seyfarth, R. Wrangham and T. T. Struhsaker (eds.), pp.
83-97. The University of Chicago Press, Chicago.
Valderrama, X., Srikosamatara, S. and Robinson, J. G. 1990.
Infanticide in wedge-capped capuchin monkeys, Cebus
olivaceus. Folia Primatol. 54: 171-176.
Waser, P. M. 1987. Interactions among primate species. In:
Primate Societies, B. Smuts, D. Cheney, R. Seyfarth, R.
Wrangham and T. Struhsaker (eds.), pp.210-226. The
University of Chicago Press, Chicago.


A listing of the World's top 25 most endangered primates
was published in the 17 January 2000 edition of Time
magazine (pp.76-79). The list was prepared by Conservation
International, Washington, DC, in collaboration with the
IUCN/SSC Primate Specialist Group Chairman, Russell A
Mittermeier, and Deputy Chairmen, William R. Konstant
and Anthony B. Rylands, and released at a press conference
at the National Press Building, Washington, DC, on 10
January. The press release was also attended by Primate
Specialist Group members Devra G. Kleiman (National
Zoological Park, Washington, DC), Karen B. Strier (Univer-
sity of Wisconsin, Madison) and Thomas T. Struhsaker (Duke
University, Durham, North Carolina).

Of the 25 primates listed, 24 are endemic to seven of the 25
biodiversity hotspots identified for priority conservation action
by Conservation International (see Mittermeier et al., 1999;
Myers et al., 2000). The hotspots identified by Conservation

International cover only 1.4% of the Earth's land surface, but
claim more than 60% of all plant and animal diversity. The
mountain gorilla (Gorilla beringei) occurs in the Democratic
Republic of Congo, Rwanda and Uganda). The hotspots and
the critically endangered primate species occurring in them
are as follows:

Madagascar and Indian Ocean Islands: Golden bamboo
lemur (Hapalemur aureus), Lac Alaotra bamboo lemur (H.
griseus alaotrensis), Perrier's sifaka (Propithecus diadema
perrieri), the silky sifaka (P d. candidus), and the golden-
crowned sifaka (P tattersalli).
Atlantic forest region: Golden lion tamarin (Leontopithecus
rosalia), black lion tamarin (L. chrysopygus), black-faced lion
tamarin (L. caissara), the buff-headed capuchin (Cebus
xanthosternos), and the northern muriqui (Brachyteles
Tropical Andes: Yellow-tailed woolly monkey (Lagothrix
Guinean Forests of West Africa: Miss Waldron's red colobus
(Procolobus badius waldroni), white-naped mangabey
(Cercocebus atys lunulatus), Sclater's guenon (Cercopithecus
sclateri), the drill (Mandrillus leucophaeus), and the Cross River
gorilla (Gorilla gorilla diehli).
Eastern Arc Mountains and Coastal Forests of Tanzania and
Kenya: Sanje mangabey (Cercocebus galeritus sanjei)
Indo-Burma: Delacour's langur (Trachypithecus delacouri),
Cat Ba Island golden-headed langur (T poliocephalus), grey-
shanked douc langur (Pygathrix nemaeus cinerea), Tonkin
snub-nosed monkey (Rhinopithecus avunculus) and Hainan
gibbon (Hylobates concolor hainanus).
Sundaland: the Sumatran orangutan (Pongo abelii) and the
Javan gibbon (Hylobates moloch).

Russell A. Mittermeier, William R. Konstant and Anthony
B. Rylands, Conservation International, Suite 200, 2501 M
Street NW, Washington, DC 20037, USA.


Mittermeier, R. A., Myers, N., Gil, E R. and Mittermeier, C.
G. (eds.). 1999. Hotspots: Earth's Biologically Richest and
Most Endangered Terrestrial Ecoregions. Cemex, Conserva-
tion International and Agrupaci6n Sierra Madre,
Monterrey, Mexico.
Myers, N., Mittermeier, R. A., Mittermeier, C. G., Fonseca,
G. A. B. da and Kent, J. 2000. Biodiversity hotspots for
conservation priorities. Nature, Lond. 403: 853-858.


What exactly is the diversity of primates that we need to con-
serve? Are there primate populations which are being ignored
in worldwide conservation efforts just because they are not
currently recognized as distinct taxa? These were two of the
questions which stimulated the IUCN/SSC Primate Special-
ist Group (PSG) to hold a workshop, "Primate Taxonomy for

Neotropical Primates 8(1), March 2000

the New Millennium", held at the Disney Institute,
Orlando,Florida, 25-29 February 2000. The workshop was
organized by the PSG Chair Russell A. Mittermeier in col-
laboration with Don Melnick, Executive Director of the Cen-
ter for Environmental Research and Conservation., Colum-
bia University, NY, and John E Oates, Hunter College, City
University of NewYork, NY, and was sponsored by the Margot
Marsh Biodiversity Foundation, Virginia, and the Disney In-
stitute, Orlando, Florida, which kindly provided the venue
and excellent accommodation.

The Workshop involved the collective brainstorming of ap-
proximately 25 field primatologists, taxonomists, biogeogra-
phers, morphologists and geneticists in discussions of three
major questions: 1) the identification oftaxa 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; 2) the estab-
lishment of a single taxonomic listing for the primates, based
on the evidence available today, most particularly from mor-
phological and genetic research; and 3) the degree to which
the identifiable taxa are currently threatened, based upon the
most reliable information.

The participants were divided into working groups dealing
with each of the four major primate regions: the Neotropics,
Africa, Madagascar and South-east Asia. Those who contrib-
uted to the group for the Neotropical primates included:
Russell A. Mittermeier (Conservation International, Wash-
ington, DC), Colin P. Groves (Australian National Univer-
sity, Canberra), Horacio Schneider (Universidade Federal do
Pari, Belkm, Brazil), Ernesto Rodrfguez-Luna (Universidad
Veracruzana, Xalapa, Mexico), Alfredo Langguth
(Universidade Federal do Paraiba, Joao Pessoa, Brazil), Peter
Grubb (London, UK), and Anthony B. Rylands, (Center for
Applied Biodiversity Science, Conservation International,
Washington, DC). Particularly important was the contribu-
tion of Colin Groves, whose book on primate taxonomy is
soon to be published by the Smithsonian Institution Press,
Washington, D.C.

Participants in other groups included: Simon Bearder (Noc-
turnal Primate Research Group, Oxford Brookes University,
Oxford, UK), Douglas Brandon-Jones (Natural History Mu-
seum, London), Thomas M. Butynski (PSG Vice Chair for
Africa, Africa Biodiversity Conservation Program, Zoo At-
lanta, Nairobi), Todd R. Disotell (New York University, New
York), Ardith Eudey (PSG, Vice Chair for Asia, Upland, Cali-
fornia), Jbrg Ganzhorn, (PSG Vice Chair for Madagascar,
Hamburg University, Hamburg), Thomas Geissmann
(Institut fuir Zoologie, Tierirztliche Hochschule, Hannover),
Kenneth Glander (Duke Primate Research Center, Durham,
North Carolina), William R. Konstant (PSG Deputy Chair,
Conservation International, Washington, DC), Don Melnick
(Executive Director of the Center for Environmental Research
and Conservation), Juan Carlos Morales (Center for Envi-
ronmental Research and Conservation, Columbia University,
New York), Myron Shekelle (Washington University, Saint

Louis, Missouri), Caro-Beth Stewart (University of Albany
SUNY, New York), Thomas T. Struhsaker (Duke University,
Durham, North Carolina) and Ian Tattersall (American Mu-
seum of Natural History, New York).

The results of this workshop are still being compiled, and will
form the basis for an Action Plan for Critically Endangered
and Endangered Primates, currently being prepared by the
PSG and coordinated by William R. Konstant. The full, an-
notated taxonomic listing for the platyrrhines which resulted
from the workshop will be published in the next issue of Neo-
tropical Primates 8(2).

Ernesto Rodriguez-Luna, Director, Instituto de
Neuroetologfa, Universidad Veracruzana, AP 566, Xalapa,
Veracruz, 91000 Mexico, and Anthony B. Rylands, Senior
Director, Conservation Biology, Center for Applied
Biodiversity Science, Conservation International, 2501 M
Street, NW, Suite 200, Washington, DC 20037, USA.


Groves, C. P. In press. The Taxonomy ofPrimates. Smithsonian
Institution Press,Washington, DC.


Carla Castro defended her doctoral thesis, entitled "Diet,
Patterns of Range Use, and Agonistic Intergroup Interactions
of the Common Marmoset (Callithrix jacchus)", for the
postgraduate program in Ecology and Natural Resources at
the Federal University of Sao Carlos (UFSCar), Sio Paulo,
Brazil. Her supervisors were Professor Manoel Martins Dias
Filho and Professor Cleber Jos6 Rodrigues Alho. The Brazilian
Higher Education Authority (CAPES) supported the study.
The following is an abstract of the thesis.

The diet, range use, and agonistic interactions of three groups
of common marmosets (Callithrixjacchus) were studied from
October 1996 to January 1998, at the Experimental Forestry
Station (060 05'S, 350 12' W) EFLEX/IBAMA, Rio Grande
do Norte, Brazil. The study involved 728 hours of direct
observation of the groups. The phenology of 231 trees was
monitored. During the course of the study, the marmosets
used all of the 231 trees (16 species, eight families) for their
fruits and/or gums. Gum feeding was negatively related to
the availability of fruits, but not to animal prey ingestion.
Invertebrates such as grasshopers (Orthoptera: Acrididae and
Tettigoniidae), crickets (Gryllidae), Coleoptera
(Chrysomelidae), Lepidoptera, cicadas (Cicadidae), and
occasionally lizards and nesting birds comprised the majority
of animal prey items taken. Although gum provide a
significant part of their diet, fruits were preferred when
available. Home-range size varied from 0.7 to 2.4 ha. Fruit
and gum trees were patchy in their distribution, affecting

NeotropicalPrimates 8(1), March 2000 51

patterns of range use. Two kinds of agonistic interactions were
observed: a) intergroup interactions, and b) interactions
between groups and intruders. Agonistic intergroup
interactions were more frequent (n = 278) than those between
groups and intruders (n = 26). Adult and subadult marmosets
of both sexes participated in these interactions, but juvenile
individuals were rarely involved. Agonistic behaviors shown
at these interactions included vocalizations, arch-bristle
movements, general piloerection, chases, and tail-raised
present. The frequency of intergroup interactions did not
change significantly between dry and wet months, suggesting
that defense of food sources was not the primary function of
agonistic intergroup interactions. However, breeding
marmosets presented higher levels of agonistic behaviors
during intergroup interactions when individuals of the same
sex were present, indicating mate defense. In contrast, non-
breeding marmosets showed affiliative behaviors such as
grooming during interactions between groups and intruders.
Non-breeding marmosets also presented high rates of extra-
group copulations following or during an agonistic
intergroup interaction. Two males from strange groups
emigrated into one of the study groups. Before the
immigrations at least one of them had been observed
copulating with two non-breeding females from the study
group during agonistic intergroup interactions. At the end
of the observation period the two females and one of the males
that emigrated were still residing in the study group and
became the breeding individuals. This reinforces the idea that
intergroup interactions play a role in locatng mates and
breeding opportunities outside of their group.

Carla Castro, Departamento de Zoologia, Universidade Fe-
deral do Rio Grande do Norte (UFRN), Centro de
Biociencias, BR 101, Lagoa Nova, Natal 59072-070, Rio
Grande do Norte, Brazil.


Paulo Auricchio defended his master's thesis "Relac6es
Filogen6ticas de Pithecia (Atelidae)" on 4 December 1999,
at the Department of Zoology, University of Sio Paulo. His
supervisor was Mirio C. C. de Pinna, and the study was
supported by the Brazil Science Council (CNPq). The
following is an abstract of the thesis.

Uma analise cladfstica entire 6 das 8 espdcies e subespdcies
recentes do genero Pithecia (Primates, Atelidae) foi feita com
base em caracteres de pelagem, palo, crinio, de p6s-crinio e
cari6tipo. P aequatorialis e P. monachus miller nio foram
induidos na analise devido a falta de material para estudo.
Sessenta e seis caracteres foram analisados usando-se o
program Hennig86 (versdo 1,5) para implementar andlise
de parsim6nia estrita. Os caracteres foram polarizados por
comparaq6es com os grupos-externos, representados por
especies dos generos Cebus (macaco-prego), Chiropotes (cuxid),
e Cacajao (uacari). fndices de Consistencia (CI) e de Retencao
(RI) tambem foram calculados para cada carter e para o

cladograma total. Duas irvores mais parsimoniosas (L = 110,
CI = 59, RI= 55) foram encontradas. 0 consenso (L = 113,
CI = 57 e RI = 52) indica a seguinte hip6tese: (Chiropotes)
(Cacajao) (P.p. pithecia, P.p. chrysocephala) (P albicans) (P m.
monachus, P i. irrorata, P i. vanzolinii). Os resultados desta
analise evidenciam o monofiletismo de Pithecia e que Cacajao
Sseu grupo-irmao. Os tAxons setentrionais ao Rio Amazonas,
P.p. pithecia e Pp. chrysocephala, formam grupo monofiltico
que e o grupo-irmao de todo o restante do genero. Duas outras
andlises experimentais foram efetuadas separadamente
incluindo caracteres de pelagem e de p&lo para avaliar a
quantidade de informac(o filogen&tica em cada um destes
grupos de caracteres. A relagqo entire P i. irrorata, P i.
vanzolinii e P m. monachus nao estd resolvida corn base no
conjunto total de caracteres, mas aparecem plenamente
resolvidos na andlise de palo: (Pithecia i. irrorata) (P m.
monachus) (P i. vanzolinii).

Phylogenetic relations of Pithecia (Atelidae)

The study involved a cladistic analysis of six of the eight recent
taxa of the genus Pithecia (Primates, Atelidae). It was based
on the following characters: pelage, hair, cranium,
postcranium, and karyotype. P aequatorialis and P monachus
milleriwere not included in the analysis due to lack of material.
Sixty-six characters were analysed by strict parsimony with
the help of the program Hennig86 (version 1.5). Characters
were polarized by outgroup comparisons, represented by
species of the genera Cebus (capuchin monkey), Chiropotes
(bearded saki monkey), and Cacajao uakarii). Consistency and
retention indices were calculated for each character and the
whole cladogram. Two most parsimonious trees were found
(L = 110, CI = 59, RI = 55). Their consensus (L = 113, CI =
57 and RI = 52) indicates the following hypothesis
(Chiropotes) (Cacajao) (P p. pithecia, P p. chrysocephala) (P
albicans) (P m. monachus, P i. irrorata, P i. vanzolinii). The
results of the analysis indicate monophyly of Pithecia, and
that Cacajao is its sister group. Taxa to the north of the Amazon
basin (P. p. pithecia, P. p. chrysocephala), form a monophyletic
group, which is the sister group to the rest of the genus. Two
other experimental analyses were performed using fur and
hair characters separately to evaluate the amount of
phylogenetic information in each subset of the data base. The
relationships between P i. irrorata, P i. vanzolinii and P m.
monachus were unresolved on the basis of the present total
data set, but were completely resolved in the hair analysis
(Pithecia i. irrorata) (P m. monachus) (P. i. vanzolinii).

Paulo Auricchio, Instituto Pau Brasil de Hist6ria Natural,
Caixa Postal 282, 07400-970 Aruji, Sio Paulo, Brazil. E-mail:


Auricchio, P. 1999. Rela6es Filogeneticas de Pithecia (Pri-
mates, Atelidae). Master's thesis, Departamento de Zoologia,
Universidade de Sio Paulo, Slo Paulo. December 1999, 92pp.

Neotropical Primates 8(1), March 2000


Jtilio C&sar Bicca-Marques defended his PhD thesis entitled
"Cognitive aspects of within-patch foraging decisions in wild
diurnal and nocturnal New World monkeys" at the
Department of Anthropology of the University of Illinois at
Urbana-Champaign, USA, on 29 October 1999. The study
was supervised by Dr. Paul A. Garber. Financial support was
provided by the Brazilian Higher Education Authority
(CAPES/MEC) World Wildlife Fund-Brazil, Fundacao 0
Boticirio de Protegio a Natureza, Wenner-Gren Foundation
for Anthropological Research, the American Society of
Primatologists, Department of Anthropology and the Tinker
Fund/Center for Latin American and Caribbean Studies/
University of Illinois at Urbana-Champaign. Logistical support
was provided by the Zoobotanical Park/Federal University of
Acre and Fundacio S.O.S. Amaz6nia. The following is an
abstract of the thesis.

In this dissertation, I examine the ability of free-ranging
diurnal and nocturnal New World monkeys (black-chinned
emperor tamarins, Saguinus imperator imperator, Weddell's
saddleback tamarins, Saguinus fuscicollis weddelli; red titi
monkeys, Callicebus cupreus cupreus; and southern red-necked
night monkeys, Aotus nigriceps) to use visual cues, olfactory
cues, spatial information, associative cues, landmark cues, and
quantitative information in making within-patch foraging
decisions. This was accomplished through a controlled
experimental field study conducted at the Zoobotanical Park
of the Federal University of Acre (Rio Branco, Acre, Brazil)
from August 1997 to July 1998.

Analyses at the group level indicated that all four species were
capable of learning the spatial distribution of food items within
a patch and used this knowledge to return to previously
exploited feeding sites. Diurnal monkeys used visual cues to a
greater extent than did night monkeys. Night monkeys,
however, failed to show evidence of the use of olfactory cues
more effectively than did emperor tamarins. In addition, when
faced with conflicting spatial and perceptual information, all
four species relied on visual or olfactory cues in selecting
feeding sites. Results on diurnal and nocturnal species suggest
that one of the main challenges faced by early primates in
their shift from a nocturnal to a diurnal lifestyle was taking
advantage of available visual cues (e.g., hue) in locating feeding

A comparison of the performances of emperor and saddleback
tamarins when in and out of mixed-species association
indicated that the dominant emperor tamarins may use
information from the foraging behavior of saddlebacks to
improve their searching efficiency. In contrast, saddlebacks
performed better when out of association. Moreover, when

forming mixed-species troops, both emperor and saddleback
tamarins experienced costs of decreased time spent feeding.

Analyses at the individual level indicated that among emperor
and saddleback tamarins, group members may be classified
as producers, scroungers, or opportunists, depending on the
time invested searching for food. Producers focused on
ecological information in making foraging decisions, whereas
scroungers used primarily social information. Opportunists,
on the other hand, balanced the use of both kinds of
information. The adoption of these strategies is likely to be
determined by factors such as social status and age. Differences
in cognitive skills are unlikely to have an important role in
the individual choice of foraging strategies. In addition,
saddlebacks were more tolerant than emperors of sharing food
rewards with other group members. Only adult male emperor
tamarins, however, deferred to immatures and breeding adult
females at feeding sites.

The results of the present study suggest that a model of primate
cognition that integrates both social and ecological
information provides the most appropriate framework for
addressing questions concerning the evolution of primate
intelligence and decision-making, and that this issue can be
studied experimentally in the wild by manipulating food
availability and distribution, and the presence of potential

Jfilio Cesar Bicca-Marques, Department of Anthropology,
University of Illinois at Urbana-Champaign, 109 Davenport
Hall, 607 South Mathews Avenue, Urbana, IL 61801, USA.
E-mail: .


Norka E Rocha S. defendi6 en abril 1999 su Licenciatura en
Biologia de la Universidad Gabriel Rene Moreno de Santa
Cruz, con la tesis "Censo de Primates y Evaluaci6n de la
Metodologfa de Lineas de Transectas en Lago Caiman, Parque
Nacional Noel Kempff Mercado". Fue supervisada por el Dr.
Robert Wallace, quien realize un studio de 2 afios en el mismo
sitio (Wallace, 1998) y por el Dr. Damin Rumiz, especialista
en Vida Silvestre del Proyecto de Manejo Forestal Sostenible
BOLFOR y de Wildlife Conservation Society. El studio fue
financiado con fondos de USAID y el Gobierno de Bolivia.
A continuaci6n se present el resume de la tesis.

Se estim6 en base a censos por transectas lineales la abundancia
y densidad de Ateles chamek y Cebus apella en la parcela de
investigaci6n del "Lago Caimin", Parque Nacional Noel
Kempff Mercado, en el Noreste del Departamento de Santa
Cruz, Bolivia. La parcela de investigaci6n contaba con mis
de 440 ha y un sistema de sendas que la cuadriculaban cada
100 m. Inclufa cuatro tipos de habitats; bosque alto, bosque
bajo de lianas, bosque sartenejal estacionalmente innundable
y bosque pie de monte en la base de la serrania de Huanchaca.

Neotropical Primates 8(1), March 2000 53

Se realizaron censos diurnos caminando un total de 289.51
km en una series de 6 replicaciones, donde se registraron
encuentros con primates tomando en cuenta la hora, especie,
ntImero, composici6n y comportamiento del grupo, distancia
ya recorrida a lo largo de la transecta, metodo de detecci6n,
distancia perpendicular desde la transecta al centro del grupo,
habitat general, calidad de la observaci6n, condiciones del
tiempo, hora inicial y hora final del recorrido. Los analisis de
densidad se realizaron por medio del program Distance y la
abundancia fue expresada el nmiero de encuentros por 10
kmin de recorrido. La densidad de Ateles chamek se estim6
utilizando la distancia perpendicular al centro del grupo y a
cada individuo por separado, y se compare con el valor real
de densidad de la misma poblaci6n conocida segun studios
de largo plazo obtenidos a trav6s de radiotelemetria y
seguimiento de grupos (Wallace, 1998).

En las parcelas de investigaci6n del Lago Caimin se
encontraron tres species de primates diurnos; con mis
frecuencia el marimono (Ateles chamek), seguido por el mono
martin (Cebus apella) y raramente el mono leoncito (Callithrix
melanura). La densidad estimada para Ateles chamek usando
las distancias al centro del grupo (n = 84 encuentros) fue 6.18
grupos/km2 o 18 individuos/km2, que fue igual (18
individuos/km2) a lo estimado usando las distancias
individuals (n = 245). Los valores obtenidos por ambos
m6todos fueron muy similares de 18 a 20 individuos/km2
calculada segfin el registro de 45 a 50 individuos en un total
de 250 ha (Wallace, 1998).

La densidad de Cebus apella en base a 29 encuentros fue de
3.17 grupos/km2 y 11.4 individuos/km2. La abundancia se
estim6 con nmmero de encuentros de grupo por cada 10 km
de recorrido y fue mayor para Ateles chamek (x = 2.9/10 kin,
DS. 0.89) que para Cebusapella (x = 0.99/10 km, DS. 0.40)
usando las seis replicas de trayecto. En base al analysis de chi
cuadrado se observaron preferencias en cuanto al uso de los
habitats entire las diferentes species y se comprob6 que Ateles
chamek seleccion6 los habitats de bosque alto, sartenejal y pie
de monte mientras que Cebus apella no mostr6 preferencia o
aversion por algtin habitat. Los patrons de uso de habitat de
Ateles y Cebus fueron diferentes y variaron estacionalmente.

La linea de transectas es un metodo adecuado para realizar
estimaciones de abundancia y densidad, donde otros m&todos
son impracticables. Adennis esta metodologia require de poca
inversion y de poco equipo, pero de considerable esfuerzo
segin la precision deseada. Las estimaciones de densidad de
Ateles chamek en base a las distancias al centro de grupo y a
cada individuo fueron muy cercanas a la densidad real, por
lo que de ambas variantes pueden recomendarse la distancia
al centro del grupo porque es menos laboriosa. Este metodo
de transectas lineales sirve para realizar censos de primates y
puede ser aplicado para establecer programs de monitoreo
en zonas donde se permiten actividades extractivas.

Norka Rocha, Museo de Historia Natural, Noel Kempff
Mercado, Area de Mastozoologia, Casilla 2489, Proyecto
Bolfor, Santa Cruz, Bolivia, e-mail: < museo @museo .sczbo .com>.


Rocha, N.E 1999. Censo de Primates y Evaluaci6n de la
Metodologia de Lineas de Transectas en Lago Caimin -
Parque Nacional Noel Kempff Mercado. Tesis Licenciatura,
Facultad de Biologia, Universidad Aut6noma Gabriel Rene
Moreno, Santa Cruz, Bolivia.
Wallace, R. 1998. The Behavioural Ecology of Black Spider
Monkeys in North-eastern Bolivia, Tesis de PhD, University
of Liverpool, Liverpool.


Promoting the development of scientific research in the areas
of biology and conservation in the Central American
Isthmus, the Central American Institute for Biological
Research and Conservation is offering short field training
courses. Undergraduate and graduate courses on Biology and
Conservation subjects include: Field ecology, 15-31 April,
2001 Application deadline January 30, 2001, and
Phylogenetics cladisticc analysis), 21 January- 4 Febuary, 2001,
application deadline, November 24, 2000. Further
information can be found at:
or email .


Botanist Alwyn Gentry died tragically in a plane crash on 3
August 1993 in western Ecuador. The Missouri Botanical
Garden is, however, continuing his project on the study of
the floristic diversity of the world's tropical forests. Gentry
and his collaborators had surveyed nearly 250 sites on six
continents, establishing and collecting data from 0.1 ha
transects. A review of these studies has been compiled by James
Miller, Oliver Phillips, and Nancy Hediger, and the raw data
is available on the Garden's web site: MOBOT/research/applied_research/gentry.html>. The data
for each site are being analyzed, and a volume summarizing
the results will be published by the MBG. In addition to
summarizing the transect data, the book will review the
historical development of Gentry's ecological studies, the
methods by which the date were collected, and their
significance in contributing to our understanding of global
patterns of plant diversity. Missouri Botanical Garden, Tel:
314 577 5169, Fax: 314 577 0830. From: Tropinet, 10(3),
September 1999.


A Primate Mixed Species Symposium was held outside Dallas,
Texas on February 27, 1999. More than 80 zoo professionals
from approximately 45 institutions attended to participate in

54 Neotropical Primates 8(1), March 2000

presentations of 11 papers and poster sessions followed by an
open roundtable discussion of the success and problems of
mixed primate species combinations and exhibits. The Dallas
Zoo has published the Proceedings. This includes the
roundtable discussion, which was edited and categorized as
follows: How can success of a mixed-species exhibit be assessed?
General advice for mixed-species exhibits: Record keeping,
Management, Exhibit Mixed species successes and failures:
New World Primates, Old World Primates, Prosimians, Non-
primate mixed species combinations.

The Proceedings are available to anyone interested, for a cost
of $15.00, which will be donated to the New World Primate
Taxon Advisory Group Conservation Fund. Requests from
primate habitat countries will be processed free. Please make
check payable to the Dallas Zoological Society. Send your
request to: Ken Kaemmerer, Dallas Zoo, 650 South R.L.
Thornton Freeway, Dallas, Texas 75203, USA.

The Canopy Citations Database is now available on the World
Wide Web. It contains over 1,300 citations regarding canopy
ecology. Search for authors, titles, dates, journals, keywords
or words within an abstract. Web site: canopycitations>.


The Primate Information Center Primate Literature Database,
PrimateLit, indexing over 140,000 research publications from
1940 to the present is now available on the web. Access to
PrimateLit is by password only. In order to obtain your
password contact the Primate Information Center at: email:
, subject line: Password Request
Message: Include your full name, affiliation and email address.
A password and instructions for accessing the database will
be e-mailed to you.


Eric Bairrdo Ruivo, EEP Co-ordinator for the Emperor
Tamarin, Saguinus imperator, published the 1999 Studbook
(6" edition, data current to 31" December 1999). He was
assisted by Prof. Pereira da Silva, Patricia Vilarinho, Orlando
Silva and Jose Dias Ferreira. The Studbook includes the 1998
Annual report, and a report of a research project, entitled,
"Hybrids: Relationships between Emperor Tamarins" by
Anne-Sophie Blandin. The Studbook includes a full historical
listing of S. i. imperator and S. i. subgrisescens and hybrids,
births, deaths and transfers during 1999, a listing of the living
population by location, a comprehensive demographic and
genetic analysis (including age pyramids, analyses of age

distributions, mortality, fecundity, and inbreeding),
recommendations regarding breeding and transfers for 2000,
and an address list of holding institutions.

The full listing includes data on 730 tamarins (608 S. i.
subgrisescens, 47 S. imperator, 75 hydrids), 145 of which were
alive on 31 December, 1999. The current population in
Europe comprises one pure S. i. imperator, a very old female
in the Frankfurt Zoo, and 131 S. i. subgrisescens (69:54:8, sex
ratio 1:0.79) in 42 institutions. The hybrid population has
been transferred to Peaugres with subsequent sterilisation of
all hybrid females.

The S. i. subgrisescens population arose from 34 founders, 29
wildborn and 5 animals of unknown origin. The population
has been increasing since 1980, however there was little
population growth in 1999 due to the deaths of several
important breeding females and the loss of two genomes from
the death of a founder without descendant and the death of
the only living descendant of another founder. The number
of institutions keeping emperor tamarins has increased with
many other institutions waiting to receive animals. Three new
institutions, Lille in France, Stuttgart in Germany and Zagreb
in Croatia, received tamarins for the first time and Jersey will
receive a group sometime during 2000. The goal for 2000
continues from 1999 with the hopes of publishing husbandry
guidelines as part of the Callithrichids EEP Primate TAG
Husbandry guidelines.

Eric Bairrio Ruivo, Jardim Zool6gico e de Aclimatagko em
Portugal, SA., Estrada de Benfica 158-160, 1549-004 Lisboa,
Portugal, e-mail: .


The editors of the Laboratory Primates Newsletter, published
quarterly by the Schrier Research Laboratory, Psychology De-
partment, Brown University, Rhode Island, USA, offer space
to all Spanish- and Portuguese-speaking primatologists to
publish in their native language announcements, news,
plans and summaries of the results of their investigations in
the section "Primates of the Americas the Page" (see Lab.
Prim. Newsl. 36[4]: 20, 1997). The norms for preparation of
summaries, notes and announcements are the same as for the
English language material, but contributions must be brief,
preferably not more than half a page. Please send all
correspondence related to "Primates of the Americas the
Page" to: Juan Carlos Serio Silva, Department of Plant Ecology,
Institute de Ecologia, A. C., A.P 63, CP 91000, Xalapa,
Veracruz, Mexico, Tel: 52 28 42 1800 x1201; Fax: 52 28 42
1800 x 1204, e-mail: < serioju@ sun. ieco.conacyt.mx>, or T.
Elva Mathiesen, c/o Judith Schrier, Psychology Department,
Box 1853, Brown University, Providence, RI 02912, USA,
Tel: 401 863 2511, Fax: 401 863 1300, e-mail:

Neotropical Primates 8(1), March 2000


An extensive website is now available on gibbons
(Hylobatidae). Thomas Geissman offers an an informative
introduction to the natural history of the lesser apes, a review
of gibbon systematics and gibbon singing behavior, a photo
and sound gallery of all Hylobatidae species, and news and
complete research articles on gibbon biology. The site can be
viewed at .


Mandated to promote high quality research on the manage-
ment, use, and conservation of biological resources and their
enviornment, the International Foundation for Science pro-
vides small research grants to scientists in and from a devel-
oping country or those employed in a developing country
institution. For further information contact: International
Foundation for Science, Grev Turegatan 19, 114 38
Stockholm, Sweden, Tel: (46) 8 545 818 00, e-mail:


The Fundacgo Biodiversitas, President Aspisia Camargo,
Director Luiz Carlos Cardoso Vale, took over the coordination
of the Brazilian Committee of the World Conservation Union
(IUCN) in February 2000. This decision was ratified during
the last meeting of the Committee, 13-14 April 2000, at the
Salto Morato Natural Reserve in Parana.

The FundacIo has changed its address. The new address of
the Fundacgo Biodiversitas is: Rua Ludgero Dolabela 1012,
7. Andar, 30430-130 Belo Horizonte, Minas Gerais, Brasil,
Tel: (0)31 292 8235, Fax: (0)31 291 7658, e-mail:
. Home page: biodiversitas.org>.


The IUCN/SSC Re-Introduction Specialist Group office has
moved from Nairobi, Kenya to Abu Dhabi, United Arab
Emirates with chairman Dr. Mark Stanley-Price being re-
placed by Dr. Frederic Launay, head of the National Avian
Research Center, Environmental Research and Wildlife De-
velopment Agency.

The Specialist Group is currently developing a project on
drawing up re-introduction guidelines for primates. If you

have publications or opinions regarding such a document,
please get in touch with Pritpal Soorae at the address below.

The new contact details for the Re-Introduction group are:
Pritpal S. Soorae, Senior Conservation Officer, IUCN/SSC
Re-Introduction Specialist Group, Environmental Research
and Wildlife Development Agency, P.O. Box 45553, Abu
Dhabi, United Arab Emirates. Tel: (D/L) 971 2 693 4506
or 693 4628. Fax: 971 2 693 4628.


The Primate Society of Great Britain (PSGB) awards small
grants in support of primate conservation and education.
These grants are administered by the Conservation Working
Party, which considers applications at its biannual meetings.
The following notes give details of eligibility and application
procedure and should be read carefully before preparing an

Proposals are invited for grants to assist-: research of benefit to
primate conservation; short surveys to identify locations of
value to primate conservation; projects involving conservation
education relevant to primates.

Obligations ofgrantees are as follows: to present a report on the
progress of the project within 6 months of commencement;
to present a final report on completion of the project to be
used by PSGB at its discretion in publications or in any way
thought to be of value to primate conservation; to
acknowledge the support received from PSGB in any
publication resulting from the project and to supply PSGB
with two copies of each publication; to produce, where
appropriate, slides and/or sound recordings for non-
commercial use by PSGB or others in the promotion of
primate conservation.

Eligibility. Grants will be awarded to members of PSGB or to
citizens of primate range states who are sponsored by a
member. Only those projects which are judged to have
attainable goals that will benefit primate conservation or
conservation education will be considered. Group training
projects will not be considered for these grants. Awards are
made on a competitive basis and the decision of the
Conservation Working Party is final. In some cases applicants
may be invited to submit an amended application.

Application and award details. Individual awards tend to be
in the range of 250 to 500. Two closing dates apply: the
last day of February and the last day of August. Applications
must be made on the Application Form, or following the same
format, and should be sent by post to reach the Convener on

56 Neotropical Primates 8(1), March 2000

or before the relevant closing date. Applications by e-mail or
fax will not normally be accepted.

Application forms are available on the PSGB web site
can be obtained directly from: David A. Hill (Convenor of
the CWP), School of Biological Sciences, University of Sus-
sex, Falmer, Brighton BN1 9QG, UK, Tel: +44 1273 606755
ext. 2755;Fax: +441273678433, e-mail: .


The new official journal of the Instituto Pau Brasil de Hist6ria
Natural (IPBHN), Director Dr. Paulo Auricchio, focuses on
various themes in natural history, including Biology, Zoology,
Botany, and Ecology and gives special emphasis to
Environmental Education. Manuscripts can be sent in
Portuguese, Spanish and English. The main objective of the
IPBHN is to divulge scientific studies as the first step to
understanding the world we live in and promote the
preservation of its biodiversity and natural ecosystems. For
subscriptions to Publicaroes Avulsas (three issues are already
available) please write to < or bulletin@institutopaubrasil.org.br>
or write to IPBHN (Revista), c/o Ana Maria de Souza (Edi-
tor), Caixa Postal 282, Aruji 07400-970, Sao Paulo, Brazil.


Blackwell Science has launched a new magazine -
Conservation Biology in Practice. It is designed for people who
are short on time but long on information needs. It includes
the following sections: Reviews; Features (new information
and thought provoking concepts and practices); Case studies
(successes, failures and lessons to be learned, new approaches
and strategies); Tools and Techniques; Numbers in Context
(graphs, charts and tables and other data with minimal text
to give readers a quick accessible way to graps conservation
trends); and Resources (book reviews, web sites and other
sources of practical conservation information). The editors
welcome articles that reflect clear and innovative thinking,
ideas that can be translated into management action, and
outstanding, jargon-free writing. Manuscript outlines
suggestionsn) and texts can be sent to: Conservation Biology
in Practice, Department of Zoology, Box 351800, University
of Washington, Seattle, WA 98195-1800, USA, Tel: 206 685-
4724; Fax: 206 221-7839, e-mail: .

It is published quarterly, and subscription rates are as follows:
US$30 in the USA, US$35 for Canada and Mexico, and
US$40 overseas. Institution rates: US$75 in the USA, US$80
for Canada and Mexico, and US$85 overseas. Write to:
Journal Subscription Department, Blackwell Science, Inc.,
350 Main Street, Maiden, MA 02148, USA.


Primate Communities, edited by J. G. Fleagle, C. H. Janson,
and K. E. Reed, 1999, 329pp. Cambridge University Press,
UK ISBN 0-521-62967-5. Price: 18.95. Comparing the
communities of living non-human primates throughout the
world, Primate Communities examines the underlying
similarities and differences between the composition,
behaviour and ecology of primate communities in Africa, Asia,
Madagascar and South America. Focusing on the phylogenetic
history, climate, rainfall, soil type, forest type, competition
with other vertebrates, and human activities as it relates to
primate communities, the text is an excellent source for
researchers and students of anthropology, ecology and
conservation. Contents: African primate communities:
Determinants of structure and threats to survival C. A.
Chapman, A. Gautier-Hion, J. E Oates & D. A. Onderdonk,
pp.1-37; Biomass and use of resources in south and south-
east Asian primate communities A. K. Gupta & D. J. Chivers,
pp.38-54; Species coexistence, distribution and
environmental determinants of neotropical primate richness:
A community-level zoogeographic analysis C. A. Peres & C.
H. Janson, pp.55-74; Primate communities: Madagascar J.
U. Ganzhorn, P. C. Wright & J. Ratsimbazafy, pp.75-89;
Primate diversity J. G. Fleagle, C. H. Janson and K. E. Reed,
pp.90-91; Phylogenetic and temporal perspectives on primate
ecology J. G. Fleagle & K. E. Reed, pp.92-115; Population
density of primates in communities: Differences in
community structure K E. Reed, pp.116-140; Body mass,
competition and the structure of primate communities J.
U. Ganzhorn, pp.141-157, Convergence and divergence in
primate social systems P M. Kappeler, pp.158-170; Of mice
and monkeys: Primates as predictors of mammal community
richness L. H. Emmons, pp.171-188; Comparing
communities J. G. Fleagle, C. H. Janson and K.E. Reed,
pp. 189-190; Large-scale patterns of species richness and
species range size in anthropoid primates H. A. C. Eeley &
M. J. Lawes, pp.191-219; The recent evolutionary past of
primate communities: Likely environmental impacts during
the past three millennia C. Turtin & L. White, pp.220-236;
Resources and primate community structure C. H. Janson
and C. A. Chapman, pp.237-267; Effects of subsistence
hunting and forest types on the stucture ofAmazonian primate
communities C. A. Peres, pp.268-283; Spatial and temporal
scales in primate community structure J. G. Fleagle, C. H.
Janson & K. E. Reed, pp.284-288; Primate communities in
Africa: The consequences of long-term evolution or the
artifact of recent hunting? T. T. Struhsaker, pp.289-294;
The future of primate communities: A reflection of the

Neotropical Primates 8(1), March 2000 57

present? P. C. Wright &J. Jernvall, pp.295-309; Concluding
remarks J. G. Fleagle, C. H. Janson & K.E. Reed, pp.310-
314. Available from: Cambridge University Press, The
Edinburgh Building, Cambridge CB2 2RU, UK, or 40 West
20th Street, New York, NY 10011-4211, USA Web site:

Primate Encounters Models of Science, Gender and Society,
edited by Shirley C. Strum and Linda M. Fedigan, 2000,
The University of Chicago Press. ISBN 0 226 77754 5. Price
US$35. An attempt to answer the diversity of primatologies
and on science in general by reflecting on the relations of
both to cultural, historical and social issues. Sections:
Introduction and History, What do the Pioneers say? The
Advantages of Hindsight, A diversity of other Primatologies:
Other National Traditions, Enlarging the Lens: Closely
Related Disciplines, Models of Science and Society,
Reformulating the Questions, Index. Includes chapters by
Maria Emilia Yamamoto and Anuska Irene Alencar (Some
Characteristics of Scientific Literature in Brazilian Primatology)
and Karen B. Strier (An American Primatologist Abroad in
Brazil). Available from: The University of Chicago Press, 11030
S. Langley Avenue, Chicago, IL 60628, USA, Tel: (800) 621-
2736, Fax: (800) 621-8476. e-mail: .
Website: http://www. press. uchicago.edu.

Lista Vermelha das Espicies Ameafadas de Extinfdo da Flora de
Minas Gerais, edited by Fundaclo Biodiversitas, Belo Hori-
zonte, June 2000. Fundacao Biodiversitas and the Fundacqo
Zoobotdnica de Belo Horizonte, Belo Horizonte. A list e uma
ferramenta essencial para o planejamento e execucgo de ati-
vidades relacionadas a conservaqao da flora mineira, alem de
alerta para a necessidade da atuacdo eferiva das entidades
responsiveis pela conservaqio dessas especies" explica Livia
Lins, bi6loga e coordenadora de projetos da Biodiversitas. Das
537 especies listadas, 221 (41%) sdo consideradas vulneriveis,
129 (24%) em perigo, 108 (20%) estio criticamente em
perigo e 79 (15%) foram consideradas provavelmente extintas
por nao terem sido encontradas na natureza nos liltimos 30
anos. A destruigio da vegetacao native e a coleta predat6ria,
principalmente para fins ornamentais, sdo as principals causes
da degradagao da flora mineira. Quase todas as 27 especies
de bromdlias, 31 de cactAceas e 20 de orqufdeas presents na
lista possuem valor onramental e sofrem pressdo de coleta
indiscriminada. 0 mesmo ocorre corn indmeras espdcies de
sempre-vivas, que tern sua situacao agravada por geralmente
se distribufrem em uma area restrita, nos campos rupestres
da Serra do Espinhago, principalmente na regiao de
Diamantina, maior p6lo fornecedor de sempre vivas para o
comercio. 0 pouco conhecimento existente sobre a flora mi-
neira foi evidenciado no estudo para elaboracao da lista. Um
exemplo e o da pterid6fita Huperzia rubra, induida na lista
como provavelmente extinta e que foi posteriormente encon-
trada no alto da Serra da Caraqa. Antes de significar uma
imprecisdo do mitodo, o fato indica necessidade premente
de incremento dos inventirios biol6gicas no estado. Essa
preocupaI.o ji estava present as durante a elaboracao do
Atlas da Biodiversidade de Minas Gerais, que incluiu as areas

onde ainda existem remanescentes significativas da vegetacao
native mas cujo conhecimento biol6gico e insuficiente. A
indicacao dessas Areas objetivou incentivar o direcionamento
de pesquisas e prioridades aos pesquisadores e 6rgaos
financiadores de fomento. Alim de center a lista das especies
ameagadas, a publicaqo traz uma caracterizacao geral dos
biomas e dos grandes grupos vegetais que ocorrem em Minas
Gerais. 0 livro poderi ser adquirido na: Fundacgo
Biodiversitas, Rua Ludgero Dolabela 1012, 7o. Andar,
30430-130 Belo Horizonte, Minas Gerais, Brasil, Tel: (0)31
292 8235, Fax: (0)31 291 7658, e-mail: < biodiversitas@
biodiversitas.org>. Home page: .

Conservation of Biological Resources, by E. J. Milner-Gulland
and Ruth Mace (with contributors), 1998, 416pp, 114
illustrations. Blackwell Science, Oxford. Paperback ISBN 0
86554 2738 0. Price: 24.95. A student textbook presenting
the issues surrounding the biological conservation of species
and ecosystems used by humans. It is aimed at final year
undergraduate and Master's level students as well conservation
professionals, including managers, policy-makers and
researchers. Contents: Part One. Introduction to Biological
Conservation and Sustainable Use. Part Two. Theoretical
Background. 1. The ecological and economic theory behind
sustainable harvesting. 2. Harvesting and ecological realities.
3. Decision-making by users of natural resources. 4. Practical
considerations when applying the theory. Part Three. Case
Studies. Part Four. Making Conservation Work. Part 3
includes contributions by Kathy Mackinnon (Sustainable use
as a conservation tool in the forests of South-east Asia), R. E.
Gullison (Will bigleaf mahogany be conserved through
sustainable use?), Vivienne Solis Rivera and Steven Edwards
(Cosigiiina, Nicaragua: A case study in community-based
management of wildlife), Sophie des Clers (Sustainability of
the Falkland Islands loligo squid fishery), Andrew Price,
Callum Roberts and Julie Hawkins (Recreational use of coral
reefs in the Maldives and Caribbean), Joel Freehling and
Stuart A. Marks (A century of change in the Central Luangwa
Valley of Zambia), M. Norton-Griffiths (The economics of
wildlife conservation policy in Kenya), Tom Butynski and Jan
Kalina (Gorilla tourism: A critical look), Anne Gunn (Caribou
and muskox harvesting in the Northwest Territories), and
Leonid Baskin (Hunting of game mammals in the Soviet
Union). Available from: Anna Van Opstal, Blackwell Science
Ltd., Osney Mead, Oxford OX2 OEL, UK, Tel: +44 (0)1865
206206, Fax: +44 (0)1865 721205. Website: http://

Developing a Social Psychology of Monkeys and Apes, by John
Chadwick-Jones, 1998. 208pp. Psychology Press, London.
Hardback ISBN 0 86377 820 8. Price: 29.95. Contents:
Introduction: social psychology and primates; Intentionality,
deception and social intelligence; Primate communication and
social psychology; Facial expressions; Gestures, postures and
touch; Vocal signals: apes; Vocal signals: monkeys; Tactics and
social devices; Social exchange and grooming partnerships;
Sexuality in monkeys and apes; Dominance and social
relationships; Primate studies and social psychology.

Neotropical Primates 8(1), March 2000

Appendices. Available from: Afterhurst Ltd. Mail Order
Service, c/o The Book Ordering Dept., Taylor and Francis
Ltd., Rankine Road, Basingstoke, Hants RG24 8PR, UK, Tel:
+44 (0)1256 813000, Fax: +44 (0)1256 479438, e-mail:

Species Coexistence, by M. Tokeshi, 1998, 464pp. 159
illustrations. Blackwell Science, Oxford. Paperback ISBN 0
86542 744 5. Price: 37.50. Contents: Introduction:
conceptual threads; Origination: the basis of coexistence;
Origination and evolution of communities; Patterns in species
richness: temporal dimension; Patterns in species richness:
spatial dimension; The niche, resources and species assembly;
Niche apportionment and relative abundances of coexisting
species; Competition, co-operation and coexistence; Agent-
mediated coexistence: predation and disturbance; Patchiness,
heterogeneity and stochasticity; Traits and coexistence;
Stability and conservation of coexisting species; Concluding
remarks. Available from: Anna Van Opstal, Blackwell Science
Ltd., Osney Mead, Oxford OX2 OEL, UK, Tel: +44 (0)1865
206206, Fax: +44 (0)1865 721205. Website: http://

Old World Monkeys, edited by Paul E Whitehead and Clifford
J. Jolly, 2000, 528pp. Cambridge University Press, UK.
Hardback ISBN 0 521 57124 3. Price: US$115. Old World
monkeys, Cercopithecoidea, are the most successful and
diverse group of non-human primates. The species
represented in this group have much to teach us about
evolution and ecology. This book provides a broad technical
account of cercopithecoid biology, including molecular and,
behavioral and morphological approaches to phylogeny;
population structure; allometry; fossil history; functional
morphology; ecology; cognitive capabilities; social behavior
and conservation; it is a definitive reference for those
professionals and students wishing to further their
understanding of primatology. Available from: Cambridge
University Press, The Edinburgh Building, Cambridge CB2
2RU, UK, or 40 West 20th Street, New York, NY 10011-
4211, USA. Web site: www.cup.org>.


Anonymous. 2000. Guidelines for the treatment of animals
in behavioral research and teaching. Anim. Behav. 59(1):
Baker, J. V., Abbott, D. H. and Saltzman, W 1999. Social
determinants of reproductive failure in male common mar-
mosets housed with their natal family. Anim. Behav. 58(3):
Boubli, J. P. 1999. Feeding ecology of black-headed uakaris
(Cacajao melanocephalus melanocephalus) in Pico da Neblina
National Park. Int. J. Primatol. 20(5): 719-749.
Chaves, R., Sampaio, I., Schneider, M. P., Schneider, H., Page,
S. L. and Goodman, M. 1999. The place of Callimico goeldii

in the calltrichine phylogenetic tree: Evidence from von
Willebrand factor gene Intron II sequences. Molec.
Phylogenet. Evol. 13(2): 392-404.
Cincotta, R. P., Wisnewski, J. and Engelman, R. 2000. Hu-
man population in the biodiversity hotspots. Nature, Lond.
404: 990-992.
Defler, T. R. 1999. Locomotion and posture in Lagothrix
lagothricha. Folia Primatol. 70(6): 313-327.
Ferrari, S. E, Iwanaga, S., Coutinho, P. E. G., Messias, M. R.,
Cruz Neto, E. H., Ramos, E. M., and Ramos, P. C. S. 1999.
Zoogeography of Chiropotes albinasus (Platyrrhini, Atelidae)
in southwestern Amazonia. Int. J. Primatol. 20: 995-1004.
Ferrari, S. E, Iwanaga, S., Messias, M.R., Ramos, P.C.S., Cruz
Neto, E. H., and Coutinho, P. E. G. 2000. Titi Monkeys
(Callicebus spp., Atelidae Platyrrhini) in the Brazilian state
of Rondonia. Primates 41: 191-196.
Garcia, E, Noguds, C., Garcia, M., Egozcue, J. and Ponsi,
M. 1999. Characterization of constitutive heterochroma-
tin in Cebus apella (Cebidae, Primates) and Pan troglodytes
(Hominidae, Primates): Comparison to human chromo-
somes. Am. J. Primatol. 49(3): 205-221.
Giminez, J. A. G. 2000. Perfil del primat6logo en la Peninsula
Ibdrica. Boletin de la Associacion Primatoldgica Espanola 7(1):
Gullison, R. E., Rice, R. E. and Blundell, A.G. 2000. 'Mar-
keting' species conservation. Nature, Lond. 404: 923-924.
Harcourt, A. H. 2000. Coincidence and mismatch of
biodiversity hotspots: A global survey for the order Primates.
Biol. Conserv. 93(2): 163-175.
Herrera, E. R. T, Knogge, C. and Heymann, E. W 2000.
Infanticide in a group of wild saddle-back tamarins,
Saguinus fuscicollis. Am. J. Primatol. 50: 153-157.
Heymann, E. W 2000. Primate behavioral ecology and dis-
eases some perspectives for a future primatology. Primate
Report (55): 53-65.
Heymann, E. W and Smith, A. C. 1999. When to feed on
gums: Temporal patterns of gummivory in wild tamarins,
Saguinus mystax and Saguinusfiuscicollis (Callitrichinae). Zoo.
Biol. 18: 459-471.
Heymann, E. W and Buchanan-Smith, H. M. 2000. The
behavioral ecology of mixed species troops of callitrichine
primates. Biol. Rev. 75: 169-190.
Heymann, E. W., Knogge, C. and Herrera, E. R. T. 2000.
Vertebrate predation by sympatric tamarins, Saguinus mystax
and Saguinus fiscicollis. Am. J. Primatol. 51: 153-158.
Krishnamani, R. and Mahaney, W. C. 2000. Geophagy
among primates: Adaptive significance and ecological con-
sequences. Anim. Behav. 59(5): 899-915.
Mittermeier, R. A. and Konstant, W R. 2000. Worlds van-
ishing primates. Australasian Primatology 14(2): 2-3.
Mittermeier, R. A., Fonseca, G. A. B., Rylands, A. B. and
Mittermeier, C. G. 1999. Atlantic Forest. In: Hotspots: The
Earths Biologically Richest and Most Endangered Terrestrial
Ecoregions, R. A. Mittermeier, N. Myers, P Robles Gil & C.
G. Mittermeier (eds.), pp.136-147. Cemex, Conservation
International, Agrupaci6n Sierra Madre, Monterrey,
Myers, N., Mittermeier, R. A., Mittermeier, C. G., Fonseca,

NeotropicalPrimates 8(1), March 2000

G. A. B. da and Kent, J. 2000. Biodiversity hotspots for
conservation priorities. Nature, Lond. 403: 853-858.
Norcross, J. L., Newman, J. D. and Cofrancesco, L. M. 1999.
Context and sex differences exist in the acoustic structure
of phee calls by newly-paired common marmosets (Callithrix
jacchus). Am. J. Primatol. 49(2): 165-181.
Oates, J. F 2000. Why a prime model for saving rain forests
is a failure. Australasian Primates 14(2): 10-12.
Passamani, M. & Rylands, A. B. 2000. Feeding behavior of
Geoffroys marmoset (Callithrix geoffroyi) in an Atlantic for-
est fragment of south-eastern Brazil. Primates 41(1): 29-40.
Passamani, M. and Rylands, A. B. 2000. Home range of a
Geoffroys marmoset group, Callithrix geoffroyi (Primates,
Callitrichidae) in south-eastern Brazil. Rev. Brasil. Biol.
60(2): 275-281.
Pimm, S. L. and Raven, P. 2000. Extinction by numbers. Na-
ture, Lond. 403: 843-845.
Richard-Hansen, C., Vie, J. C., de Thoisy, B. 2000. Translo-
cation of red howler monkeys (Alouatta seniculus) in French
Guiana. Biol. Conserv. 93(2): 247-253.
Roush, R. S. and Snowdon, C. T. 1999. The effects of social
status on food-associated calling behaviour in captive cot-
ton-top tamarins. Anim. Behav. 58(6): 1299-1306.
Schneider, H. 2000. The current status of the New World
monkey phylogeny. An. Acad. Bras. Cienc. 72(2): 165-172.
Setz, E. Z. E, Enzweiler, J., Solferini, V. N., Amnndola, M. P.
and Berton, R. S. 1999. Geophagy in the golden-faced saki
monkey (Pitheciapithecia chrysocephala) in the central Ama-
zon. J. ZooL, Lond. 247: 91-103.
Sousa e Silva, J. de., Jr. and Noronha, M. de A. 1998. On a
new species of bare-eared marmoset, genus Callithrix
Erxleben, 1777, from central Amazonia, Brazil (Primates:
Callitrichidae). Goeldiana Zoologia (21): 1-28.
Silver, S. C., Ostro, L. E. T., Yeager, C. P. and Dierenfeld, E.
S. 2000. Phytochemical and mineral components of foods
consumed by black howler monkeys (Alouattapigra) at two
sites in Belize. Zoo Biol. 19(2): 95-109.
Strier, K. B. and Zeigler, T. E. 2000. Lack of pubertal influ-
ences on female dispersal in muriqui monkeys, Brachyteles
arachnoides. Anim. Behav. 59(4): 849-860.
Tabarelli, M., Mantovani, W. and Peres, C. A. 1999. Effects
of habitat fragmentation on plant guild structure in the
montane Atlantic forest of southeastern Brazil. Biol. Conserv.
91: 119-127.
Visalberghi, E. and Addessi, E. 2000. Response to change in
food palatability in tufted capuchin monkeys, Cebus apella.
Anim. Behav. 59(1): 231-238.


Dunbar, R. 1999. The number of males in primate groups.
Primate Eye (69): 11-12.
Herndndez-L6pez, L. and Mondrag6n-Ceballos, R. 1999.
The reproductive physiology of the black-handed spider
monkey (Ateles geoffloyi). Primate Eye (69): 17.
Prescott, M. J. 1999. Social learning in mixed-species troops

of Saguinus fuscicollis and Saguinus labiatus: Tests of forag-
ing benefit hypotheses in captivity. Primate Eye (69): 29.
(Ph.D. abstract).
Rodriguez, S. S. 2000. Costos e beneficios del transport de
las crias en el sistema de crianza cooperative del titi de cabeza
blanca (Saguinus oedipus) (Callitrichidae, Primates). Boletin
de la Associacidn Primatoldgica Espanola 7(1): 24. Thesis

IUCN the World Conservation Union World Conservation
Congress, 4-11 October, 2000. Amman, Jordan. The IUCN/
SSC meeting will be held on 2-3 October, 2000. For more
information: IUCN-The World Conservation Union, Rua
Mauverney 28, CH-1196 Gland, Switzerland, Tel: +41 22
999 0001, Fax; +41 22 999 0002, e-mail:

Nonhuman Primate Pathology Seminar and Workshop
2000, 7-8 October, 2000, Wisconsin Regional Primate
Research Center, University of Wisconsin- Madison, USA,
in conjunction with the Symposium on Nonhuman Primate
Models for Aids (4-7 October). The conference will be held
at the Pyle Conference Center. For more information and
registration materials please contact either Dr. Amy Usborne,
e-mail: or Dr. Iris Bolton, e-
mail: or at WRPRC, University
Of Wisconsin, 1220 Capitol Court, Madison, WI 53715-
1299, USA.

Associazione Primatologica Italiana Congress (XIVth), 4-6
October 2000, Pisa Italy. For more information contact: Pro-
fessor Silvana Borgognini Tarli, Dipartimento di Etologia, Eco-
logia, Evoluzione, Via Volta, 4-56126 Pisa, Italy, Tel: (050)
24613 or (050) 44484, Fax: (050) 24653, e-mail:
. Home Page: www.unipv.it/webbio/api/congl4/14con.htm

II Congress Brasileiro de Unidades de Conservaqio, 5-9
de November 2000, Centro de Conven96es de Campo
Grande, Mato Grosso do Sul, Brazil. Commemorating 60
years after the creation of the Brazils 1 National Park. For
more information contact: Fundacqo 0 Boticirio de Proteqdo
a Natureza, Av. Rui Barbosa 3450, Afonso Pena, 83065-260
Sdo Jose dos Pinhais, Parana, Brazil, Tel: (041) 382 3456,
Fax: (041) 382 4179, E-mail: fundacao@fbpn.org.br. Home
Page: .

Primates in Biomedical Research: Diseases and Pathology -
2"d Gottingen Symposium, 8-9 November 2000,
Department of Veterinary Medicine and Primate Husbandry,
Deutsches Primatenzentrum. Focus: Spontaneous and
induced primate pathology. A special EUPREN Workshop
"The Future of Primates in Biomedical Research" will be held

60 Neotropical Primates 8(1), March 2000

in conjunction with this symposium on 10 November 2000.
All symposium participants are invited to attend. Contact:
Conference Secretariat: Ingrid Rossbach, Department of
Veterinary Medicine, Deutsches Primatenzentrum,
Kellnerweg 4, D-37077 Gbttingen, Germany, Tel: 49 (0) 551-
3851 119, Fax: 49 (0) 551-3851 277, e-mail:

Association for the Study of Animal Behaviour Winter
Meeting, 30 November 1 December, 2000, Zoological
Society of London Meeting Rooms, London, UK. Organised
by Andrd Gilburn. Theme: "Sexual Conflict". For further
information: Dr. Andrd S. Gilburn, Department of Biology,
University of Leicester, Adrian Building, University Road,
Leicester LE1 7RH, England, UK, Tel: +44 (0)116 252 3488,
Fax: +44 (0)116 252 3330.

Association of Primate Veterinarians Annual Workshop, 3-
5 November, 2000, San Diego, California. A case report
presentation dealing with aspects of clinical medicine or
husbandry/management of nonhuman primates. Those
veterinarians enrolled in laboratory animal, primate medicine,
or zoological medicine training programs or residencies may
apply by September 30, 2000 for a travel award of $500.00
to attend the workshop and present a clinical case report or
research findings. Contact: Dr. Nick Lerche, Tel: 530 752
6490, Fax: 530 752 2880, or email


British Ecological Society '2000' Winter Meeting, 3-5
January 2001, University of Birmingham, England, UK. For
more information: British Ecological Society, 26 Blades Court,
Deodar Road, Putney, London SW15 2NU, England, UK.

XVIIIth Congress of the International Primatological
Society, 7-12 January 2001, Adelaide, Australia. Hosted by
the Australasian Primate Society, President Mr. John Lemon,
Western Plains Zoo, Dubbo, NSW. Theme: "Primates in the
New Millenium". Mr. Graeme Crook is Chairman of the
Organizing Committee. Symposia Participants wishing to
register a symposium title must submit a 200 word abstract
by 31 July 1999. E-mail to Carla Litchfield
. Titles of accepted symposia will be
published on the webpage from August 1999. Papers An
abstract of 100 words is required. E-mail to Carla Litchfield
. Closing date for first call for papers:
31 January 2000. Closing date for second call for papers: 31
May 2000. A final list of papers will be published on the
Internet by 30 June 2000. For more information, and to be
put onto the Congress Organizer's mailing list, write to:
Conventions Worldwide, PO Box 44, Rundle Mall, SA 5000,
Australia, Tel: +61 8 8363 0068, Fax: +61 8 8363 0354, e-
mail: , sending your postal address.

International Conference "Ecology of Insular Biotas", 12-
16 February 2001, Victoria University of Wellington,
Wellington, New Zealand. Focus: ecological patterns and

processes of importance to isolated biotas, including true
islands, and natural and artificial habitat islands. Examples of
topics for papers include: dispersal and gene flow within and
among isolated populations; ecology of small populations;
ecological consequences of disharmonic floras and faunas; the
relevance of island biogeography principles in conservation;
islands as model ecosystems; and comparative ecology of true
islands vs. habitat islands. Abstracts may be submitted
electronically on the webpage (http://www.vuw.ac.nz/sbs/
conferences/island.shtml) and submitting the requested
information to . Deadline for
abstracts is 1 October 2000. If web access is not available,
contact Dr. Christa Mulder, School of Biological Sciences,
Victoria University of Wellington, P.O. Box 600, Wellington,
New Zealand. The complete scientific program will be
available on the conference website by 15 November 2000.

14th Annual Meeting of the Society for Tropical Ecology,
14-16 February 2001, Zentrum fiir Marine Tropenokologie
(Center for Marine Tropical Ecology), Bremen, Germany.
Topics: Aquatic ecosystems, Neotropics, Interdisciplinary
research. Free topics in tropical ecology. Contact: Zentrum
fiir Marine Tropen6kologie, Fahrenheitstr. 1, D-28359
Bremen, Germany, Tel: +49 421-2380-029, Fax: +49 421-
2208-330, e-mail:< gtoe2001@zmt.uni-bremen.de>.
Homepages: , bremen.de>.

Association for the Study of Animal Behaviour Spring
Meeting 2001, University of Liverpool, Liverpool, UK.
Organized by Jane Hurst. For more information.: Prof. J.
Hurst, Division of Animal Husbandry, Faculty of Veterinary
Science, University of Liverpool, Leahurst, Neston, South
Wirral L64 7TE, England, UK, Tel: +44 (0)151 7946100,
Fax: +44 (0)151 7946107, e-mail: .

Association for the Study of Animal Behaviour Summer
Meeting 2001, University of Glasgow, Scotland, UK.
Organized by Felicity Huntingford. Theme: "Interfacing
Behaviour with Other Disciplines". For more information:
Prof. F. A. Huntingford, Division of Environmental and
Evolutionary Biology, Graham Kerr Building, Glasgow
University, Glasgow G12 8QQ, Scotland, UK, Tel: +44
(0)141 330 5968, Fax: +44 (0)141 330 5971, e-mail:

The First International Conference on Distance Sampling,
30 July-3 August, 2001. St. Andrews School, Scotland.
Organized by Rhona Rodger. Theme: Estimating Wildlife
Abundance Ecology, Management and Conservation Tel: +44
(0)133 446 3228, e-mail:,
Homepage: .


The journal/newsletter aims to provide a basis for conserva-
tion information relating to the primates of the neotropics.
We welcome texts on any aspect of primate conservation, in-
cluding articles, thesis abstracts, news items, recent events,
recent publications, primatological society information, and


Please send all English and Portuguese contributions to:
Jennifer Pervola, Conservation International, Center for Ap-
plied Biodiversity Science, 1919 M. St. NW, Suite 600, Wash-
ington, DC 20036, Tel: 202 533 9533, Fax: 202 331 0570,
e-mail: , and all Spanish con-
tributions to: Ernesto Rodriguez-Luna, Instituto de
Neuroetologfa, Universidad Veracruzana, Apartado Postal
566, Xalapa 91000 Veracruz, Mexico. Tel: 281 8-77-30, Fax:
281 8-77-30, 8-63-52, e-mail: speedy.coacade.uv.mx>


Manuscripts can be in English, Spanish or Portuguese, and
should be double-spaced and accompanied by the text on
diskette for PC compatible text-editors (MS-Word,
WordPerfect, Excel, and Access), and/or e-mailed to <
j.pervola @ conservation. org>.(English, Portuguese) or <
saraguat @speedy. coacade.uv.mx> (Spanish) Hard copies
should be supplied for all figures (illustrations and maps) and
tables. The full name and address for each author should be
included. Please avoid abbreviations and acronyms without
the name in full. Authors whose first language is not English,
please have texts carefully reviewed by a native English speaker.

Articles. Each issue of Neotropical Primates will include up to
three full articles, limited to the following topics: Taxonomy,
Systematics, Genetics (when relevant for systematics), Bioge-
ography, Ecology and Conservation. Texts for full articles
should not exceed about 20 pages in length (1.5 spaced, and
including the references). Please include an abstract in En-
glish, and (optional) one in Portuguese or Spanish. Tables and
illustrations should be limited to six, excepting only the cases
where they are fundamental for the text (as in species de-
scriptions, for example). Full articles will be sent out for peer-

Short articles. These are reviewed only by the editors. A
broader range of topics are encouraged, including such as
behavioral research, in the interests of informing on general
research activities which contribute to our understanding of
platyrrhines. We encourage reports on projects and conser-
vation and research programs (who, what, where, when, why,

etc.) and most particularly information on geographical dis-
tributions, locality records, and protected areas and the pri-
mates which occur in them. Texts should not exceed 10 pages
in length (1.5 spaced, including the references).

Figures and maps. Articles can include small black-and-white
photographs, high quality figures, and high quality maps and
tables. Please keep these to a minimum. We stress the impor-
tance of providing maps which are publishable.

News items. Please send us information on projects, field sites,
courses, recent publications, awards, events, activities of Pri-
mate Societies, awards, etc.

References. Examples of house style can be found throughout
this journal. Please refer to these examples when citing refer-

Journal article. Stallings, J. D. and Mittermeier, R. A. 1983.
The black-tailed marmoset (Callithrix 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
with Conservation International, Center for Ap-
plied Biodiversity Science, 2501 M. St. NW, Suite
200, Washington, DC 20037.

Neotropical Primate'
journal and Newsletter of the IUClN/SSC Primate Specialist Group
VoL 8(1), March, 2000


Editorial .... i 3 R .:.'.'. ., E- ..cr A .i .'. : L ... ..... ............ ...... ... ..................


Two new species of marmoser. genu_ ',al.t/tr-.v ErxeJben. 1-'- iCallirnchuiae, Primatesi,
from the Tapai/4l/Madeira minerthihum. south Ceonal Amaronia. Brazil
1.. G. .il ., .; .A.... ..-. 1:-... .... ., :.,I..,-/, s ,,.. .l1. *:. ..... ,...1:.1., .,, 8.; 1 ,
Reconocrumento de Ia poblacion del mono aullador negro (.lIouiarm pt'ra, en Palenque.
Cluapas Mexico
/. ,, t, .:,, : .. d. i, .. ... ...-*., i-.,.;' ,? .i, , .C .a ,,.,.,. .4. ( ...,^ .T ... ... . ". ........ ..
".. ,,. i t ..... :... h r .....
.4 ,.5 _"; ,' c. i. :. .,i,; G. .'; L/ '.; "- r '. ,:, : '. w, b e i:.e l .a -r ...... .. . .. .... ... .. ... .. :.. .. ....... . ... ... 1 -'
Prim.ate diversity. disriburion and relative abundance in the Rion RBanco ) Negro \\ildlife Reserve.
Sa.nL. Craz Depar-mcnt. Bolivia
A.'u -er b. ln'.dla...r tI P...: L I' *?, L *.-: 1. :i;, d 'a-, 3. 1 . ...... .. .. .. ......

Short Artic .

Novos regiseros de. 4/owarra no crado do-Ceari iPrinraie.. Arelida
I:,..r... .'G.C L ..._I B :- .,jO -".. a'.J ". 1 .. L 1:,' L.-i..,, . ........ . . .. .....
-Noa ocorrentiade Brialte-./i an-I/woidw'- no Parque Estdual da Siraa do-Mir. Sao Piulo. Brasil
.',.,'. 4,, i, /':.- IL..* LJ,,, p _" ,r ';. F. r,.', .4/ .. ,' . . ... .. . . :-... . ,
S- Fal;Adispersal in the Bdlizan -[ak houi linig monnkey A/a ri l . I.
. ~. tS.- .. .- H ,-ia ./ Cl '. B i ,.c ... ... ........ .. .. .... .. .... .. .. .. .:
-pd-arfid ht sam orfth% O I rgaraiu island capuchAn. Ce(- apei miarwri," .
.. l -.l .-t..-j ,,-, .j..;;i\ ..a ./n.A ..1^a ..- ....-; .-- ........:. _..- --i
r- mare rcordis from thePotaro Plateau. esteem m Gmuan-x including the fir-Tfdr Liu Ibifio ns .
-east ofihe Rio Branco.- Brazil
-. 1. '- ._.r. .1 Eb __ ai! ";:!. . / ci.*'.>- i i .... -
'1/1-:, .-. A I .' ;. .t .,'. P br it v '..; .. .... ... . .. .. .... . ..... . ... .. .. ...... . .... .... i .
Sr- .of .lnunta. p .d/ t;Ir.the Bilsa Biological-Remer'c. norrhn-esr Ecuador -
S-,' ; -i-:. r..-r ' L / ..,..... ~. ., '. i .- ;o. -: '.,: ., .. .: .... .. ..... 4 "
tmogriapbr of a group of utited capuchun maknlkes iLcebts ip;//an ngrintsi at the fstacao -
Bioldgica-de Caratinga, Minims Gerak Brail

N . ... .........F.'................... -................
N.- . . .. . . .. .. .

Prim ate -ocienes ... . ...... .. ...... ............. .............. ... .. .. ..... . .......... .... ....... .. .

Rc L .Publkatior. ... .. .. .. .................... ......... ..... ... -. ..

N.M stings .. .

University of Florida Home Page
© 2004 - 2010 University of Florida George A. Smathers Libraries.
All rights reserved.

Acceptable Use, Copyright, and Disclaimer Statement
Last updated October 10, 2010 - - mvs