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Title: Neotropical primates
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Permanent Link: http://ufdc.ufl.edu/UF00098814/00050
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Title: Neotropical primates a newsletter of the Neotropical Section of the IUCNSSC Primate Specialist Group
Abbreviated Title: Neotrop. primates
Physical Description: v. : ill. ; 27 cm.
Language: English
Creator: IUCN/SSC Primate Specialist Group -- Neotropical Section
IUCN/SSC Primate Specialist Group -- Neotropical Section
Conservation International
Center for Applied Biodiversity Science
Publisher: Conservation International
Place of Publication: Belo Horizonte Minas Gerais Brazil
Belo Horizonte Minas Gerais Brazil
Publication Date: December 2005
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).
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Table of Contents
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    Back Matter
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    Back Cover
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Full Text
ISSN 1413-4703



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


Anthony B. Rylands
Ernesto Rodriguez-Luna
Assistant Editors
John M. Aguiar
Liliana Cort6s-Ortiz
PSG Chairman
Russell A. Mittermeier
PSG Deputy Chairman
Anthony B. Rylands





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

Center for Applied Biodiversity Science
Conservation International
2011 Crystal Drive, Suite 500, Arlington, VA 22202, USA

ISSN 1413-4703 Abbreviation: Neotrop. Primates
DOI: 10.1896/ci.cabs.2005.np.13.3

Anthony B. Rylands, Center for Applied Biodiversity Science, Conservation International, Arlington, VA, USA
Ernesto .1 ,,_. -T ..... UniversidadVeracruzana, Xalapa, Mexico

Assistant Editors
John M. Aguiar, Center for Applied Biodiversity Science, Conservation International, Arlington, VA, USA
Liliana Cortes-Ortiz, Universidad Veracruzana, Xalapa, Mexico

Editorial Board
Hannah M. Buchanan-Smith, University of Stirling, Stirling, Scotland, UK
Adelmar E Coimbra-Filho, Academia Brasileira de Ciencias, Rio de Janeiro, Brazil
Liliana Cortes-Ortiz, Universidad Veracruzana, Xalapa, Mexico
Carolyn M. Crockett, Regional Primate Research Center, University of i ...... Seattle, WA, USA
Stephen E Ferrari, Universidade Federal do Para, Belem, Brazil
Eckhard W. Heymann, Deutsches Primatenzentrum, C -1...... Germany
Russell A. Mittermeier, Conservation International, Arlington, VA, USA
Marta D. Mudry, Universidad de .
Horacio Schneider, Universidade Federal do Para, Belem, Brazil
Karen B. Strier, University of Wisconsin, Madison, WI, USA
Maria Emilia Yamamoto, Universidade Federal do Rio Grande do Norte, Natal, Brazil

Primate Specialist Group
Chairman Russell A. Mittermeier
Deputy Chair Anthony B. Rylands
Co-Vice Chairs for the Neotropical Region Anthony B. Rylands & Ernesto - .
Vice Chair for Asia Ardith A. Eudey
Vice Chair for Africa Thomas M. Butynski
Vice Chair for Madagascar Jorg U. Ganzhorn

Layout: Kim Meek, Center for Applied Biodiversity Science, Conservation International, Arlington, VA, USA

Editorial Assistance:
Mariella Superina, University of New Orleans, Department of- I i Sciences, New Orleans, LA

IUCN/SSC Primate Specialist Group logo courtesy of Stephen D. Nash, 2002.

Front cover: A juvenile Ateles belzebuth from the Peruvian Andes. Photo by Russell A. Mittermeier.

This issue of Neotropical Primates was kindly sponsored by the Margot Marsh Biodiversity Foundation, 432 Walker Road, Great Falls, Virginia 22066,
USA, and the Los Angeles Zoo, Director John R. Lewis, 5333 Zoo Drive, Los Angeles, California 90027, USA.

Neotropical Primates 13(3), December 2005


Pedro G. Mendez-Carvajal


The taxonomic identity of the Azuero howler monkey has
been controversial. Lawrence (1933) initially described this
taxon as a subspecies ofA. palliata. In 1987, Froehlich and
Froehlich analyzed the fingerprint pattern of different spe-
cies of howler monkeys as a proxy to infer genetic distance.
They concluded that A. p. coibensis should be considered
a distinct species, A. coibensis. In this analysis they also
found A. p. trabeata to be closer to the coibensis form than
to other A. palliata forms, and suggested that this taxon
be a subspecies of A. coibensis. Cortes-Ortiz et al. (2003)
carried out genetic studies (mitochondrial DNA) of howler
monkeys along their entire geographic range and, inferring
the phylogenetic relationships among the species, they con-
cluded that both trabeata and coibensis share mitochondrial
haplotypes with other forms of howler monkeys in Central
America (coibensis with mexicana and palliata, and trabeata
with aequatorialis), supporting the initial classification of
Lawrence, and suggesting that they are at best subspecies
of A. palliata.

The original distribution of the Azuero howler monkey
covered most of the Azuero peninsula in Panama, includ-
ing portions of the provinces ofVeraguas, Herrera and Los
Santos (Mendez, 1970; Froehlich and Froehlich, 1986,
1987; Arauz, 1998; Mendez, 1999). This subspecies is an
endemic taxon under high risk of extinction if the present
trends of forest destruction continue (Rodrfguez-Luna et
al., 1996; Rylands et al., 2000; Mendez, 2002).

In order to better understand the current status of the
Azuero howler monkey, I conducted a survey in the north-
ern part of its range in the province of Herrera (Fig. 1).
Demographic and behavioral data were recorded, as well
as the quality of the habitat in the area. I also used the
information collected on these howler monkeys to initiate
an education program with local people aiming at the con-
servation of this primate in Azuero.

Study site

The Azuero peninsula is in southwestern Panama. The
annual average temperature is 28.1C (22.5C to 33.7C)
and annual rainfall averages 1,410 mm/year (Contraloria
General de la Rep dblica, 2001). The dry season is from De-
cember to April and the rainy season from May to Decem-
ber (Suarez, 1981). The area surveyed was quite flat with

small hills of 90 to 150 m (Mendez, 2001). Remnant forest
can be found on the hilltops and along the rivers. The tall-
est trees are about 15 to 20 m high (Mendez, 2001). There
is only one patch of forest that the farmers have left more
or less untouched. The landscape in the region is mostly
grassland and cattle pasture, with a few fragments of forest,
some of them connected by narrow strips of trees and
sparse riparian forest.


I first traveled through the region (15 towns) asking local
people about the presence of monkeys in order to get some
understanding of the location of the howler populations.
Fieldwork was subsequently conducted for five consecutive
days each month, from April to December 2001. I surveyed
all the trails and roads between the towns of Ocd and Parita
(Fig. 1), from 6:00 am to about 2:00 pm. On finding a
group of monkeys, I recorded their location, the number of
individuals and the composition (following Milton, 1982),
and also noted aspects of their behavior and the quality of
the habitat. I spent as much time as I could observing the
behavior of the group and whenever possible following and
recording the activities of particular individuals I was able
to identify. Remaining with the group until the howlers
vocalized made it possible to detect other groups in the vi-
cinity. When hearing other groups, I recorded the compass
bearing, inferred the distance, and went in search of them.
Care was taken not to double-count groups or individuals.


Three hundred and sixty hours were spent in fieldwork.
Thirty percent of this time was devoted to searching for
the howler monkeys and the remaining 70% was dedicated
to accompanying the groups, identifying sex-age composi-
tion, and conducting direct observations on behavior. In
total I visited 15 towns and their surrounding areas, along
24 linear kilometers (Fig. 1, Table 1). Howler monkeys
were found around only three of the towns: Santa M6nica,
Llano Grande, and Llano Hato. Eleven howler monkey
groups were located. In Santa M6nica I also found captive
monkeys of two other species, Cebus capucinus and Saguinus
.. rr.., that were taken from Eastern Panama (Cerro Azul
and Darien respectively). In the town of Camaron (Fig. 1)
I was told of an introduction of a male howler monkey that
was brought from La Chorrera, in the Panama Canal area,
and therefore A. p. aequatorialis. However, I was unable to
find any howler monkeys in that area. Local people also re-
ported a case when a howler monkey was hunted and eaten
in the town of Pedregosito (Fig. 1).

I counted 119 howler monkeys in five groups by direct ob-
servations. The average size of the groups was 23.8 (range
15-39) individuals. Six more groups were heard but never
found in a forest of approximately 400 ha between the
towns of Llano Grande and Llano Hato. The forest around
Santa M6nica totaled approximately 245 ha in two patches

Neotropical Primates 13(3), December 2005

of forest and a riparian forest corridor. All these forest frag-
ments are surrounded by grassland. In total I estimated
262 howler monkeys in the area between Ocd and Parita
towns, in the central part of Herrera Province. Population
density in the area was calculated to be 40.6 individuals/
km2 and 1.7 troops/km2. The average sex-age composition
of the groups was 6.0 adult males, 7.8 adult females, 6.6 ju-
veniles and 3.4 infants. Female/male and immature/female
ratios were 1:1.3 and 1:1.39 respectively (Table 2).

Habitat use

wire fences to travel from one tree to another, and even
run through the grassland to reach other trees. They evi-
dently have a taste for mango fruits (Mangifera indica), a
very common introduced tree in Panama. Table 3 gives a
list of trees that were used by howler monkeys (as food or
to rest in) during the survey. The majority of the tree spe-
cies reported in this survey have also been reported by other
authors as sources of food for howler monkeys in different
localities (e.g., Milton, 1982; Terborgh, 1983).


At least in this part of the Azuero peninsula, the howler
monkeys showed no strong preference for any particular
types of forest. Our observation did, however, demonstrate
their attachment to certain trees. On some occasions we
watched them spend up to 60% of the day in the same
group of trees (Enterolobium cyclocarpum, Bursera simaru-
ba, Ficus yoponensis and Spondias mombin) in the middle of
the thin lines of trees that separate cattle ranching proper-
ties. According to local people, the monkeys sometimes use

I estimated 262 individual howler monkeys in 11 troops
living in highly fragmented forest and along the sparse cor-
ridors of trees which customarily delineate properties in
central Herrera Province. In general, the land use in the
area is a fine-grained mosaic, with small forested areas in-
terspersed with crops and enormous areas of grassland and
cattle pasture. People take wood from the forest patches and
hunt deer, rabbit, and armadillo in them. Although local
people did not admit to hunting howlers, I did witness one

Figure 1. Location of the towns between Oc6 and Parita, Azuero, Herrera Province, Republic of Panama.

Neotropical Primates 13(3), December 2005

case where an individual was hunted and eaten. This part
of the range of the Azuero howler monkey is not protected
in any way by the National Environmental Authority (Au-
toridad Nacional del Ambiente ANAM) and yet is one
of the few lowland areas in the region that still has forest
patches large enough (645 ha in total) to hold small num-
bers of groups of howler monkeys.

Despite the fragmentation of their habitat, the howler
monkeys in the region have a robust and healthy appear-
ance, suggesting that they are not lacking in food. There is
a good variety of trees in the patches of forest that the howl-
ers can exploit as food sources (Table 3), and some of the
trees left standing to delineate properties are mature and

evidently appear to be providing sufficient food and shelter.
Furthermore, the howlers are able to eat fruits from species
such as mango trees that have been introduced, likely an
important supplement to their diet, or even a mainstay at
certain times of the year.

The presence of juveniles and infants suggests that the
population is growing. Average group size for A. p. trabeata
in this region of Herrera was similar to that of other popu-
lations of howler monkeys studied in Azuero (Brandaris,
1983) and in Barro Colorado Island, Panama (Carpenter,
1934; Milton, 1982). The large number of individuals
per group may be a consequence of the impossibility of
dispersal among fragments. When howler monkeys cross

Table 1. Localities, distance and the presence of Azuero howler monkeys during the survey in Herrera Province. *We found Saguinus
and Cebus capucinus in captivity. (h) = hearing; (o) = observed.
Linear distance No. of monkeysber of troops
Town from Ocfi Habitat estimated by local Numbidentified oops
(km) people
Ocd 0 Human settlements 0 0
Las Animas 1.5 Human settlements 0 0
La Polonia 10 Human settlements, riparian forest 0 0
El Calabazal 11 Human settlements, riparian forest ? 0
Santa Mnica 11.5 Human settlements, riparian forest, forest 30-35 2(h)

Llano Grande 12 Human settlements, riparian forest, forest 20 5 (h)
fragment, grassland 3(o)

La Chavarria-Pes6 12.5 Human settlements, riparian forest, forest 1 0
fragment, grassland
Human settlements, riparian forest, forest 4(h)
Llano Hato 12.5 40
fragment, grassland 2(o0)
Llano de la Cruz 14 Human settlements, riparian forest. 3 0
Los Cantos-Parita 22.4 Human settlements, riparian forest, forest 2 0
fragment on a hill
Los Castillos-Parita 24.9 Human settlements, riparian forest, grassland 12 0
Pedregoso-Pes6 21 Human settlements, riparian forest, forest 0 0
Pedregosito-Pes6 23 Human settlements, riparian forest, grassland 1 0
Camaron 5 Human settlements, Forest Reserve

Tijeras 8 Hill 1 (h)
_____________________ _______________________________________ ___________________5(o)

Table 2. Social structure of Azuero howler monkey troops observed in Santa M6nica, Llano Grande and Llano Hato, Herrera Province.
Classification according to Milton (1982).
Troops Male Female J3 J2 J1 13 12 11 Total
T-1 5 6 1 1 0 0 1 1 15
T-2 4 5 3 3 2 0 0 1 18
T-3 10 10 8 5 1 1 1 3 39
T-4 7 11 3 1 1 1 1 1 26
T-5 4 7 3 1 0 0 1 5 21
Totals 30 39 18 11 4 2 4 11 119
Average 6.0 7.8 3.6 2.2 0.8 0.4 0.8 2.2 23.80
%0 25.2 32.8 15.1 9.2 3.4 1.7 3.4 9.2 100

Neotropical Primates 13(3), December 2005

Table 3. List of tree species used byAzuero howler monkeys and collected in Herrera Province. = monkeys eating from these trees during
the survey.
Common name Scientific name Family
Espav6* Anacardium excelsum Anacardiaceae
Mango Mangifera inlica Anacardiaceae
Malagueto hembra Xylopia aromatica Annonaceae
Malagueto macho Xylopia frutescens Annonaceae
Caracucha Plumeria sp. Apocynaceae
Lagartillo Sciadodendron excelsum Araliaceae
Palma negra Astrocaryum standleyanum Arecaceae
Uvito Bactris major Arecaceae
Barrig6n* Pseudobombax septenatum Bombacaceae
Indio desnudo* Bursera simaruba Burseraceae
Guarumo Cecropia peltata Cecropiaceae
Camaroncillo Hirtella racemosa Chrysobalanaceae
Sastra Rheedia sp. Clusiaceae
Guachapali Albizia guachapele Fabaceae
Harino Andira inermis Fabaceae
Corotu* Enterolobium cyclocarpum Fabaceae
Bobo Erythrinafusca Fabaceae
Liana Erythrina sp. Fabaceae
Sigua Ocotea dendrodaphne Lauraceae
Nance Byrsonima crassifolia Malpighiaceae
Pasmo hediondo Siparuna guianensis Monimiaceae
Higuer6n* Ficus insipida Moraceae
Higuer6n* Ficus yoponensis Moraceae
Guayabo de montafia Eugenia sp. Myrtaceae
Arbol carne Roupala montana Proteaceae
Jagua Genipa americana Rubiaceae
Naranjo Citrus sinensis Rutaceae
Mam6n* Melicoccus bijugatus Sapindaceae
Guizimo* Guazuma ulmifolia Sterculiaceae
Peine de mono Apeiba membranacea Tiliaceae

grassland to reach other fragments there is a high risk of (Crockett and Eisenberg, 1987). It is interesting to note
predation by dogs a situation reported by local people, that the unimale social system reported by Milton and Mit-
confirmed by direct observations, and reported elsewhere in termeier (1977) for A. p. coibensis (always considered as the
Azuero (Brandaris, 1983). Another potential predator that same species as A. p. trabeata but then believed to be a dif-
has been reported in Azuero is the coyote (Canis latrans), ferent species of Alouatta) differs from the one observed in
which reached the peninsula at least five years ago (2000), the population of Azuero howler monkeys.
and is currently considered a serious problem for howler
monkeys and other populations of wild animals in Azuero. I recorded only one case of aggression among the howlers,
The natural predators of howler monkeys, such as harpy but I did note that many males had scars on their bodies.
eagles (Eason, 1989; Sherman, 1991) and jaguars (Kinzey, The dominant male in group T3, for example, had lost an
1997), are no longer believed to occur in the region. eye. Another male in group T4 had a recent cut on his left
leg and was unable to use it when I first saw him, although
Age-sex composition of A. p. trabeata troops is similar to he was fully recovered when I returned a year later. Other
other A. palliata populations studied in different loca- males had easily visible scars from old wounds and fractured
tions (Carpenter, 1934; Chivers, 1969; Milton, 1982). tails. The one aggressive event occurred between two males
The social system observed in the Azuero howler monkey evidently fighting over a female. The male that was guarding
troops is multi-male/multi-female, similar to what has been the female chased and fought off an approaching male. The
reported for A. p. mexicana (Cortis-Ortiz, 1998; Dias and presence of scars, injuries and fractures in the males of these
Rodrfguez-Luna, 2003) and for other species of the genus groups suggest that there are sporadic but serious fights.

Neotropical Primates 13(3), December 2005

Although howler monkeys suffer botfly (Alouattamyia
baeri) parasitism in most of their range (Milton, 1982,
1996 for Barro Colorado Island; Cowlishaw and Dunbar,
2000 for Brazil; Cortes-Ortiz, pers. comm. for Mexico) I
was unable to find signs of botflies in the population I ob-
served in Azuero. Special conditions of soil humidity are
necessary for botflies to complete their pupal phase (K.
Milton, pers. comm.). The more arid conditions of the
Azuero peninsula may be responsible for the absence of this
parasite. Nonetheless, the cattle surrounding the howler
monkeys undoubtedly increase the probabilities of infec-
tion by screw worm larvae (Cochliomyia hominivorax), an-
other parasite reported for A. palliata in Panama (Milton,
1982). The risk of infection by this type of larvae increases
with open wounds, such as those observed in the males of
this population.

The Azuero howler monkeys in the central part of Herrera
Province, Panama, are highly endangered for the reasons
already mentioned above, including forest clearance and
fragmentation, as well as hunting for food or pets. The iso-
lation of this population of howler monkeys is jeopardiz-
ing its long-term genetic viability. As a consequence of this
survey, and in order to help with the conservation of the
Azuero howler monkeys, the Mammal Society of Panama
(SOMASPA) has initiated a conservation campaign and en-
vironmental education project in the region. Furthermore,
we will continue monitoring this population to understand
the population dynamics and social behavior of this highly
threatened primate.

Acknowledgements: I thank the Rafael Quintero Villarreal
secondary school from Ocu, and their teachers, especially
Nidia Aguirre. Thanks also to Mr. Didio Gonzalez and the
Gonzalez family, owners of the pasturelands from Llano
Hato and Llano Grande respectively, and to Mr. Luis Car-
rasco and the people from Ocd and Parita for their hospital-
ity and help in finding and protecting the howler monkeys
in the area. I greatly appreciate the enthusiasm of the stu-
dents Emigdio Mitre, Wedlis Gonzalez and Donald Osorio
who helped with observations in the field. Thanks to Ivel-
isse Ruiz-Bernard, Alonso Santos and Ricardo Moreno for
supporting this research, and Agustin Somosa for the plant
identifications. My sincere gratitude to Liliana Cortes-Ortiz
for her advice and assistance on this manuscript.

Pedro G. M6ndez-Carvajal, Sociedad Mastozool6gica de
Panama, Comisi6n de Primatologfa, Apartado 797 (0816-
07905), Zona 1, Panama, Republica de Panama. Current
address: Florida Museum of Natural History, Dickinson
Hall, PO Box 117800, University of Florida, Gainesville,
FL 32611-7800, USA. E-mail: .


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Rylands, A. B., Schneider, H., Langguth, A., Mittermeier,
R. A., Groves, C. P. and Rodrfguez-Luna, E. 2000. An
assessment of the diversity of New World primates. Neo-
trop. Primates 8(2): 61-93.
Sherman, P. T. 1991. Harpy eagle predation on a red howler
monkey. Folia Primatol. 56: 53-56.
Suirez, J. 0. 1981. Hombres y Ecologia en Panamd. Ed.
Universitaria, Smithsonian Tropical Research Institute,
Terborgh, J. 1983. Five New World Primates: A Study in
Comparative Ecology. Princeton University Press, Prince-
ton, NJ.


Hugo Medeiros Garrido de Paula, Renata Souza Tdvora
Marcos Vinicius de Almeida, Larissa Sbeghen Pelegrini
Graziela Valenca da Silva, Rosdngela Lopes Zaganini
Anderson Lucindo


0 municipio de Bauru localiza-se na regiao centro-oeste do
Estado de Sao Paulo (Fig. 1, painel superior) e apresenta
67.937 hectares de drea, sendo 5,54% da superficie coberta
por vegetagao native. A maioria dessa vegetagao esti con-
centrada numa unidade de conservacao chamada Area de
Protecao Ambiental (APA) Vargem Limpa Campo Novo,
localizadana regiao leste do municipio (22o20'S, 49001'W),
e rodeada pelo Jardim Botanico e Zool6gico Municipal de
Bauru e por dreas de ocupacao rural e urbana (Fig. 1, painel
inferior). A vegetacao da APA foi extensivamente estudada
por Cavassan e colaboradores (1984), que descreveram-na
como sendo tfpica de cerradao, com manchas de cerrado
senso restrito e de floresta subtropical semi-decfdua (Ca-
vassan, 1990). 0 clima e predominantemente tropical,
com uma estagao seca entire margo e outubro (na qual a

Figura 1. Ilustragoes relatives a area de estudo. Painel superior: lo-
calizayao do municipio de Bauru no Estado de Sao Paulo, Brasil.
Painel inferior: area de protegao ambiental (APA) municipal
Vargem Limpa-Campo Novo.

umidade relative do ar pode chegar a 15%) e um perfodo
quente e imido estendendo-se pelos meses de novembro a

Ao long dos iltimos anos, surgiram numerosos relatos
sobre a presenga de sagiiis nas proximidades da drea em
questao. Vale destacar que pouqufssimos levantamentos
sobre a fauna do local foram realizados, sendo inexistente
qualquer mencao na literature sobre a presenga de sagiiis na
drea. Por esse motivo, o objetivo do present trabalho foi
realizar um levantamento preliminary da presenga de calitri-
quideos na APA e suas adjacencias, com vistas a contribuir
para o mapeamento das especies de primatas no territ6rio
sul-americano pelo projeto BDGEOPRIM (Hirsch et al.,

Neotropical Primates 13(3), December 2005

2002). Alim disso, foram estudados alguns aspects das in-
teraqoes dos calitriquideos com diversos fatores locals, entire
eles corn as populao6es humans que habitam a Area. Por
fim, foi feita a dissecaqao de tries individuos encontrados
mortos, corn a finalidade de extrair informaqoes biol6gicas
sobre os animals viventes no local.


Estudo 1: Levantamento preliminary dos grupos
Como primeira abordagem, 16 entrevistas foram realizadas
no mes de abril de 2002 corn pessoas residents ou que
trabalham em bairros distribufdos ao long da Rodovia
SP/225 Comandante Joao Ribeiro de Barros, a qual de-
limita a parte norte da APA (ver Fig. 1). A escolha dos
individuos a serem entrevistados seguiu uma amostragem
oportunistica, no sentido de que se buscou incluir indivi-
duos que, por alguma indicaqao, teriam avistado sagiiis. Os
questionarios utilizados foram preparados de tal forma a re-
quisitar uma certa descrigao dos animals, corn a finalidade
de se evitar falsos relatos. Em seguida, ocorreu uma etapa
de busca de registros in loco da presenca dos animals, corn
base nos dados levantados pelos questionarios. Observacoes
diretas dos sagiiis, bem como registros indiretos (detecqao
de marcas de gomivoria e a identificaqao de vocalizaqoes
tfpicas das esp&cies), foram recursos utilizados nessa etapa.
Entre marso e junho de 2004, realizamos 18 novas entre-
vistas na mesma regiao corn a finalidade de atualizar infor-
maqoes quanto ao ndmero, composicao e distribuicao de
cada grupo. Os dados de todas as etapas foram reunidos e,
corn a ajuda de mapas e fotos areas da regiao, foi feita uma
plotagem dos bandos identificados no present estudo.

Estudo 2: Aspectos ecoldgicos e comportamentais de um grupo
local de Callithrix jacchus
Apenas um grupo de sagiiis da especie ( .- jacchus
foi escolhido para ser estudado quanto a aspects ecol6-
gicos e comportamentais, tais como Area de vivencia, hAi-
bitos alimentares e perfodos de atividade. Esse grupo se
situava na periferia da Area de Reserva Legal da UNESP,
pr6xima ao Jardim Zool6gico Municipal e margeando a
Rodovia SP/225 (Fig. 2). A regiao apresenta pouco decli-
ve e terreno de latossolo textura arenosa, corn vegetadao
caracteristica de cerradao, sendo que no lado oposto ao
da rodovia ha um pequeno alagadico corn a nascente do
c6rrego Vargem Limpa. Tal Area foi dividida em seis qua-
drantes, de 50 x 50 metros cada, totalizando uma Area de
aproximadamente 1,42 hectares (Fig. 2, painel inferior).
Foi realizado o levantamento floristico dentro dos qua-
drantes utilizando-se o m&todo de parcelas, no qual as
Arvores amostradas deveriam center mais de 0,3 metros
de diametro na altura de 1,5 metros (Miiller-Dombois
e Ellemberg, 1974). Os exemplares selecionados tive-
ram seus ramos coletados, marcados e herborizados para
confirmaqao em chave especifica da regiao e comparaqao
corn exemplares existentes no herbArio do Departamento
de Ciencias Biol6gicas da UNESP de Bauru. A presenca
de marcas de mordidas de sagiiis era anotada para cada

exemplar identificado. Durante o perfodo de atividade
dos animals, das 06:00 as 18:00 horas, os quadrantes
eram visitados para registros cursivos ad libitum de com-
portamentos obtidos por observacao direta.

Estudo 3: Relaroes entire calitriquideos e series humans
Os questionArios aplicados no Estudo 1 continham per-
guntas sobre como os humans que habitavam as Areas ex-
ploradas pelos sagiiis se relacionavam corn estes animals.
Foram feitas quest6es sobre o oferecimento de comida e
sobre possiveis incomodos causados pela presenca dos ani-
mais. Tambem foi perguntado quais os perfodos (tanto do
dia como do ano) em que a presenca dos sagiiis nas proxi-
midades das Areas urbanas era mais freqiiente, e sobre pos-
sfveis ameacas aos sagiiis.

Estudo 4: Andlise post-mortem de trs sagiiis
Entre 2001 e 2004, tr&es sagiiis foram encontrados mortos
nas imediao6es da APA e encaminhados ao Departamen-
to de Ciencias Biol6gicas da UNESP / Bauru, onde foram
mantidos congelados para posterior analise post-mortem.
Nas dissecaq6es, inicialmente foram investigadas as possf-
veis causes de morte, por procura de les6es externas e inter-
nas. 0 trato digest6rio foi tambem dissecado para fornecer
informaqoes sobre o conteddo alimentar e presenca de pa-
rasitas. A determinaqao do estAgio de desenvolvimento dos
indivfduos foi feita pela pesagem e medicao do corpo, bern
como pela analise macrosc6pica dos 6rgaos genitals.


Estudo 1: Levantamento preliminary dos grupos
Identificou-se a presenca de 13 grupos de calitriqufdeos,
distribufdos na APA Municipal e em suas adjacencias,
abrangendo uma Area total de aproximadamente 1.600 hec-
tares e totalizando 150 indivfduos. Sobre os grupos, nao foi
possivel determinar corn certeza se todos eram constitufdos
por ndcleos familiares, ou se seriam meros agrupamentos
de animals na Area sem uma organizacao verdadeiramente
grupal e/ou familiar. De qualquer forma, cada grupo foi
nomeado por uma letra, indo de A a M, como esta mos-
trado naTabela 1. Essa tabela tambem exp6e a composicao
e o ndmero de indivfduos de cada grupo. Cinco grupos
(B, D, E, F e G) foram compostos por exemplares de duas
especies: ( .-' jacchus e ( .- penicillata, e por
isso foram designados como grupos mistos. Outros grupos
considerados mistos, aparentemente, foram compostos por
subgrupos monoespecificos distintos. Foi o caso de grupo
A, composto por um casal de adults C. penicillatae por um
subgrupo de C. jacchus que eram observados na mesma Area
de vivencia. 0 grupo M foi um caso semelhante, em que
dois pequenos grupos de C. penicillata viviam no mesmo
territ6rio de um grupo de C. jacchus. Os demais grupos
foram monoespecificos e, corn exceqao do grupo J, eram
formados por ndcleos familiares, isto e, compostos por um
casal de adults reprodutivos e seus descendentes jovens e
filhotes. 0 grupo J pareceu reunir mais de um ndcleo fami-
liar devido ao elevado ndmero de indivfduos.

Neotropical Primates 13(3), December 2005

A localizacao de cada grupo estA plotada na Fig. 2 (painel
superior), mostrando que os grupos de sagiiis se concen-
tram nas areas de vegetacao native, mas tambdm ocupam
manchas de vegetacao incrustradas em Areas urbanas.

Estudo 2: Aspectos ecoldgicos e comportamentais de um grupo
local de Callithrix jacchus
O subgrupo de C. jacchus pertencente ao grupo M, e cons-
titufdo por seis adults, dois subadultos, dois juvenis e um
infante, foi observado por um perfodo de tempo mais pro-
longado (15 horas acumuladas de observacao direta ao long
de tries meses) com o objetivo de permitir a caracterizadao
de seus aspects comportamentais e ecol6gicos. Conside-
rando todos os locais em que os animals foram avistados,

chegou-se a Area de vivencia do grupo correspondent a 2,4
hectares, representada na Fig. 2 (painel inferior).

Os resultados relatives a presenca dos animals em cada qua-
drante estao mostrados na Tabela 2. A Arvore-dormit6rio
esta localizada no quadrante 1 (QI), onde os animals foram
avistados is 06:00 horas deixando esse local e se locomoven-
do silenciosamente para o quadrante QII, onde era muito
comum ve-los perfurando arvores por volta das 07:00
horas. Em seguida, os animals locomoviam-se rapidamente
pelos quadrantes QIII, IV e V emitindo vocalizaqoes do
tipo longphee tais como as descritas por Epple (1968). Por
volta das 11:00 horas, os sagiiis permaneciam no quadran-
te QVI tomando sol ou engajados em comportamentos

Figura 2. Mapas localizando as areas de vivencia dos sagtiis na APA Vargem Limpa Campo Novo e seus arredores. Painel superior: As
letras "A" a "M" indicam os locals de permanencia de cada grupo descrito na Tabela 1. Painel inferior: Fotografia area mostrando os
limits da area de vivencia do grupo de Callithrixjacchus descrito no Estudo 2. Os ndmeros I a VI indicam os quadrantes utilizados na
divisao da area.

Neotropical Primates 13(3), December 2005

socials do tipo brincadeira e catagao. Na parte da tarde, eles
foram mais freqiientemente avistados nos quadrantes QVI,
IV e II, quando entao retornavam para a drvore-dormit6rio
ao por-do-sol.

As esp&cies arb6reas mais comumente amostradas foram
Vochysia tucanorum, Qualea multiflora e Qualea, r.- .
possuindo abundancias variaveis por quadrante. Em toda a
Area de vivencia, 76 Arvores apresentaram marcas de denta-
das dos sagiiis nos troncos e nos galhos, sendo que as tries
esp&cies supracitadas foram acometidas pela gomivoria.

Tabela 1. Ndmero estimado de individuos de Callithrixpenicillata
e Callithrixjacchus em cada grupo (de "A" a "M") localizado na
Figura 2.
Refer&ncia Nimero de
no mapa Espcie individuos (n)
A Callithrixpenicillata 2
Callithrixjacchus 3-4

B C. penicillata 12 20
C. jacchus
C C. penicillata 4- 6

D C. penicillata 6 10
C. jacchus
E C. penicillata 20
C. jacchus
F C. penicillata 10
C. jacchus
G C. penicillata 16
C. jacchus
H C. jacchus 5
I C. jacchus 6
J C. penicillata 24
K C. jacchus 7
L C. jacchus 8
M C. penicillata 9
C. jacchus 11

Estudo 3: Relaqres entire calitriquideos e series humans
Como a aplicaqao dos questionArios seguiu uma amostra-
gem que visava reunir o mAximo de relatos sobre a presenga
dos sagiiis na Area, todas as pessoas entrevistadas afirmaram
ver os animals com certa freqiencia. A maioria (55%) de-
clarou avistar os sagiiis pr6ximos a Areas residenciais e o
restante (45%) relatou avista-los apenas em locals de mata
fechada e distant de areas urbanas. Nao houve um perf-
odo preferencial de aparecimento dos sagiiis, sendo que
50% dos entrevistados visualizaram esses animals tanto nos
perfodos da manha como da tarde, enquanto que 27,8%
avistaram apenas no perfodo da manha, e 22,2% somente
durante a tarde. Nao houve tambem alguma 6poca do ano
mais propicia a visualizacao dos animals, ja que cerca de
50% dos entrevistados afirmam ver os sagiiis ao long de
todo o ano. Vale mencionar, entretanto, que apenas 5,5%
dos entrevistados relataram avistar os animals durante dias
mais frios de outono e de inverno.

A grande maioria dos entrevistados (94,5%) relatou gostar
da presenga destes animals nas proximidades das regi6es
urbanas. Cerca de metade das pessoas (53,8%) declararam
que fornecem algum tipo de alimento aos sagiiis, como ba-
nanas e mamao, colocadas em comedouros improvisados,
ou em outros locals elevados do chao. 0 restante, relative a
46,2% dos entrevistados, nao oferece nenhum tipo de ali-
mentagao. Apesar da percepcao favoravel em relagao a exis-
tdncia dos calitriquideos nas imediac6es de suas habitac6es,
11,1% chegou a notar algum tipo de ameaga humana aos
sagiiis, principalmente em relagao a caga, ja que a presenga
de predadores naturals nao foi identificada por nenhuma
das pessoas que vivem no local.

Estudo 4: Andlise post-mortem de trs sagiiis
O primeiro indivfduo submetido a necropsia foi uma
femea de ( .-'. penicillata pesando 260 gramas e corn
idade estimada de 10 a 12 meses. Ela apresentava les6es na
regiao inguinal da pele, costelas quebradas do lado direito e
pulmao dilacerado. Juntamente corn o fato de que o animal

Tabela 2. Dados de permanencia e de gomivoria relatives as observag6es de um grupo de Callithrixjacchus (descrito no Estudo 2) discri-
minado por quadrantes da area de vivencia.
aaa Nimero de Perodo do dia Total de Arvores corn
Quadrantes imetosodo do dia goivoria Arvores mais comuns por quadrante

I 1 6h 7h 25 Vochysia tucanorum
QualeaS .,...'. -W ./

6h 9h Vochysia tucanorum
II 10 14h 17h 14 Qualeal, ,.S ,
Qualea multif ora
Vochysia tucanorum
III 5 6h 9h 8 Qualea .,...., >.,.
Qualea multifora

IV 6 7h llh 12 Vochysia tucanorum
14h 17h Qualea ...t.. i ./.r

V 1 8h 9h 5 Vochysia tucanorum
QualeaS .,.*. ,.:,. .,
8h 12h Vochysia tucanorum
14h 17h Qualea .,.*,! /,-

Neotropical Primates 13(3), December 2005

fora encontrado ao lado de uma estrada, esses achados su-
gerem que o sagiii tenha morrido atropelado. 0 segundo
indivfduo era um ( .-'. jacchus macho, que pesava
240 gramas e apresentava idade estimada de 6 a 10 meses,
ja que os testiculos ainda nao haviam migrado para a bolsa
escrotal. Este animal foi encontrado num bosque dentro
do Campus da UNESP de Bauru sem les6es aparen-
tes, e a causa da morte nao foi determinada. 0 terceiro
sagiii era um ( .-'. jacchus femea adulto (corn mais de
12 meses de idade), e que pesava 280 gramas. Ele apresen-
tava crInio fraturado e provavelmente tambem morreu por

O conteddo estomacal dos animals apresentava uma grande
massa esbranquigada, que foi identificada como goma ve-
getal dilufda. Outros materials vegetais tambem foram en-
contrados, como pedagos de folhas, gravetos e sementes.
Havia ainda parties corporals de artr6podes, como patas,
asas e antenas. Essas parties foram identificadas como per-
tencentes a aracnfdeos, cole6pteros, ort6pteros, himen6p-
teros, hom6pteros e lepid6pteros. No intestino, o conteddo
era demasiadamente modificado para ser analisado, mas o
que merece destaque na analise desse compartimento di-
gest6rio foi a presenca de endoparasitas: uma media de
80 nemat6ides encontrada em cada individuo, principal-
mente na porgao do ceco intestinal.


Nas condio6es do present trabalho, que correspondeu a
primeira tentative de se estudar a presenca de sagiiis na
regiao de Bauru, Sao Paulo, foi contabilizada a presenca de
13 grupos de ( .-.'- jacchus e ( .-.'- penicillata na
regiao da APA Vargem Limpa Campo Novo. Admite-se
a possibilidade de que esse ndmero seja maior se for con-
siderado que vArios grupos familiares podem estar incluf-
dos num dnico grupo avistado pelos observadores. Parece
pouco provivel, contudo, que esse ndmero esteja super-
estimado, dado a amplidao da Area estudada e o tamanho
da Area de vivencia dos grupos, que num caso particular
mostrou-se relativamente pequena: 2,4 hectares para o
grupo de 11 individuos descrito no Estudo 2. Areas de
vivencias desse porte ja foram relatadas anteriormente na
literature (Digby e Barreto, 1993).

A maioria dos grupos de C. jacchus e C. penicillata possui
distribuigao marcadamente associada com a APA, o que
pode ser explicado pela abundancia de recursos alimentares
oferecida pela vegetacao local, como a presenca das Arvores
gomiferas Vochysia sp. e Qualea sp. (Cavassan, 1990). Se-
gundo Vilela e Faria (2002), o exsudato vegetal na forma de
goma constitui o principal recurso alimentar dessas especies
de sagiiis em qualquer 6poca do ano ou mesmo em ambien-
tes ricos em outros itens alimentares. No caso dos sagiiis de
Bauru, corrobora para essa ideia o resultado da analise do
conteddo alimentar que revelou que a goma vegetal e inten-
samente ingerida pelos animals. Contudo, a mesma anali-
se mostrou tambem o consume de outros itens de origem

vegetal e animal. Devido a esses dados, e possivel interpreter
que do ponto de vista alimentar, os sagiiis encontram con-
dio6es propicias para sua proliferaqao na Area da APA e ime-
diao6es. Soma-se a isso o fato dos animals se concentrarem
em locals relativamente protegidos da ocupacao humana e
com baixo nfvel de exposicao a predadores naturals, sendo
ameagados apenas por serpentes e gavi6es pouco presents
no local. Dessa forma, e possivel concluir que a populadao
de calitriquideos da Area deve aumentar rapidamente. As
conseqiiencias desse crescimento constituem mattria a ser
investigada, uma vez que esses animals podem ameagar o
equilfbrio de especies vegetais e animals nativos.

Outro achado interessante foi a presenca de sagiiis nas pro-
ximidades de Areas urbanas. Observou-se que esses grupos
de calitriquideos aproximam-se de residencias a procura de
comida, que na maioria dos casos e oferecida pelas pessoas
em comedouros. Desse tipo de interaqao surge a questao
da dependencia que estes animals podem desenvolver pelos
alimentos oferecidos pelos humans, uma vez que as frutas
constituem fontes atrativas e facilmente acessiveis e podem
perturbar padres naturals de forrageamento. Outra ques-
tao e a proximidade continue, que pode levar os sagiiis a se
habituarem aos humans, tornando-os mais vulnerAveis a
caca (como relatado por alguns entrevistados). Por fim, a
presenca em Areas urbanizadas aumenta o risco desses pri-
matas sofrerem atropelamentos, como ja documentado no
present trabalho.

Considerando a outra parte da interaqao, ou seja, a parte
humana, sabe-se que os sagiiis podem transmitir indmeras
doencas infecciosas, como raiva, atraves do contato fisico
(Favoretto et al., 2001). Sendo assim, torna-se necessirio
avaliar o risco de acidentes envolvendo humans e sagiiis.
A possibilidade de transmissao de doenyas, como parasitoses
intestinais, constitui mais um ponto a merecer atengao, es-
pecialmente porque as Areas urbanizadas freqiientadas pelos
sagiiis na region apresentam precaria infra-estrutura saniti-
ria e criancas foram comumente observadas brincando no

A ocorrencia de C. jacchus e C. penicillata na regiao se deve
provavelmente a introducao antr6pica. JA ha vArios anos,
estes animals tem sido repetidamente introduzidos na
Regiao Sudeste do Brasil, com enfase para os estados do Rio
de Janeiro e de Sao Paulo (Mittermeier et al., 1982). Co-
nhecidas por apresentarem grande adaptabilidade e capaci-
dade de explorer Areas perturbadas (Stevenson e Rylands,
1988; Faria, 1989), estas esp&cies tem se espalhado por
diferentes regi6es do territ6rio brasileiro, chegando ate o
extremo sul (Alexandre de Menezes, comunicaqao pessoal).
Finalmente, o present trabalho consiste em mais um relato
de ocorrencia de primatas que visa contribuir com o proje-
to BDGEOPRIM de localizaqao de primatas neotropicais
(Hirsch et al., 2002).

Agradecimentos: Agradecemos ao Prof. Dr. Osmar Cavassan
por estimular os estudos na Area da Reserva da UNESP, e ao

Neotropical Primates 13(3), December 2005

botInico Luiz Carlos de Almeida Neto por incentivar os es-
tudos na Area do Jardim BotInico de Bauru. Agradecemos
ao DAE pelo fornecimento dos mapas.

Hugo Medeiros Garrido de Paula, Renata Souza Tivora,
Marcos Vinicius de Almeida, Larissa Sbeghen Pelegrini,
Graziela Valenya da Silva, Rosingela Lopes Zaganini e
Anderson Lucindo, Departamento de Ciencias Biol6gi-
cas, Faculdade de Ciencias, UNESP-Campus de Bauru,
Av. Eng. Luiz Edmundo Carrijo Coube, 14-01, CEP:
17033-360, Bauru, Sao Paulo, Brasil. E-mail: fc.unesp.br>.


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genus ( .-'. Em: Ecology and Behavior of Neotrop-
ical Primates, R. A. Mittermeier, A. B. Rylands, A. F.
Coimbra-Filho e G. A. B. da Fonseca (eds.), pp.131-222.
World Wildlife Fund, Washington DC.
Vilela, S. L. and Faria, D. S. 2002. Dieta do .-
penicillata (Primates, Callithrichidae) em Areas de cerra-
do no Distrito Federal, Brasil. Neotrop. Primates 10(1):


Diego E Cisneros-Heredia
Andres Ledn-Reyes
Sylvia Seger

Although it is thought that predation has played a major
role in the evolution of primate sociality, actual predation
events involving primates are rarely documented (Van
Schaik, 1983; Boinski et al., 2000). Birds of prey, felids,
mustelids, and snakes are known predators of Neotropi-
cal primates. Most reported attacks by Neotropical snakes
on monkeys are attributed to Boa constrictor, which feeds
on callitrichids (Saguinus) and cebids (Saimiri, Cebus,
Alouatta and Chiropotes), as well as a wide variety of small-
and medium-sized mammals (didelphids, dasypodids,
vespertilionids, molossids, procyonids, agoutids, dasyproc-
tids, echimyids, murids and sciurids), birds (falconids, cot-
ingids and formicarids), and reptiles teiidss) (Janzen, 1970;
Greene, 1983; Chapman, 1986; Trail, 1987; Henderson et
al., 1995; Martins and Oliveira, 1998; Thorstrom and Mo-
rales, 2000; Shahuano Tello et al., 2002; Perry et al., 2003;
Urbani, 2003; Ferrari et al., 2004; pers. obs.).

Here we report an instance of predation by Boa constrictor
on ( .... discolor, observed during fieldwork at the Ti-
putini Biodiversity Station (TBS), a field station located in
the Ecuadorian Amazon (0037'05"S, 7610'19"W, elev.
215 m; see Cisneros-Heredia, 2003). A total of 12 pri-
mate species have been recorded at TBS: (.,. pygmaea,
Saguinus tripartitus, Aotus vociferans, C .... discolor,
Pithecia monachus, Pithecia aequatorialis, Saimiri sciureus,
Cebus albifrons, Cebus a,//la, Alouatta seniculus, Lagothrix
lagotricha and Ateles belzebuth. This is the first report of boa
predation on monkeys of the genus Callicebus.

On 28 September 2003, at 11:30 am, we heard the calls
of at least two ( .. discolor. Following the calls, we
discovered an adult Boa constrictor (total length ca. four
meters) constricting a ( ... discolor in a tree, ap-
proximately five meters above ground. The boa was coiled
around the monkey, still shifting and squeezing. A second
monkey was about four meters from the boa at the same
height and called out once. No physical interactions were
observed between the second monkey and the boa. The boa
remained coiled around the carcass for some 45 minutes
and then took approximately one hour to swallow it.

Reducing the risk of predation has been hypothesized to be
one of the benefits of group living, and group behaviours
such as alarm calls, mobbing and counter-attacks have
been reported as primate responses to predation attempts
by snakes (Chapman, 1986; Bartecki and Heymann, 1987;
Shahuano Tello et al., 2002). During this predation event,
the only response behaviour we recorded was the calling
from the second individual (rather short, classified into the

Neotropical Primates 13(3), December 2005

second group of Robinson, 1979). The absence of other re-
sponse behaviours cannot be assumed, however, because we
arrived when the boa was already constricting the monkey.
It is unknown how predation events may have functioned
in the evolution of sociality in Callicebus, but this observa-
tion, together with similar reports (Chapman, 1986; Bar-
tecki and Heymann, 1987; Martins and Oliveira, 1998;
Shahuano Tello et al., 2002; Perry et al., 2003; Ferrari et
al., 2004), suggests that snakes play a major role as preda-
tors of Neotropical primates.

Acknowledgements: We are grateful to Eckhard W. Hey-
mann, Stella de la Torre and Kelly Swing for their critical
reading of the manuscript. Susan Perry, Eckhard W. Hey-
mann and Stella de la Torre provided relevant literature.
Maria Elena Heredia, Laura Heredia and Hector Le6n
provided financial and moral support. Tiputini Biodiver-
sity Station / Universidad San Francisco de Quito provided
institutional and logistical support. Video files of this event
are deposited in the archives of the Tiputini Biodiversity
Station, Universidad San Francisco de Quito. This is a
publication of the project "Study of the Herpetofauna of
the Tiputini Biodiversity Station" (D. F Cisneros-Heredia,

Diego F. Cisneros-Heredia, College of Biological & Envi-
ronmental Sciences, Universidad San Francisco de Quito,
Quito, Ecuador, Andras Le6n-Reyes, Tiputini Biodiversity
Station, Universidad San Francisco de Quito, Ecuador, and
Sylvia Seger, School for International Training, Hernando
de la Cruz N31-37, Quito, Ecuador. Address for correspond-
ence: Diego E Cisneros-Heredia, King's College London,
Department of Geography, Strand, London WC2R 2LS,
UK. E-mail: .


Bartecki, U. and Heymann, E. W. 1987. Field observa-
tion of snake-mobbing in a group of saddle-back tama-
rins, Saguinus fuscicollis nigrifrons. Folia Primatol. 48:
Boinski, S., Treves, A. and Chapman, C. A. 2000. A critical
evaluation of the influence of predators on primates: Ef-
fects on group travel. In: On the Move: How and Why An-
imals Travel in Groups, S. Boinski and P. A. Garber (eds.),
pp.43-72. The University of Chicago Press, Chicago.
Chapman, C. A. 1986. Boa constrictor predation and group
response in white-faced Cebus monkeys. Biotropica 18(2):
Cisneros-Heredia, D. E 2003. Herpetofauna de la Estaci6n
de Biodiversidad Tiputini, Amazonfa Ecuatoriana:
Ecologfa de una comunidad taxon6micamente diverse con
comentarios sobre metodologfas de inventario. In: Ecologia
y Ambiente en el Ecuador: Memorias I Congreso de Ecologia
y Ambiente, S. de la Torre and G. Reck (eds.), pp.1-21.
Universidad San Francisco de Quito, Ecuador.
Ferrari, S. F, Pereira, W. L. A., Santos, R. R. and Veiga,
L. M. 2004. Fatal attack of a Boa constrictor on a

bearded saki (Chiropotes satanas utahicki). Folia Primatol.
75: 111-113.
Greene, H. W. 1983. Boa constrictor (boa, bequer, boa
constrictor). In: Costa Rican Natural History, D. H.
Janzen (ed.), pp.380-382. The University of Chicago
Press, Chicago.
Henderson, R. W., Micucci, T. W. P., Puorto, G. and Bour-
geois, R. W. 1995. Ecological correlates and patterns
in the distribution of Neotropical boines (Serpentes,
Boidae): A preliminary assessment. Herpetological Natu-
ralHistory 3: 15-27.
Janzen, D. H. 1970. Altruism by coatis in the face of preda-
tion by Boa constrictor. J. Mammal. 51: 387-389.
Martins, M. and Oliveira, M. E. 1998. Natural history of
snakes in forests of the Manaus Region, Central Amazo-
nia, Brazil. Herpetological NaturalHistory 6(2): 78-150.
Perry, S., Manson, J. H., Dower, G. and Wikberg, E. 2003.
White-faced capuchins cooperate to rescue a groupmate
from a Boa constrictor. Folia Primatol. 74: 109-111.
Robinson, J. G. 1979. Vocal regulation of use of space by
groups of titi monkeys ( .... moloch. Behav. Ecol. So-
ciobiol. 5: 1-15.
Shahuano Tello, N. S., Huck, M. and Heymann, E. W.
2002. Boa constrictor attack and successful group defence
in moustached tamarins, Saguinus mystax. Folia Primatol.
73: 146-148.
Thorstrom, R., Ramos, J. D. and Morales, C. M. 2000.
Breeding biology of barred forest-falcons in northeastern
Guatemala. Auk 117: 781-786.
Trail, P. W. 1987. Predation and antipredator behavior at
Guianan Cock-of-the-Rock leks. Auk 104: 496-507.
Urbani, B. 2003. Utilizaci6n del estrato vertical por el
mono aullador de manto (Alouatta palliata, Primates) en
Isla Col6n, Panama. Antropo 4: 29-33.
Van Schaik, C. P 1983. Why are diurnal primates living in
groups? Behaviour 88: 120-143.


Gabriela Ludwig
Lucas M. Aguiar
VlamirJ. Rocha


Na porcao sul e parte da porcao sudeste do Brasil, Cebus
nigritus (Goldfuss, 1809) e a esp&cie de macaco-prego ca-
racteristica da Mata Atlantica. Ao norte, sua distribuicao
limita-se a margem esquerda do Rio Doce (Silva Jdnior,
2001; Vilanova et al., 2005) e ao sul limita-se as municipa-
lidades de Sao Lourenco do Sul, Rio Grande do Sul (Printes
et al., 2001). A leste, a distribuicao e limitada pelo oceano
Atlantico e a oeste pelo Rio Parana (Silva Jdnior, 2001; Vi-
lanova et al., 2005).

Neotropical Primates 13(3), December 2005

Esta especie de macaco-prego esta entire os mamiferos mais
facilmente encontrados em fragments florestais do norte
do estado do Parana (Rocha, 2001), cujo patrim6nio natu-
ral vem sofrendo intense devastacao por atividades agricolas
e agropecuarias que dizimaram a mata native dessa regiao,
reduzindo a floresta original a valores pr6ximos de 1% a
2% (Parand, 1987). Cebus nigritus e uma especie onfvora
com grande adaptabilidade aos ambientes alterados pelo
home e pode sobreviver em Areas de florestas fragmenta-
das e degradadas, desde que tenham acesso a outras fontes
alimentares, como as plantaq6es ao redor de seu ambiente
(Rocha, 2000). E justamente nessa situacao que a especie
e vista como uma praga florestal por produtores rurais,
pois invadem planta6es e passam a consumer pomares,
milharais, canaviais, plantios de mandioca e ate mesmo
plantac6es de Pinus (Rocha, 2000; Ludwig et al., no prelo).
Assim, esse lamentavel conflito pode futuramente alterar o
status de conservacao desses primatas em estados agricolas
como o Parana.

Os poucos trabalhos que abordaram o comportamento e a
ecologia dessa especie em ambiente natural na regiao mos-
traram flexibilidade comportamental e estrategias ecol6gi-
cas oportunistas para que os animals conseguissem sobre-
viver ao confinamento de fragments pequenos e isolados
(Rocha, 1995, 2001; Rocha et al., 1998, no prelo; Ludwig
et al., no prelo). Portanto, visando a future conservadao
da especie e um melhor entendimento da adaptabilidade
desses animals frente aos efeitos antr6picos, surgiu o prop6-
sito deste trabalho que objetivou verificar e analisar a dieta,
a drea de vida, os percursos diarios e as estimativas popula-
cionais de C. nigritus no fragmento florestal Mata Doralice,
Ibipora, Parani.

Material e Metodos

Area de estudo
A Mata Doralice (2316'S, 5103'W) situa-se no muni-
cfpio de Ibipora, norte do estado do Parand, na porcao
baixa da bacia do Rio Tibagi, a 484 m de altitude. E um

fragmento florestal de 170 ha, coberto em sua maior part
por uma vegetacao primAria alterada do tipo Floresta Es-
tacional Semidecidual. 0 fragmento limita-se ao sul com
o Rio Tibagi e esta circundado por plantios de monocul-
turas, pomares e pastagens (Fig. 1). 0 solo da regiao e
classificado segundo o Sistema Brasileiro de Classificaqao
de Solos (EMBRAPA, 1999) como Nitossolo Vermelho
Eutr6fico (terra roxa) e caracteriza-se por ter alta fertili-
dade natural. 0 clima e subtropical 6mido, apresentando
as quatro estaq6es bem definidas. As medias anuais para
temperature e precipitacao sao 21,8C e 1558 mm, res-
pectivamente (dados obtidos em Soares-Silva et al., 1992;
Carmo, 1995). Estudos fitossociol6gicos na drea apon-
taram densidade absolute de 1396 individuos/ha, diver-
sidade especifica de H' = 3,6 (I = 0,786), e as families
mais representatives sendo Meliaceae, Myrtaceae, Faba-
ceae, Euphorbiaceae, Moraceae e Mimosaceae. A floresta
e constitufda por um estrato arbustivo e dois arb6reos,
alum de arvores emergentes com alturas superiores a 20 m
(Soares-Silva et al., 1992; Carmo, 1995).

Os dados foram coletados entire outubro de 2001 e setem-
bro de 2002, atraves de quatro visits mensais a campo di-
vidas igualmente entire o perfodo matutino e vespertino.
Foram acompanhados dois grupos focais (GA: 35 indivfdu-
os e GB: 25 individuos; 155 e 25 h de acompanhamento,
respectivamente); os dados de dieta foram coletados atrav5s
do m&todo de frequencias de observacao (atraves de ob-
servagao direta dos animals) e freqiuencia de ocorrencias
dos itens alimentares em fezes. Para padronizar as amostras
fecais e as visuais na somat6ria final, considerou-se uma
ocorrencia tanto uma especie encontrada nas fezes como
cada especie ingerida pelo animal na observagao direta
do grupo (Rocha, 2001). Itens vegetais nao identificados
foram mencionados como indeterminados (sp. 1, sp. 2,
etc.). Fragmentos de parties animals (artr6podos) encontra-
dos nas fezes foram identificados pelo laborat6rio de ento-
mologia da Universidade Estadual de Londrina, Londrina,

Figura 1. Mapa do fragmento florestal Mata Doralice (170 ha), Ibipora, Parand.

Neotropical Primates 13(3), December 2005

Todas as porcentagens finals obtidas dos itens que fazem
parte da dieta foram obtidos atraves da somat6ria do total
de itens vegetais (descritos na Tabela 1) e do total de itens
animals (Tabela 2), totalizando em 100% (e. g., 769 + 226
= 100%). Assim como as anilises sazonais: I itens vegetais
+ Y itens animals de cada estagao = 100%. (P. ex., a porcen-
tagem de Maclura tinctoria na primavera e de: (18 + 23) x
100% / (29 + 61 + 108) + (4 + 27 + 13) = 16,9%.)

Para o estudo da area de vida, GA foi acompanhado por
180 h. 0 mapeamento da area de vida foi feito atraves da
plotagem dos pontos de GPS (Garmin, modelo eTrex Ven-
ture). Os pontos e rotas foram marcados pelo aparelho
conforme a atividade dos animals, mudangas de diregoes



Figura 2. FreqUiencia total (%) dos itens alimentares (especies veg-
etais e artr6podos) consumidos por Cebus nigritus.

do grupo e possibilidade de contato corn satdlites dentro da
mata (Aguiar et al., 2002). A Area foi calculada atraves do
program AutoCAD 2000 (Autodesk, 1999). Alguns per-
cursos diArios foram possiveis de serem mensurados pelo
pr6prio GPS.

Para estimar a densidade da espucie no fragmento foi em-
pregado o m&todo da Area de vida (adaptado de Brockel-
man e All, 1987), levando-se em conta o nimero medio
de individuos encontrados por grupo ao long do estudo
e tanto a dimensao total da Area de vida quanto a de uso
exclusive do grupo.


A dieta constituiu de itens vegetais e animals, registrados
em 995 ocorrencias sendo a porcentagem de frutos (67,4%)
substancialmente mais elevada que as dos demais itens ve-
getais (sementes, 3,6%; flores, 2,1%; n&ctar, 1,1%; folhas,
0,8%; meristema, 0,7%; raiz, 0,7%; caule, 0,5%; brotos,
0,4%) e animals (22,7%) (Fig. 2).

Quanto aos itens vegetais (77,3%) foram registradas 73 es-
p$cies (alum do genero Ficus), incluindo seis ex6ticas (Zea
mays, Citrus sp. 1, Citrus sp. 2, Hovenia dulcis, Caryota
urens e Musa paradisiaca), totalizando 32 families identifica-
das (Tabela 1). A familiar Meliaceae foi a que obteve maior

Tabela 1. Especies vegetais, item consumido, ndmero de vezes que o item foi consumido sazonalmente e porcentagem total que apareceu
na dieta de Cebus nigritus. IC = Item Consumido; I itens vegetais + Y itens animals = 100%. Uma tabela aumentada, corn informao6es
detalhadas sobre os itens menos comuns, pode ser encontrada no sitio de internet do Neotropical Primates em org/np/LudwigTabelal a.pdf>.
Primavera Verao Outono Inverno
Espdcie IC I %
________ __ ON D J F MA M J J A S _
Sorocea bonplandii Frutos 7 1 9 37 57 6 117 11,8
Euterpe edulis Frutos 1 8 7 7 21 20 14 78 7,8
Zea mays Frutos 2 4 2 5 10 5 4 2 8 13 55 5,5
Pereskia aculeata Frutos 3 2 16 23 5 3 52 5,2
Guarea kunthiana Frutos 1 3 1 2 1 1 13 23 4 49 4,9
Maclura tinctoria Frutos 18 23 1 1 1 44 4,4
Syagrus : Frutos 8 6 7 4 3 4 32 3,2
Miconiapusilliflora Frutos 5 20 5 30 3,0
Piper amalago Frutos 5 6 3 3 1 18 1,8
Ficusspp. Frutos 3 4 2 5 2 1 17 1,7
Piper aduncum Frutos 1 8 4 13 1,3
Nectandra megapotamica Frutos 12 12 1,2
Psidium guayava Frutos 11 1 12 1,2
Cabralea canjerana Sementes 3 9 12 1,2
Piper crassinervium Frutos 9 1 1 11 1,1
Jacaratia spinosa Frutos 3 6 2 11 1,1
Macfadyena dentata Sementes 10 1 11 1,1
Indeterminada sp. 3 Frutos 1 9 10 1,0
Euterpe edulis Flores 4 1 5 10 1,0
Outros (67 itens) 17,5
Total ____29 61 108 75 61 37 66 35 62 99 107 29 769 77,3

Neotropical Primates 13(3), December 2005

ndmero de especies consumidas, seis no total. A familiar
Moraceae (17,9%) destacou-se na dieta principalmente pela
utilizacao dos frutos de Sorocea bonplandii (11,8%). A Are-
caceae e a Poaceae foram freqiientes na dieta, representadas
pelo consumo de Euterpe edulis, palmito (9,5%, sendo 7,8%
de fruto; 1,0% de flor; 0,7% de meristema) e Zea mays,
milho (5,5%), respectivamente. Especies como Guarea kun-
thiana, Ficus spp. e Z mays estiveram presents em todas as
estaq6es do ano (Tabela 1). Esta 6ltima espucie foi coletada,
juntamente com Manihot esculenta mandiocaa), pelos ani-
mais nas plantaq6es localizadas ao redor da mata.

P6de-se constatar que durante a primavera e o verao ocor-
reu maior riqueza de especies vegetais quando comparada
ao outono e ao inverno. Na primavera, os itens vegetais mais
consumidos foram frutos de Maclura tinctoria (16,9%) e
Miconia, p',,'..j,,. (12,4%); no verao, foram os frutos de
Euterpe edulis (10,8%) e de Zea mays (9,8%). No outono,
novamente prevaleceram os frutos do palmito (28,2%), se-
guidos dos frutos de Pereskia aculeata (10,7%). Ressalta-se
que nessas duas 6ltimas estao6es os animals foram regis-
trados consumindo o meristema de palmito (0,7%). No
inverno, predominaram os frutos de Sorocea bonplandii
(28,2%) e Guarea kunthiana (11,3%). Nas duas 6ltimas
estaq6es, observou-se individuos visitando os plantios de
Manihot esculenta para desenterrarem os tub&rculos e pos-
teriormente consumi-los dentro da mata.

Em relagao aos itens de origem animal (22,7% da dieta
total), foram registrados aracnfdeos (0,2%) e seis ordens de

insetos (Hemiptera: 3,4%; Hymenoptera: 3,4%; Coleopte-
ra: 1,3%; Orthoptera: 1,1%; Diptera: 0,3%; Lepidoptera:
0,2%) alum de invertebrados nao identificados (12,8%)
(Tabela 2). 0 consumo de itens animals foi mais elevado
no inverno (34%) do que nas outras estao6es do ano.

Area de vida e percursos didrios
O grupo de estudo, GA, totalizou 56 ha de Area de vida,
resultando uma exigencia espacial de 1,6 ha/indivfduo. A
Area nuclear (a Area com maior concentracao de pontos
marcados pelo GPS e de uso exclusive do grupo) foi de 14
ha e correspondeu a 25% do total de sua Area. Duas outras
sub-Areas utilizadas pelo grupo foram constatadas, uma ao
noroeste e outra a sudeste da Area de vida (Fig. 3). Foram
mensurados sete percursos didrios que variaram entire 900 e
2000 m (media de 1083 m).

Cerca de 90% da Area de vida dos animals abrangeu flo-
resta primAria alterada e somente 10% abrangeu floresta
secundAria. Foi marcante o uso da borda da mata (4000
m de borda de mata utilizada) para forrageio, descanso e
para acesso aos plantios de milho, mandioca e pomares. 0
grupo adentrou de 15 a 25 m nos plantios dessas cultures.

0 grupo focal sobrepos a periferia de sua Area de vida
com mais dois outros grupos vizinhos, verificando-se 14
ha (25%) de Area sobreposta com GB (grupo que habitat a
porgao leste da mata) e 10 ha (18%) corn GC (grupo que
habitat a porgao norte da mata). GA obteve uma Area de
uso exclusive de 32 ha (57%), na qual estava contida a Area

Tabela 2. Itens artr6podos, fragments encontrados, ndmero de vezes que o item foi consumido sazonalmente e porcentagem total em que
apareceram na dieta de Cebus nigritus. CT = Categoria taxonomica; ME = modo encontrado: fragments em fezes (Frag.) ou observagao
direta (Obs.); NI = Nao Identificado; Y itens vegetais + Y itens animals = 100%.
Primavera Verao Outono Inverno
CT ME / Animal I 0 %
Coleoptera Fragm. / Besouros 2 4 1 1 2 2 1 13 1,3
Arachnida Fragm. / Aranhas 2 2 0,2
Lepidoptera Fragm. / Indeterminados 1 1 2 0,2
Orthoptera Fragm. / Gafanhoto 2 1 1 4 2 10 1,0
Orthoptera Obs. / Gafanhoto 1 1 0,1
Hemiptera Ninfa de percevejo 1 1 0,1
Hemiptera Ninfa de cigarrinha 1 1 0,1
Hemiptera Fragm. / Cigarrinha 1 1 0,1
Hemiptera Fragm. / Percevejo 5 3 2 1 2 2 4 8 1 2 30 3,0
Hemiptera Obs. / Percevejo 1 1 0,1
Hymenoptera Fragm. / Formiga 1 5 1 1 1 2 1 1 4 3 1 21 2,1
Hymenoptera Fragm. / Indeterminados 2 2 1 5 0,5
Hymenoptera Fragm. / Vespa 1 1 1 3 6 0,6
Hymenoptera Obs. / Formiga 1 1 0,1
Hymenoptera Obs. / Vespa 1 1 0,1
Diptera Fragm. / Indeterminados 1 1 2 0,2
Diptera Pupa 1 1 0,1
Artr6podo (NI) Fragmentos 1 1 2 0,2
Artr6podo (NI) Observaygo direta 3 4 6 7 2 7 3 9 40 31 13 125 12,6
Total 4 27 13 11 10 9 9 6 17 60 38 22 226 22,7

Neotropical Primates 13(3), December 2005

nuclear. Foram observados dois encontros intergrupais: no
primeiro, GA estava pr6ximo a sua area nuclear e expul-
sou GB. Em outro, quando GA estava em porgao perifkrica
da area de vida, foi o grupo expulso por GC. Na primeira
ocasiao, verificou-se que o grupo resident formou um sub-
grupo de machos adults e sub-adultos em linha de frente
e emitiram gritos de alarme, quebravam galhos e exibiam
os caninos em direqao ao outro grupo, enquanto as femeas
com infants e os juvenis afastaram-se.

Estimativas populacionais
Foram levantados tries grupos de C. nigritus no fragmento.
O tamanho dos grupos GA e GB foi de 35 e 25 individuos,
respectivamente (media: 30 individuos/grupo). As estima-
tivas indicaram de tries a cinco grupos no local e densidade
populacional de 0,54 a 0,94 individuos/ha.


O predomfnio de algumas esp&cies vegetais na dieta dos
animals pode ter sido reflexo do tipo do ambiente utilizado,
seja porque as esp&cies foram abundantes dentro da floresta
ou abundantes em plantaq6es (Galetti e Pedroni, 1994). A
familia Meliaceae familiar de maior riqueza floristica, e de
maior densidade e frequiencia nos estudos fitossociol6gicos
da Mata Doralice foi justamente a familia de maior rique-
za de species na dieta de C. nigritus. Sorocea bonplandii,
esp&cie vegetal que apresentou o maior indice de valor de
importancia para o fragmento, se destacou entire as demais
pelo consumo de frutos. Euterpe edulis (palmito), quinta
esp&cie com maior indice de valor de importIncia e Zea
mays, abundante nos plantios durante todo o ano, tambem
foram bastante utilizadas no consumo de frutos pelos

animals deste fragmento. Essas esp&cies tambem foram
bastante consumidas por C. nigritus em outros fragmen-
tos da regiao do baixo Tibagi (Rocha, 1995, 2001), sendo
consideradas altamente importantes para esses animals no
context regional.

Sazonalmente, a maior riqueza de esp&cies vegetais con-
sumidas durante a primavera e verao pode ter refletido a
maior disponibilidade de frutos durante estes perfodos. Por
outro lado, durante o outono e o inverno, foram poucas
as esp&cies que produziram frutos que foram consumidos
pelos primatas (e.g. Sorocea bonplandii, Guarea kunthiana,
Pereskia aculeata), o que pode refletir a menor riqueza de
especies vegetais na dieta. Nesses mesmos perfodos, o con-
sumo dessas especies foi tao elevado que as tries se destaca-
ram na frequencia total obtida. Alim disso, o consumo de
tub&rculos de Manihot esculenta pode refletir a adaptadao
alimentar dos animals em perfodos de escassez (Ludwig et
al., no prelo). Durante o inverno tambem pode-se perceber
um consumo mais elevado de itens animals, o que superou
ate mesmo a frequiencia observada em relacao aos frutos
mais consumidos nessas 6pocas. Itens animals, principal-
mente insetos, tambem foram mais procurados durante as
estaqoes mais secas como foi mostrado em outros traba-
lhos com vArias especies de Cebus (Robinson, 1986; Ro-
drigues, 1992; Izar, 1999; Rocha, 2001; Spironello, 2001)
- demonstrando uma estrategia em resposta a menor dis-
ponibilidade de frutos.

Area de vida e percursos didrios
0 grupo de estudo apresentou uma Area de vida pequena
e bem definida, bastante inferior em relacao aos trabalhos
com outras esp&cies de Cebus em Areas de mata continue:
80-125 ha (C. 7.c/q.i e C. ... Terborgh, 1983), 275
ha (C. olivaceus; Robinson, 1986), 108 ha (C. capucinus;
Chapman, 1988), 240 ha (C. nigritus; Izar, 1999), 81-293
ha (C. nigritus; Di Bitetti, 2001) e 900 ha (C .cl//; Spi-
ronello, 2001). Porem, esti mais pr6ximo aos valores en-
contrados em estudos de fragmentacao na regiao norte do
estado do Parana: 50-75 ha para grupos que tambem uti-
lizaram plantios e pomares (Rocha, 1995). Os efeitos de
fragmentacao na diminuicao da Area de vida podem estar
relacionados tanto as pr6prias restrio6es de tamanho im-
postas pelos fragments, quanto tambem a capacidade de
"forrageio extra" vista para o g&nero ao aproveitarem recur-
sos externos a mata, como por exemplo, os plantios e os
pomares. Estas fontes extras de recursos podem aumentar
a densidade populacional nessas respectivas Areas e conse-
quientemente reduzir as dimens6es do espago utilizado pelos
animals em ambientes confinados. Isso ajudaria explicar a
baixa exigencia espacial encontrada neste trabalho quando
comparada aos trabalhos citados acima.

Nos casos aqui verificados, a oferta de recursos relativa-
mente frequientes como os plantios poderiam estar fun-
cionando como abundantes agregados de alimento. Estes
por sua vez, reduziriam o tamanho da Area de vida do
grupo, ja que Areas de vida deveriam ser menores se os

Figura 3. Area de vida de Cebus nigritus, GA, corn 56 ha. Area
nuclear corn 14 ha (cinza escuro) e sub-areas noroeste e sudeste
(circulos em preto). Observe os plantios de milho (Zea mays) ao
redor do fragmento.

Neotropical Primates 13(3), December 2005

Tabela 3. Tabela comparative das densidades de Cebus nigritus encontradas por diversos autores ao long da distribuicao -... .. i da
Densidade (ind/ha) Mdtodo Localidade Fonte
0,54 0,94 Area de vida e tamanho de grupos Mata Doralice, Brasil este trabalho
0,29 -0,36 Area de vida e tamanho de grupos PE Mata dos Godoy, Brasil Rocha (2001)
0,66 0,76 Transecto linear PE Vila Rica do ES, Brasil Vidolin e Mikich (2004)
0,19 0,32 Transecto linear Mata Sao Jose, Brasil Bernardo e Galetti (2005)
0,057 Transecto linear PN Iguazu, Argentina Brown e Zunino (1994)
0,474 Transecto linear RB Augusto Ruschi, Brasil Pinto etal. (1993)
0,07 Transecto linear Reserva Puriti, Brasil Chiarello (2000)
0,01 Transecto linear Reserva M7/317, Brasil Chiarello (2000)
0,035 0,053 Transecto linear Serra Paranapiacaba, Brasil Gonzalez-Solfs et al. (2001)
0,16 Area de vida e tamanho de grupos PN Iguaztl, Argentina Di Bitetti (2001)
0,51 Transecto linear Mata ciliar (PR)/Rio Parand/Brasil L. M. Aguiar, dados nao publicados

aglomerados de alimentos fossem abundantes (Pough et
al., 1999). Di Bitetti (2001) tambdm verificou o efeito dos
agregados alimentares na redugao da Area de vida em um
grupo de C. nigritus no Parque Nacional do Iguazd, ao
fornecer de modo freqilente, alimentos em plataformas
para os animals.

Aldm do uso caracteristico e freqiiente de uma Area nucle-
ar (Terborgh, 1983), a utilizagao de outras duas sub-Areas
p6de star associada ao ficil acesso destas Areas aos plantios
de Zea mays em frutificacao. Aldm disso, estas sub-areas de
acesso tambdm pareciam ser estrategicas contra a possibi-
lidade de ataque de caes domesticos, pois eram localizadas
distantes dos canis. Deste modo, essas Areas propiciavam ao
grupo adentrar aos plantios com maior seguranga.

Estimativas populacionais
As estimativas populacionais partiram da hip6tese de a
Mata Doralice ser homogenea. Todavia, e salutar ressal-
tar que esta densidade pode sofrer alteragoes se for con-
siderada a heterogeneidade da mata. A estimativa de tries
a cinco grupos pareceu estar plausivel, pois ao long do
estudo foram identificados tries grupos sem a possibilidade
de existencia de outros. Ainda, a quantidade de grupos aqui
encontrada e muito semelhante a verificada por Siemers
(2000) em um fragmento de tamanho tambdm semelhante
ao da Mata Doralice. Em relagao is densidades populacio-
nais, os resultados apresentaram-se elevados em compara-
gao is densidades estimadas por outros autores ao long
da distribuigao -, ..- i,. i da esp&cie (Tabela 3). Isso ilustra
a flexibilidade de C. nigritus em prosperar uma populagao
relativamente grande em um fragmento pequeno e estreito,
frente is imposigoes de um mosaico ambiental constitufdo
por fragments isolados, entremeados por monoculturas e

Agradecimentos: Ao Pedro Favoreto, Bidd e Ivonete pelo
apoio pessoal em campo. Ao Josd Marcelo D. Torezan,
Manoel R. C. Paiva, Edmilson Bianchini e Josd Lopes
pelo auxflio na identificaqao dos itens alimentares, e aos

comentArios de Ndlio Roberto dos Reis, Isaac P. Lima, Fer-
nando C. Passos e Josd H. E Marino.

Gabriela Ludwig e Lucas M. Aguiar, P6s-graduacao em
Zoologia, Laborat6rio de Biodiversidade, Conservacao e
Ecologia de Animais Silvestres, Departamento de Zoolo-
gia, Universidade Federal do Parana, Caixa Postal 19020,
CEP 81531-990, Curitiba, Parana, Brasil, e Vlamir J.
Rocha, Fazenda Monte Alegre s/n, Lagoa, Telemaco Borba
84279-000, Parana, Brasil. Endereqo para correspondin-
cia: Gabriela Ludwig, correio eletronico ufpr.br>.


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Helissandra Mattjie Prates
Jilio Cesar Bicca-Marques


Coprophagy, or the behavior of eating feces, is classified as
autocoprophagy when the individual eats its own feces, or
allocoprophagy when it eats the feces of others (Hirakawa,
2001; Graczyk and Cranfield, 2003). This habit is observed
in lagomorphs, rodents, marsupials, and primates. Among
leporids, coprophagy occurs in the form of caecotrophy
(the reingestion of soft feces or caecotrophs) and serves to
improve the absorption of vitamins and microbial proteins
(Hirakawa, 2001). Caecotrophy has also been observed in
a prosimian, the sportive lemur Lepilemur leucopus (Hladik,
1978). Among anthropoid primates, coprophagy has been
observed in captive and wild apes (chimpanzees, gorillas,

Neotropical Primates 13(3), December 2005

orangutans and gibbons: Hill, 1966; Gilloux et al., 1992;
Warniment and Brent, 1997; Nash et al., 1999; Faraldo
and Taylor, 2003; Graczyk and Cranfield, 2003; Krief
et al., 2004), Old World monkeys (baboons and rhesus
macaques: Brent et al., 2002; see also Graczyk and Cran-
field, 2003), and New World monkeys (marmosets, tama-
rins and capuchin monkeys: Anderson et al., 1991; Clark,
1994; Wissman, 1999; Taylor, 2002).

Krief et al. (2004) discussed a number of hypotheses to ex-
plain coprophagy by captive primates: (a) food deficiency,
(b) boredom, (c) social stress, and (d) medical problems.
The only report of coprophagy in capuchin monkeys (An-
derson et al., 1991) was of an occurrence during the inte-
gration of a tame adult female into a captive group. An-
derson et al. argued that this abnormal behavior may have
been related to food deficiency and/or social stress, since
the human-raised female used to eat feces before adjusting
to the standard primate food pellets offered in captivity.
She was also frequently involved in agonistic interactions
with other group members soon after her integration.

In this paper we report cases of coprophagy by captive
brown capuchin monkeys and examine whether they fit the
"food deficiency" and "social stress" hypotheses cited above.
We predict that if food limitation is the primary stimulus
for this behavior, coprophagy will be more frequent during
those periods in which the monkeys have no food avail-
able in the cage. On the other hand, if social pressure can
explain this behavior, we can expect to find an inverse re-
lationship between social rank and individual frequency of
coprophagy. In addition, we would expect that a given in-
dividual will be more likely to eat feces after being harassed
by other groupmates.


The social behavior of a group of 10 brown capuchin mon-
keys (Table 1) was studied over 219.5 observation hours,
from 8 April to 24 October 2003 at the Parque Farroupilha
in Porto Alegre, Rio Grande do Sul, Brazil. This urban park
has many visitors and is close to busy city streets. The study
group was kept in a wired hexagonal cage with a cement
floor, measuring approximately 5.5 m on each side with a
height of 3.5 m. For behavioral enrichment the cage was
equipped with a wheel, two tires and one movable ladder.
The monkeys were fed only once a day, at about 09:00 h,
with fruits, vegetables and sometimes peanuts. The cage
was washed with water once a week on Thursday after-
noons, so the monkeys had no food available until Friday

We recorded all occurrences of coprophagy following the
behavior sampling rule with continuous recording (Martin
and Bateson, 1993). We determined the dominance status
of each individual based on the frequency and distribution
of agonistic interactions within dyads (see Janson, 1985).
An individual was considered to have high social rank if

Table 1. Age-sex composition of the study group, social rank, and
recorded cases of autocoprophagy and allocoprophagy by each
SSocial Auto- Allo-
Age-sex class Individual S A A
Age-sex class Individual rank coprophagy coprophagy

Adult male Roger 1tr
Adult male Chico 2nd
Adult male Tiburcio 4rh
Adult male Barba 6rh 2 9
Adult male Assis 10h
Adult female Amelia 3rd 3 23
Adult female Fabi 9h 15
Juvenile male Dali 5rh 1 8
Juvenile male Guri 7rh 4 4
Infant male Fiba 8 4
Total 10 63

it was the initiator of aggression more frequently than a
recipient of aggression. In contrast, low social rank was in-
dicated by a higher frequency of aggression received than


We recorded a total of 73 events of coprophagy (0.33 events
per hour of observation). Six individuals (Amelia, Fabi,
Barba, Dali, Guri and Fiba) were observed to eat feces. The
remaining four (Roger, Chico, Tiburcio and Assis) never
demonstrated this behavior (Table 1), but were occasion-
ally seen to drink urine from the floor. Coprophagy was
more prevalent than expected in females than in males
(41 events vs. 32 events; x2 = 59.671, d.f. = 1, p < 0.001).
Allocoprophagy was more frequent than autocoprophagy.

Coprophagy was more common in the afternoon than in
the morning (47 events or 0.47 events per hour of observa-
tion vs. 26 events or 0.22 events per hour; x2 = 11.000, d.f.
= 1, p < 0.001), suggesting that food availability may play
an important role in the occurrence of this behavior. How-
ever, we recorded only nine instances of coprophagy (12%)
in which there was no food on the cage floor. The frequen-
cy of coprophagy on Thursdays (when the cage was cleaned
and devoid of food) was 0.21 events per hour, not above
the expected level based on sampling effort (14 events; x2 =
1.785, d.f. = 1, NS).

We observed a total of 326 cases of agonistic interactions
during the study (1.49 events per hour of observation).
Social rank did not explain inter-individual differences
in coprophagy (r2 = 0.002, n = 10, F-ratio = 0.016, p =
0.901). For example, the two highest-ranking individu-
als (Roger and Chico) and the lowest-ranking individual
(Assis) never ate feces, whereas Amelia and Fabi (ranked
third and ninth, respectively) showed the highest frequen-
cies of coprophagy (Table 1). In addition, only rarely had
the individual observed eating feces been harassed earlier

Neotropical Primates 13(3), December 2005

in the day (Barba: two cases; Fabi: three cases; and Guri:
two cases). Therefore, the "social stress" hypothesis was not


We observed a high frequency of coprophagy in this group
of captive brown capuchin monkeys, a behavior reported
only once before in this species (Anderson et al., 1991).
In a similar study on the social behavior of a captive
group of nine capuchin monkeys at the Parque Zool6gico
de Sapucaia do Sul, Rio Grande do Sul, Brazil, Daniel
B. Montano did not observe a single case of coprophagy
during 208 hours of observation over 16 months (pers.

Its prevalence in adult females and immature individuals
may be related to a diet insufficient in protein, as described
for marmosets by Flurer and Zucker (1988). The tendency
of females and immatures to exploit diets richer in protein
has been described for a number of primates, and is relat-
ed to the nutritional demands of gestation, lactation, and
growth (see Bicca-Marques and Calegaro-Marques, 1994).
This tendency, coupled with the fact that coprophagy was
more frequent in the afternoon, supports the "food de-
ficiency" hypothesis. However, most cases of coprophagy
were observed when there was still food available in the
cage, and its frequency on Thursdays (when the cage is
cleaned) was not greater than that expected by chance.

Although social rank may interfere with food access, and
may contribute to within-group differences in food de-
ficiency, we found no relationship between this variable
and the frequency of coprophagy. Thus, if the individual's
degree of social pressure (reflected here by frequency of
harassment) is inversely related to social rank, the "social
stress" hypothesis may also be rejected to explain the occur-
rence and distribution of coprophagy in this group.

We did not test the "boredom" and "medical problem"
hypotheses for coprophagy, both of which would have
required detailed information on individual differences
in personality and health status. Therefore, it is not pos-
sible to evaluate whether the observed distribution of co-
prophagy among group members was caused by a single
factor or by an interplay between food deficiency, bore-
dom, and medical problems. Finally, it is possible that
this uncommon behavior has been transmitted culturally
among group members, as proposed for captive chimpan-
zees by Nash et al. (1999). The observations of Roger,
Chico, Tiburcio and Assis (the only four individuals not
involved in coprophagy) drinking urine is compatible
with Stemmler-Morath's (1937, apud Hill, 1966) find-
ings that apes began coprophagy by drinking their urine.
To better understand the causes and significance of co-
prophagy for capuchin monkeys, it is important that re-
searchers report all observations of this behavior both in
captivity and in the wild.

Acknowledgements: We thank the personnel of the Parque Far-
roupilha, especially the zookeepers, for their support during
this study.

Helissandra Mattjie Prates and Jtilio C6sar Bicca-Marques,
Pontificia Universidade Cat61lica do Rio Grande do Sul, Fac-
uldade de Biociencias, Av. Ipiranga 6681, Pd 12A, Porto
Alegre 90619-900, Rio Grande do Sul, Brazil. E-mail: icca@pucrs.br>.


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Gabriela Bruno, Aldo M. Giudice
Mariela Nieves, Marta D. Mudry

Alouatta caraya posee una amplia distribuci6n en Sudame-
rica ocupando distintos tipos de bosques hasta los 29056'S,
en los cuales muestra una plasticidad comportamental des-
tacada entire los primates del Neotr6pico (Neville et al.,
1988). En condiciones de cautividad los aulladores negros
no sobreviven largos perfodos de tiempo, sugiridndose la
incapacidad de sustituir su dieta en estas condiciones, o
bien la elevada susceptibilidad a situaciones generadoras
de estres entire las razones de esta baja viabilidad (Benton,
1976; Colillas y Coppo, 1978; Giudice et al., 1995). Resul-
ta parad6jico que s6lo se reproduzca excepcionalmente en
cautiverio, bajo condiciones de cuidado intensive; mientras
logra reproducirse exitosamente en condiciones de liber-
tad, en ambientes forestales ex6ticos con presi6n antr6pica
y mas al sur de su Ifmite austral natural de distribuci6n.
Estas condiciones no parecerfan ser las mas apropiadas para
su supervivencia, ya sea por el rigor climAtico, la extrema
fragmentaci6n del habitat en cercanfa del hombre o por la
pobre calidad y cantidad de la oferta de recursos alimen-
tarios (Giudice y Ascunce, 1998). En esta oportunidad se
presentan las primeras observaciones sobre habitat, supervi-
vencia, dieta y nacimientos, de A. caraya en condiciones de
semilibertad, en bosques serranos, fuera del limited natural
de su distribuci6n marginal sur.

Sitio de Estudio

La poblaci6n de A. caraya que es objeto de este report es
parte de un studio ecol6gico y comportamental en una
tesis doctoral de la Universidad de Buenos Aires. Esta po-
blaci6n habitat dentro de los limits de una estancia privada
de 300 ha en la que se encuentra el Centro de Reeducaci6n
del Mono Aullador Negro (CRMAN), surgido hace 10
anios como una respuesta a la problemAtica relacionada con
la rehabilitaci6n y mantenimiento de ejemplares de aulla-
dores negros extraidos de su entorno natural para su venta
en el mercado illegal de mascotas y que posteriormente son
retenidos por diversas organizaciones de la provincia de
C6rdoba (Bruno et al., 2004).

El CRMAN se encuentra ubicado a 1409 m.s.n.m., en el
paraje Tiu Mayu (3058'S, 6425'0), C6rdoba, Argentina.

El clima templado serrano tiene temperatures que osci-
lan entire -8.8C y 32C en invierno y 0.80C y 38.4C en
verano, con una precipitaci6n annual de aproximadamente
700 mm (Demaio y Medina, 1999). El Area citada esta in-
serta en la provincia fitogeografica chaquefia, distrito serra-
no, caracterizado por sectors de bosque y pastizal de altura
(Cabrera, 1976). Esta zona esta notablemente alterada por
las actividades humans; las species caracteristicas del
bosque nativo han sido paulatinamente eliminadas, siendo
este bosque reemplazado casi en su totalidad por species

Sujetos de Estudio y Manejo

El CRMAN se caracteriza por recibir ejemplares proceden-
tes de donaciones de particulares, asf como de incautacio-
nes/decomisos y derivados por el Departamento de Fauna
Provincial (C6rdoba, Argentina). Los ejemplares derivados
al centro pasan un perfodo de atenci6n intensive, en el cual
son sujetos a los procedimientos clinics y terapeuticos que
los casos demanden. Posteriormente, se forman grupos para
introducir en parches de monte ex6tico de aproximadamen-
te 0.18 ha cada uno. En estos fragments de vegetaci6n se
han colocado travesanios para mejorar el desplazamiento en
el dosel, tarimas para la colocaci6n de los alimentos apor-
tados por el centro y tambores metalicos de 0.60 x 1.00 m
acondicionados para funcionar como refugios.

Una vez que los monos estan en libertad se les aporta agua y
alimentos en una raz6n diaria de 3 kg por grupo aproxima-
damente. La dieta suministrada esta compuesta por verdu-
ras de hoja, frutas, pan, huevos, suplementandose ademis
con td y leche. Todo ejemplar enfermo o que muestre signos
de incompatibilidad social, es recapturado, reiniciandose la
primera etapa de rehabilitaci6n. Hasta la fecha, se han for-
mando cuatro grupos, cuyo tamafio y composici6n sexo/
edad se detallan en la Tabla 1.

Datos Preliminares

El CRMAN ha trabajado sobre un total de 89 aulladores
desde 1994 hasta la fecha. Actualmente se mantienen 54
ejemplares, de los cuales 20 estan bajo la etapa de cuidados
intensivos y el resto, 34 ejemplares, estan formando cuatro
grupos, en distintos parches de monte ex6tico principal-
mente caducifolio (Tabla 1). Adn cuando en el Area del
CRMAN subsiste escasa vegetaci6n arb6rea native como
Fagara coco y Lithraea ternifolia (representando al bosque
serrano) y la presencia de gramineas como Stipa sp. y Festuca
sp. en el pastizal de altura (obs. pers., G. Bruno, 2004), hay
una alteraci6n en la composici6n floristica por la introduc-
ci6n de species de Arboles ex6ticos, tal como Ulmusprocera,
U. laevis, Robinia pseudoacacia, Populus nigra, Malus sylves-
tris, Thuja occidentalis, Cupressus macrocarpa y Salix fragilis,
entire otros. La reproducci6n con dxito comenz6 a partir de
1998, totalizando 26 nacimientos hasta 2005, los cuales se
produce a raz6n de cuatro ejemplares al afio. El intervalo
promedio entire nacimientos para las hembras multiparas

Neotropical Primates 13(3), December 2005

Tabla 1. Composici6n de grupos de A. caraya en semilibertad en el Centro de Rehabilitaci6n del Mono Aullador Negro.

Grupo Ndiero total de Machos Hembras Adultos Juveniles Crias

1 9 6 3 4 2 3
2 9 3 4 5 1 3 (2 sin sexar)
3 8 4 3 3 4 1(sin sexar)
4 8 4 4 6 1 1

es de 14.62 7.93 meses. Es important destacar que los
grupos formados permanecen en los parches, observando-
se cohesi6n con establecimiento de jerarqufas. Tambikn se
han registrado interacciones vocales entire grupos vecinos y
comportamientos especie-especificos, muchos de ellos rela-
cionados con el manejo del ambiente que habitan. Como
ejemplos se puede mencionar el cuidado maternal, asisten-
cia a las crfas por parte de tfas, juego entire las crias y ju-
veniles, acicalamiento entire los distintos integrantes de la
tropa, grupos t&rmicos, y alimentaci6n espontanea de hojas
y frutos de distintas species del estrato arb6reo y herbiceo.
Los monos consumieron espontaneamente hojas nuevas y
maduras de Robinia pseudoacacia, Salix fragilis, brotes de
Populus nigra, hojas y brotes de Ulmusspp., y hojas y frutos
de Rubus ulmifolius. Estos recursos estan disponibles s6lo
una parte del afio, en las estaciones de primavera y verano,
siendo suplementados con mayor intensidad con una dieta
artificial en el rest del afio, cuando los parches de bosque
pierden su follaje casi por complete.

Los datos preliminares parecen avalar un process de rehabi-
litaci6n integral de estos ejemplares extrafdos de su entorno
natural. Si comparamos la elevada tasa de reproducci6n en
el CRMAN con los escasos events que se registran en cau-
tividad (Fitchner Gomes y Bicca-Marques, 2003), estas ob-
servaciones preliminares permiten interpreter que A. caraya
proveniente del mercado illegal de mascotas puede sobrevivir
en estado de semilibertad, adaptindose a un ambiente na-
tural antropogenico que permit expresar un repertorio de
conductas compatibles con las especie-especificas descriptas
en el extreme austral de su distribuci6n -,-' Ci,. *i natural.

Si bien los studios de ecologfa de A. caraya en este am-
biente serrano antropogenizado estan en una fase inicial de
analisis, ante los hallazgos y el possible seguimiento de las
tropas en el lugar y en el tiempo, el CRMAN se constituiria
en una possible alternative para la protecci6n y conservaci6n
de A. caraya, una especie de dificil mantenimiento en el
cautiverio traditional de los zool6gicos, quedando adn para
su consideraci6n, si estos ejemplares rehabilitados pueden
ser reintroducidos con dxito en Areas particulares de su hi-
bitat natural.

Agradecimientos: Este trabajo no serfa possible sin el apoyo e
interns institucionales junto a la informaci6n brindada por
la Director del CRMAN, Prof. M. A. Juarez. Se cont6 con
la participaci6n en los subsidies MDM-UBACyT X-031,
X-107; CONICET PIP 2450/01.

Gabriela Bruno, Centro de Reeducaci6n del Mono Au-
llador Negro (CRMAN), Camino Estancia El Rosario, As-
cochinga RE66, C6rdoba, Argentina, Aldo M. Giudice,
Mariela Nieves y Marta D. Mudry, Grupo de Investiga-
ci6n en Biologfa Evolutiva gibeE), Departamento de Eco-
logfa, Gen&tica y Evoluci6n, Facultad de Ciencias Exac-
tas y Naturales, Ciudad Univ. Pabell6n II, 4to Piso, Lab.
46-47 (EHA1428), Ciudad Aut6noma de Buenos Aires,


Benton, L. 1976. The establishment and husbandry of a
black howler (Alouatta caraya) colony at Columbia Zoo.
Int. Zoo Ybk. 16: 149-152.
Bruno, G., Giudice, A. M., Nieves, M. y Mudry, M. D.
2004. Proyecto de recuperaci6n y mantenimiento de Alou-
atta caraya (Platyrrhini: Atelidae) en ambientes antro-
pizados. Actas XIXJornadas Argentinas de Mastozoologia,
p.20. SAREM, Puerto Madryn, Argentina.
Cabrera, A. L. 1976. Fitogeograffa de la Repdblica Argenti-
na. Boletin de la SociedadArgentina de Botdnica 14 (1-2):
Colillas, 0. y Coppo, J. 1978. Breeding Alouatta caraya
in Centro Argentino de Primates. En: Recent Advances in
Primate Conservation, Vol. 2: Conservation, D. J. Chivers
y W. Lane-Petter (eds.), pp.201-214. Academic Press,
Demaio, P. y Medina, M. 1999. Ecosistemas de la Provincia
de Cdrdoba. Sezo, C6rdoba, Argentina.
Fichtner Gomes, D. y Bicca-Marques, J. C. 2003. Births
of Alouatta caraya and A. belzebul (Atelidae, Alouattinae)
in captivity in Brazil. Neotrop. Primates 11(2):
Giudice, A. M., Sassaroli, J. C. y Ferraro, H. 1995. Di-
namica poblacional y estado clfnico de Alouatta caraya
en el Jardfn Zool6gico de Buenos Aires. Actas Xa Jornada
Argentina deMastozoologia, pp.32-33. SAREM, La Plata,
Giudice, A. M. y Ascunce, M. S. 1998. Presencia de Alou-
atta caraya fuera de su Area de distribuci6n natural. Neo-
trop. Primates 6(3): 82-86.
Neville, M. K., Glander, K. E., Braza, E y Rylands, A. B.
1988. The howling monkeys, genus Alouatta. En: Eco-
logy and Behavior of Neotropical Primates, Vol. 2, R. A.
Mittermeier, A. B. Rylands, A. F Coimbra-Filho y G. A.
B. da Fonseca (eds.), pp.349-453. World Wildlife Fund,
Washington, DC.

Neotropical Primates 13(3), December 2005


Margaret R. Clarke
Kenneth E. Glander


Male reproductive success in nonhuman primates has tra-
ditionally been measured by access to estrous females, and
successful matings/inseminations are attributed either to
high-ranking males in multi-male groups or to the one
male in a single-male group (e.g., Fedigan, 1983; Smuts,
1987; Dixson et al., 1993; Paul et al., 1993). Paternity
exclusion techniques and the testing of potential fathers
and infants, however, have demonstrated that access, in-
cluding copulation, does not always correlate with suc-
cess. As group size increases, there are more females for the
highest-ranking male to inseminate, and if the females are
seasonal or synchronous breeders, the male is less able mo-
nopolize estrous females (Nunn, 1999; Takahashi, 2001).
The discrepancy between dominance rank and mating
success involves more than female availability, however.
In single-male patas monkey groups, the resident male
did not sire all offspring (Ohsawa et al., 1993), and alter-
nate male strategies in multi-male groups are known from
rhesus monkeys (Berard et al., 1993, 1994, 1999; Smith,
1993, 1994) and savannah baboons (Alberts et al., 2003)
which allow for reproductive success in lower-ranking

True reproductive success is defined as the total number
of surviving offspring, which can only be measured over
the lifetime of an individual male. Given the practical dif-
ficulties, this may be reduced to a pair of simple proxies:
the length of time a male is fertile and has access to fer-
tile females, and the number of other males competing for
those same females. These two measures may serve as useful
predictors of potential reproductive success.

Mantled howlers (Alouattapalliata) may form both single-
male and multi-male groups within the same population.
Since howler groups often experience takeovers by an
outside male, there could be an advantage to living in a
multi-male group, as low-ranking males would have the
"protection" of the high-ranking males' competitive abil-
ity to repel newcomers (Nunn, 2000). Although all males
would compete for access to estrous females, occasional re-
productive synchrony could benefit low-ranking males; the
dominant male can only monopolize one female at a time,
which allows other males access to other fertile females.
Conversely, the lone male in the single-male group would
be expected to have access to all estrous females and father
all offspring born in that group (Ridley, 1986), but would
be at higher risk for predation or a takeover by an outside
male (van Schaik and Horstermann, 1994).

To evaluate the effects of male tenure on reproductive
success in both single-male and multi-male groups, we
examined our records for group membership and infant
survival in seven groups of howlers at Hacienda La Pacifi-
ca between 1970 and 2002. These included one group
that had always had only one male, two groups that were
always multi-male, and four groups that fluctuated be-
tween single- and multi-male status. As resource availabil-
ity could have affected reproductive success, we carried
out a similar analysis for males in riparian habitats (three
groups), which could be considered richer in resources
(Glander and Nisbett, 1996), vs. males in upland habitats
(four groups).

We have made two important assumptions in this analy-
sis. First, we assume that the male in a single-male group
sires all offspring; and second, we assume that over a male's
lifetime, males in multi-male groups have equal reproduc-
tive success. Takeovers by young males result in a reverse
age-graded dominance hierarchy in this population (Glan-
der, 1980; Jones, 1980). A young, dominant male would
be expected to have high reproductive success, which
would presumably decrease as the male aged and lost his


Study site
La Pacifica is a working cattle ranch / rice farm / tilapia
farm in the dry tropical forest zone of Guanacaste Prov-
ince, Costa Rica (Holdridge, 1967). Three rivers border
the farm with associated riparian habitat. The farm was
deforested in a comparatively conservative manner in the
1950s for cattle ranching The upland habitat was con-
verted to pastureland, leaving strips of forest as wind-
breaks between pastures, and large areas of forest were
left in hilly, rocky areas deemed unsuitable for grazing.
Forested areas along the three rivers were untouched. The
farm was originally irrigated through a low-maintenance,
low-impact system operated by gravity, pulling water from
the river on the north side of the farm. In 1986 the farm
was sold and irrigation patterns changed: large machinery
was used to clear larger ditches which increased waterflow
and caused extensive soil erosion. In 1991 a substantial
section was deforested for a major government irrigation
canal, and in 1998 a major shift in land use occurred as
pastures were converted to wet rice agriculture or tilapia
tanks. Many of the forest areas were disturbed, but many
others remain, and the size of the monkey population re-
mained essentially unchanged from the 1970s to 1998
(Clarke et al., 2002).

Study subjects
Animals from seven different social groups have been
tracked for various periods from 1970 to the present.
Thirty-three males were included in this analysis, eight in
single-male groups and 30 in multi-male groups. Five of
these males spent time in each type of group.

Neotropical Primates 13(3), December 2005

Data ,., .-
This analysis is based on data collected during population
surveys from 1974-76, 1984, 1991, and 1998 (Clarke et
al., 1986; Clarke and Zucker, 1994; Clarke et al., 2001),
yearly counts of groups by MRC, and repeated capture and
mark sessions from 1970 to 2002. Behavioral studies car-
ried out by M. R. Clarke, K. E. Glander, R. A. Nisbett and
E. L. Zucker during the past thirty years have characterized
group composition in detail.

To analyze male tenure, we compared the number of years
for each male in a multi-male group (n = 30) to the number
of years for each male in a single-male group (n = 8) using
an independent t-test. We also tested the number of years
spent in a single- vs. a multi-male group for the same males
in the same groups (n = 5), using a repeated measures t-test.
We compared the number of years in single-male (n = 3),
multi-male (n = 25), or both group types (n = 5) using an
independent one-way ANOVA. We used an independent
t-test to compare the number of years spent in a riparian
habitat group vs. an upland habitat group. For the habitat
comparison, total time for each male in the group was used
regardless of male composition.

To calculate male reproductive success, we divided group
history for all groups into time blocks of single- or multi-
male tenure (seven blocks of multi-male, eight blocks of
single-male). For each time block, the total number of
"male years" was calculated and divided by the years in
that time block to yield an average number of males per
year. (E.g., in a four-year time block, if there were two
males in the group for all four years and a third in the last
year, there were 4 + 4 + 1 = 9 male-years divided by the
four years of that time block, yielding an average of 2.25
males/year.) We used the same approach to calculate the
mean number of females per year. We also tabulated the
mean number of infants born per year and surviving per
year. We calculated the potential number of infants per
male (potential infant/male) by dividing the mean number
of females by the mean for males. Realized reproductive
effort was calculated by dividing the mean number of in-
fants born to the mean number of males, and dividing
that by potential infant/male, while relative infant loss was
calculated by subtracting the mean number of infant sur-
vivors from the mean of all infants born. Each of these
measures was tested by habitat (riparian vs. upland) and
by male composition (single-male vs. multi-male) using
independent t-tests.


Male tenure
We found no significant differences in time spent in a sin-
gle-male group or in a multi-male group (Table 1). For the
five males which spent time in both group types, we also
found no statistical differences between the length of time
they spent in a single-male vs. a multi-male group (Table

1). However, when total time for specific males that alter-
nated between group types is compared to males living ex-
clusively in a single-male or multi-male group, males with
the alternating strategy were in social groups significantly
longer overall (Table 1). As before, however, we found no
differences between animals living only in a single-male or
in a multi-male group (Tukey post hoc test, p < 0.05, criti-
cal diff. = 6.0: single/multi = 0.45; single/both = 6.7; multi/
both = 7.1). There were no differences in length of time in
social groups by habitat type (Table 1).

Reproductive success
We found no difference in realized reproductive effort per
male between single- and multi-male groups (see Table 2),
but the potential for infant production was significantly
greater in the single-male groups. Infant survivorship, when
expressed as a ratio of infants born minus infants surviv-
ing, was greater in a single-male group, although it only
approached statistical significance (see Table 2). More in-
fants per female per male were born in riparian habitats,
but neither the number of potential infants per male nor
infant survivorship were associated with habitat type (see
Table 2).


We expected that males would reside in multi-male groups
longer than in single-male groups, but this was not borne
out. Residence in a larger group would presumably buffer
an individual male's reproductive success against the likeli-
hood of a male takeover, ensuring that the resident male
shared group membership rather than having no member-
ship at all. In theory, the shared group membership would
be balanced by increased competition for access to fertile
females, as opposed to the presumably sole access in the
much smaller single-male group. The true situation, how-
ever, is apparently much more complex.

We found no differences in the mean number of infants
born per male between single- or multi-male groups, but
there were significantly more females available per male in
the single-male group (almost double) than in the multi-
male groups. Thus, males in single-male groups are not
achieving all possible pregnancies, which raises several
questions. Are they spending their time defending their
females rather than copulating? In the absence of male-
male competition, do they monitor female receptivity less
carefully? Even considering the approximately two-year
interbirth interval documented in this population (Glan-
der, 1980), half of the females should give birth every year
regardless of the number of available males. As reproduc-
tive success involves infant survivorship as well as number
of offspring, it should be noted that infant survivorship is
somewhat greater in single-male groups. Thus, a female
might be less likely to conceive in a single-male group, but
once she is pregnant, it would appear that her infant has
a better chance of surviving its first year in a single-male

Neotropical Primates 13(3), December 2005

Table 1. Group tenure. Comparison of time in years which males spent in single-male groups, multi-male groups or both group types,
and for groups in riparian vs. upland habitat.
Mean SD Mean SD Mean SD t / F df Significance
Single-male Multi-male Both
Sample Type:
One-multi (all) 5.6 3.0 6.2 3.6 t = 0.41 36 n.s.
Repeated 5.2 2.6 7.8 4.5 t = -1.34 4 n.s.
One/multi/both 6.3 4.0 5.9 3.4 13.0 6.0 F = 6.9 32 p < 0.01

Riparian Upland

All males 8.1 5.4 6.1 3.7 t = 1.23 31 n.s.

Table 2. Reproductive success. Realized and potential reproductive success by year for males in single-male and multi-male groups, and
by riparian vs. upland habitat type, including infant loss.
Mean SD Mean SD t df Significance
Single-male Multi-male
Sample (mean per year)
Realized reproduction/male 0.4 0.2 0.5 0.1 -1.16 13 n.s.
Potential infant/male 4.3 1.2 2.6 0.8 3.17 13 p < 0.01
Relative infant loss 0.4 0.6 0.9 0.5 -2.05 13 p = 0.06

Riparian Upland

Realized reproduction/male 0.5 0.1 0.3 0.1 2.68 13 p < 0.02
Potential infant/male 3.3 1.0 3.7 1.5 -0.54 13 n.s.
Relative infant loss 0.7 0.6 0.6 0.6 0.50 13 n.s.

Single-male groups are generally smaller (range = 4-12) than group, or form a new group with peripheral females. There
multi-male groups (range = 8-42), and while single-male is evidence that both patterns exist (Clarke and Glander,
groups occur in both habitat types, they are more common 2004).
in upland habitat (Clarke et al., 2002). Upland habitat is
more affected by the distinct seasonality of the dry tropi- The most surprising result is the almost complete lack of
cal forest, and upland habitat females are lighter than their association of either tenure or reproductive parameters with
riparian counterparts (Teaford and Glander, 1997; Glan- habitattype.The dichotomy between the upland habitat and
der, in press). These are factors which could contribute to riparian habitat in dry tropical forest is a common analyti-
females not conceiving, but they would not explain higher cal parameter, but it should be noted that while the groups
infant survivorship in single-male groups. As high-ranking labeled "upland" never used riparian habitat, all of those
females often attack the infants of lower-ranking females labeled "riparian" do make extensive use of upland habitat
(Clarke et al., 1998), it is possible that having fewer females at some times of the year (pers. obs.). A confounding effect
in a single-male group may improve infant survivorship, an is involved because single-male groups are more common
important factor unrelated to the number of males. in the upland habitat, but from this analysis, it appears that
the composition of social groups is a better predictor of
From the male standpoint, it might seem that the best strat- tenure and reproductive success than the habitat alone.
egy for long-term access to females is to maintain group
membership, regardless of whether it is in a single-male or This analysis is based on assumptions that need to be
a multi-male group. This flexible strategy is not common, confirmed through paternity exclusion tests. The overall
however, as only five of the 33 males in this sample were conclusion-that males have improved reproductive suc-
able to pursue it successfully. While males can stay in a cess through complete access to females in a single-male
multi-male group after a new male takes over, the sex-ratio group, as opposed to competing for females in a multi-
becomes less favorable thereafter, with fewer females in pro- male group-should not be accepted without question.
portion to males. Older males have two alternative strate- These results, based on long-term field records, should pro-
gies: leave a multi-male group and take over a single-male vide evidence that presumptions about male reproductive

Neotropical Primates 13(3), December 2005

success based on observations of potential access to females
alone are not valid.

Acknowledgements: The authors acknowledge the financial
support of the National Science Foundation, National
Geographic Research Grants, Explorer's Club, School for
Field Studies, Earthwatch, Duke University, Newcomb
College and Center for Latin American Studies from
Tulane University, and the Tulane National Primate Re-
search Center. We thank the management of La Pacifica,
particularly Fernando Estrada, for allowing us to continue
our research at the site, and we thank Ronald Carrera, Luis
Herra, Vreni Hagnauer, Jorge Hagnauer, Werner Hagnau-
er and all of the La Pacifica personnel who have helped us
over the years. We thank the legions of students for their
contributions to the dataset, and Norm Scott Jr., Richard
Nisbett, Evan Zucker, Carolyn Crockett, Maria Zaldivar
and Mark Teaford for both formal and informal contri-
butions to the information presented in this paper. The
impetus for this analysis came from Danielle Epstein, and
the analysis was improved by suggestions from Randall

Margaret R. Clarke and Kenneth E. Glander, Dept. of
Biological Anthropology and Anatomy, Duke University,
Durham, North Carolina 27708, USA. Corresponding
author: Margaret R. Clarke, Dept. of Neurobiology and
Anatomy, UTHSC-HOUSTON, 6431 Fannin St., MSB
7.046, Houston, TX 77030. E-mail: uth.tmc.edu>.


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Wilmer E. Pozo R.


En general, la mayoria de los primates duermen en Arboles,
pero algunas species del Viejo Mundo, como los hamadrias
(Papio hamadryas), duermen en penascos (Stammbach,
1987), los orangutanes (Pongo pygmaeus) pernoctan en
plataformas construidas sobre el suelo con ramas y hojas
(Rodman y Mitani, 1987) y los gorilas (Go. .//t gV.//1i)
pasan la noche en nidos construidos en los Arboles o sobre
el suelo (Schaller, 1963; Morris, 1991).

En los bosques Neotropicales, todos los primates duermen
en Arboles, cuyas caracteristicas difieren de una especie a

otra. Asf, por ejemplo, los monos nocturnos (Aotus spp.)
utilizan diferentes tipos de dormideros, tales como huecos
en troncos y ramas de Arboles secos o envejecidos (obs.
pers.), sitios complejos formados por una masa vegetal de
epffitos, trepadoras y enredaderas, o sitios simples de un
follaje denso (Aquino y Encarnaci6n, 1986). Los escasos
studios de campo respect a primates del gdnero C
sugieren diferencias entire sus species, pero la mayoria de
ellas aumentan el ndmero de Arboles utilizados como dor-
mideros con el tamafo de sus Ambitos hogarenos, pudiendo
tambidn usar el mismo dormidero por varias noches conse-
cutivas (Stevenson y Rylands, 1988). Los monos chichicos
(Saguinus spp.) acostumbran cambiar frecuentemente sus
dormideros, los mismos que son seleccionados estrategica-
mente a fin de minimizar el contact con sus predadores
(Snowdon y Soini, 1988). Especies del gdnero Leontopithe-
cus presentan la tendencia de dormir en huecos de Arboles
(nidos) abandonados por otras species (Kleiman et al.,

Monos de tamafo mas grande, como los aulladores (Alou-
atta spp.), duermen en las ramas horizontales de Arboles
de median a gran tamafo; casi siempre estos Arboles son
forrajeados antes de ser usados como dormideros (Neville
et al., 1988). Los chorongos del Yasunf (Lagothrix lago-
thricha) forman grandes agrupaciones sociales (Di Fiore,
1997). Estos grandes grupos forrajean juntos y al final del
dfa se dividend en subgrupos que duermen muy cercana-
mente, usando various Arboles de caracteristicas diferentes
(obs. pers.). Observaciones de campo realizadas por Rami-
rez (1988) en la Amazonfa peruana indican que un subgru-
po de cinco individuos escogi6 para dormir un gran Arbol
completamente lleno de hojas. Los muriqufs del Brasil
(Brachyteles arachnoides) generalmente duermen en el estra-
to medio del bosque, utilizando las ramas bifurcadas de sus
arboles dormideros (Nishimura et al., 1988).

Existe escasa informaci6n que caracterice los dormideros
que usa Ateles belzebuth, probablemente debido a la dificul-
tad que represent seguir a individuos de esta especie. Sin
embargo, van Roosmalen (1985) y Chapman (1989) han
reportado que los monos arana (Ateles paniscus paniscus y
Ateles ..rr .., respectivamente) prefieren dormir en Arbo-
les de los estratos altos del bosque, usando entire 11 y 43
individuos arb6reos y que muchos de estos Arboles suelen
utilizarse para dormir en multiples ocasiones. Generalmen-
te, estos primates duermen en subgrupos que ocupan uno
o various Arboles dormideros y el tamano de los subgrupos
se relaciona con las necesidades forrajeras de sus individuos
(Chapman, 1989).

En este articulo se described las caracteristicas de los sitios y
Arboles utilizados como dormideros por parte de un grupo
de Ateles belzebuth en el Parque Nacional Yasunf (PNY),
Ecuador, durante un seguimiento realizado entire diciem-
bre de 1994 y febrero de 1996, con el fin de estudiar el
comportamiento social y las costumbres alimenticias de la
especie (Pozo, 2001, 2004a).

Neotropical Primates 13(3), December 2005

Figura 1. Sistema de transectos del Area de studio. Las letras diferentes indican los nombres de los senderos; PB, PC y PD se utilizaron
para estudiar la estructura de habitat.


El studio se llev6 a cabo en el Oriente ecuatoriano, Pro-
vincia de Orellana, en el Parque Nacional Yasunf, en un
Area algo mayor a 400 ha (Fig. 1), en la que se han condu-
cido various studios primatol6gicos (Di Fiore, 1997, 2004;
Youlatos y Pozo, 1999; Cant et al., 2001, 2003; Pozo,
2001, 2004a, 2004b; Pozo y Youlatos, 2005a). El sitio se
ubica en el kil6metro 47 de la carretera Pompeya Sur -
Iro (00o42'17"S y 7628'05"0) y estA formado por valles
y colinas cuyas altitudes oscilan entire los 210 y 330 msnm.
La temperature media annual, durante el perfodo de estu-
dio, fue de 27.73C, mientras que la precipitaci6n fue de
3690.1 mm (Pozo, 2001).

Holdridge (1982) clasifica al Oriente ecuatoriano en la
zona de vida llamada bosque hdmedo tropical. La forma-
ci6n vegetal del sitio de studio es el bosque siempre verde
de tierras bajas de la amazonfa (Sierra et al., 1999), donde
Iriartea deltoidea, Oenocarpus bataua, Virola pavonis, Otoba
glycycarpa, Parkia '." Pouroma bicolor, Cedrelinga
cateniformis, Ceiba pentandra, Ficus maxima, Trichilia spp.
y Philodendron spp., entire otras, forman parte de la flora

tfpica (Sierra et al., 1999). Una lista complete de las espe-
cies de plants caracteristicas del sitio de studio se present
en Di Fiore (1997) y Pozo (2001).

En una de las revisiones mis importantes para el genero,
Kellogg y Goldman (1944) consideraron que Ateles belze-
buth estaba compuesta por tres subespecies (A. b. belzebuth,
A. b. marginatusy A. b. hybridus), mismas que recientemen-
te han sido elevadas a nivel de especie (Groves, 2001). La
localidad tipo de Ateles belzebuth es desconocida (Kellogg
y Goldman, 1944), pero se sabe que esta especie se distri-
buye desde Venezuela, al este de la bocana del rio Guap6
en el Orinoco, hasta el nororiente peruano en el Sarayacu
(Kellogg y Goldman, 1944). En Ecuador, la especie vive
s6lo al Oriente, desde el sur del rio Napo hasta la fronte-
ra sur con Perd. En espafiol, se le denomina como "mono
arafia", los Huaoranis la llaman "deiye", mientras que los
quichuas la conocen como "makisapa", que significa mano

Un grupo de Ateles belzebuth de 25 individuos entiree crias,
juveniles y adults de ambos sexos, segdn el criterio de
Izawa et al. [1979]), fue seguido diariamente por el mayor

0m 300m

Neotropical Primates 13(3), December 2005

Tabla 1. Estructura del habitat del sitio de studio. Los resulta-
dos expresan los porcentajes de las observaciones realizadas a 303
puntos de observaci6n y a 493 Arboles y palmas. BA = bosque
alto, BA/L = bosque alto con lianas, BL = bosque de lianas, BB =
bosque bajo, BT = bosque transicional, CLB = claro de bosque,
TAB = arbol con DAP tabular, PB, PC y PD = nombre de los
transectos donde se estudi6 la estructura del habitat; vea tambikn
Fig. 1.
Tipo de Subtipos Transectos Area
Estructura Subtpos PB PC PD Total
a) Topografia Cima 6 9 21 12
Ladera 48 46 42 45
Riachuelo 4 11 4 6
Terraza 31 18 30 26
Valle 12 17 4 11
Total 100 100 100 100
b) Bosque BA 42 76 56 58
BA/L 8 6 9 8
BL 11 1 5 6
BB 2 0 2 1
BT 20 7 21 16
CLB 18 10 7 12
Total 100 100 100 100
c) DAP 1-20 cm 43 37 53 44
20-40 cm 40 42 35 39
40-60 cm 7 9 5 7
60-80 cm 4 2 0 2
80-100 cm 1 0 1 1
tab 5 10 7 8
Total 100 100 100 100
d) Altura 1-5 m 3 2 1 2
5-10 m 26 15 31 24
10-15 m 28 31 36 32
15-20 m 22 33 22 26
20-25 m 14 16 8 12
25-mas 7 3 2 4
Total 100 100 100 100

tiempo possible. Una vez que los monos llegaron a un Arbol
dormidero, se tom6 la siguiente informaci6n: fecha, hora,
topografia, tipo de formaci6n boscosa en la que se ubic6
en Arbol dormidero, diametro a la altura del pecho (DAP),
altura del Arbol dormidero, altura a la cual los animals
usaron el Arbol dormidero y, en lo possible, el tax6n.

Para conocer las caracteristicas ecol6gicas del sitio de es-
tudio, se realize un analisis rapido de la estructura de ha-
bitat (Pozo y Youlatos, 2005). Este analisis se aplic6 a tres
senderos (PB, PC y PD, Fig. 1), mismos que presentaron
una orientaci6n de 320 noroeste y una longitud de 1000
m. Los senderos fueron marcados cada 10 m en un punto
medio, desde el cual se tom6 informaci6n en un ancho
aproximado de 10 m. De acuerdo con el DAP, la altura
arb6rea y el diametro de corona de 493 Arboles, asi como
los hAbitos de las plants presents en los puntos de ob-
servaci6n (palmas, epifitos, lianas, Arboles maduros, etc.),
se calific6 al tipo de vegetaci6n como bosque alto, bosque
alto con lianas, bosque de lianas, bosques bajos, bosques
transicionales y claros del bosque. Las caracteristicas bio-
fisicas de los tipos de formaciones boscosas se aprecian en
la Tabla 2.

Se hipotetiz6 (H0) que los monos arafia utilizan los Arboles
para dormir de manera aleatoria, y por tanto, las caracterfs-
ticas ecol6gicas de los sitios en donde se localizan los Arbo-
les dormideros serfan iguales a las caracteristicas ecol6gicas
generals del sitio de studio. Por ejemplo, se espera que
las proporciones de tipos de bosque donde se localicen los
Arboles dormideros sean similares a las proporciones de hi-
bitat disponibles en el Area de studio. Para contrastar esta
hip6tesis se realize un analisis de estructura del habitat y
la informaci6n obtenida (Tabla 1) se us6 como resultados
esperados en las pruebas de x2 aplicadas a los resultados ob-
tenidos, con un nivel de decision a de 0.05. En lo que res-
pecta a DAP, s6lo 14 de los 16 Arboles dormideros fueron
medidos; de ellos, cinco presentaron bases con races ta-
bulares, en cuyo caso no se consider esta media para el
analisis de los datos.

Tabla 2. Caracteristicas de las formaciones boscosas del sitio de studio (segdn Pozo y Youlatos, 2005a y 2005b). La tabla indica los re-
sultados globales de la altura en m y el diimetro a la altura del pecho (DAP) en cm de los Arboles muestreados en los senderos PB, PC y
PD; n = ndmero de Arboles medidos en cada formaci6n boscosa, D = arbol derecho, I = arbol izquierdo, PT = promedio total de los datos
obtenidos, TAB = numero de arboles con bases tabulares.
Altura DAP
Tipo de Formaci6n Boscosa TAB
n D I PT n D I PT_____
Bosque alto 352 15.98 16.05 16.01 316 22.97 26.80 24.89 36
Bosque alto/lianas 44 15.07 12.54 13.80 42 21.95 21.06 21.50 2
Bosque de lianas 14 12.21 9.46 10.83 14 22.63 18.24 20.43 0
Claros de bosque 29 11.65 10.83 11.24 29 21.68 17.10 19.39 0
Bosque transitional 46 9.15 10.73 9.92 46 16.28 16.65 15.78 0
Bosque bajo 8 7.75 7.50 7.63 8 9.90 14.28 11.09 0

Neotropical Primates 13(3), December 2005

Tabla 3. Resultados de las pruebas de Z' para determinar la preferencia de los monos arafia para dormir en los distintos tipos de bosque,
de topograffa, de estrato arb6reo y de DAP de los arboles utilizados como dormideros.
Preferencia de:
Estadisticas Bosque Topograffa Estrato DAP
Suma de categories 100 100 100 100
Z2 Bioestat 44.479 160.192 1143.458 346.057
Grados de libertad 5 4 5 5
(p) <0.001 <0.001 <0.001 <0.001
Significancia ** ** ** **
Ho Rechazada Rechazada Rechazada Rechazada
H1 Aceptada Aceptada Aceptada Aceptada

8 -"- Observada
70Figura 2. Frecuencias observadas vs. recuencias esperadas del uso

50- -.---rspdrada
so -


Tipo de ronmcion boscosaf

Figura 2. Frecuencias observadas vs. frecuencias esperadas del uso
de dormideros en cada tipo de formaci6n boscosa. BA = bosque
alto, BA/L = bosque alto con lianas, BL = bosque de lianas, BB
bosque bajo, BT bosque transicional, CLB = claro de bosque.

60 -

370 --- -pperda /

m ladera r10- chulo te152 s alle20-25m >25 m
Tipo de topografia

Figura 3. Frecuencias observadas vs. frecuencias esperadas del uso
de dormideros considerando su ubicaci6n t.. .

6 --- Rspemda

30 -


0-O5m 05-10m 10-15 I 15-20 m 20-25 m >25 m
Tipo deevtrato

Figura 4. Frecuencias observadas vs. frecuencias esperadas del uso
de estratos para dormir.


Preferencia de hdbitat
En el 75% de las ocasiones (n = 16), el grupo estudiado
durmi6 en un tipo de formaci6n boscosa denominada
bosque alto, el 18.75% lo hizo en una mezcla de bosque alto
con lianas y el restante 6.25% en bosque de lianas. Existe
una diferencia altamente significativa entire las frecuencias
de uso de tipo de bosque y las proporciones en las que se
presentan las formaciones boscosas en el sitio de studio (x2
= 44.479, gl = 5, p < 0.001, Tabla 3); lo que implica una
alta preferencia de estos monos para dormir en el bosque
alto (Fig. 2). Considerando la topograffa done se situaban
los Arboles dormideros (n = 16), los animals durmieron
en valles (50%), laderas (25%), terrazas (12.5%) y cimas
(12.5%). Los animals tampoco utilizaron las formaciones
topograficas en las proporciones en que 6stas se disponen
en el Area de studio, mas bien se observ6 la preferencia
para dormir en los valles (Z2 = 160.192, gl = 5, p < 0.001,
Tabla 3, Fig. 3). Dividiendo al bosque en estratos de 5 m
de altura, se determine que los animals usaron el estrato
> 25 m en un 69%, el estrato de 20 a 25 m en un 25% y el
estrato de 10 a 15 m en un 6%, mientras que los estratos de
0 a 5 m, de 5 a 10 my de 15 a 20 m jams fueron utilizados
para dormir. Los monos arafia mostraron una preferencia
significativa para dormir en los estratos mayores a 25 m (x2
= 1143.458, gl = 5, p < 0.001, Tabla 3, Fig. 4).

Caracteristicas de los drboles dormideros
Tan s6lo nueve de los 16 Arboles usados para dormir fueron
determinados por lo menos a nivel de familiar; de estos,
las families mas utilizadas fueron Bombacaceae (n = 3) y
Myristicaceae (n = 2), seguidas por las families Meliaceae,
Mimosaceae, Moraceae, y Papilionaceae (n = 1 cada una).
Las species identificadas como dormideros fueron Virola
pavonis (n = 2), Cecropia ,,lop,'/,y//.r (n = 1), Guarea sp. (n
= 1) y Huberodendron sp. (n = 1). Dos species de la familiar
Bombacaceae y una de la familiar Papilionaceae no pudie-
ron ser identificadas.

Los Arboles usados como dormideros (n = 16) presentaron
una altura promedio de 28.94 m (DS = + 6.05, Xmn = 14,
Xa = 35), siendo el 75% de ellos emergentes. Tan s6lo en
una ocasi6n los animals durmieron en la parte mas alta del

Neotropical Primates 13(3), December 2005

50 I
40 ~ --Espcrada
30 -

0-20 20-40 40-60 60-80 80-100 tab
Tipo de DAP

Figura 5. Frecuencias observadas vs. frecuencias esperadas del uso
de arboles dormideros con distintos DAP; tab = irbol con DAP

Arbol dormidero (30 m). A pesar de siempre usar el estrato
alto de los Arboles dormideros, los monos arafia usualmente
durmieron entire 1 y 9 m mAs abajo de la altura promedio
de dichos Arboles; es decir, que la altura promedio usada
para dormir (n = 14) fue de 23.93 m (DS = + 4.51, Xn
= 12, X = 30), la cual estuvo directamente influenciada
por la altura del Arbol escogido para dormir (R2 = 0.832, r
= 0.912, gl = 12, p < 0.001). Es interesante notar que casi
la mitad de los drboles escogidos para dormir presentaron
lianas y epffitos (43.75%, n = 16).

El DAP promedio de los Arboles dormideros medidos (n =
9) fue de 50.43 cm (DS = + 14.61, Xmn = 30, Xm = 80.1).
Creando rangos de 20 cm para el DAP de los Arboles dor-
mideros, se observ6 que los animals durmieron en Arboles
(n = 14) con DAP entire 20 a 40 cm en un 14.30% de las
veces, entire 40 a 60 cm en un 35.71%, entire 60 a 80 cm
en un 7.14%, entire 80 a 100 cm en un 7.14% y ademis
durmieron en cinco arboles (35.71%) con races tabulares
al nivel del DAP (referidas en lo subsecuente como "DAP
tabular"). Estas proporciones indican que los monos arafia
prefieren significativamente dormir en los Arboles con DAP
de 40 a 60 cm y con DAPs tabulares (2 = 346.057, gl = 5,
p < 0.001, Tabla 3, Fig. 5). De los Arboles con DAP de 40
a 60 cm (n = 5) el 80% fueron emergentes y el 100% ocu-
paron los estratos altos del bosque; mientras que s6lo uno
de los cinco arboles con bases tabulares no fue emergente,
ocupando el estrato medio del bosque.

Composicidn social de los subgrupos durmientes
El tamano de los subgrupos (n = 16) que durmieron en un
Arbol vari6 de un a tres individuos, siendo el 50% de ellos de
un solo individuo, el 12.5% de dos individuos y el restante
37.5% de tres individuos (Tabla 4). En dos de los subgru-
pos con dos individuos y en uno con tres individuos no fue
possible determinar la composici6n sexo-edad. Tan s6lo uno
de los ocho individuos solitarios fue macho adulto. El resto
fueron hembras adults con infante, mismas que fueron
contadas como miembros solitarios debido a la alta depen-
dencia de los infants con sus madres (Izawa et al., 1979).
El subgrupo de dos individuos estuvo conformado por dos
hembras adults con infante. Dos de los cuatro grupos con
tres individuos estuvieron conformados por una hembra

Tabla 4. Composici6n social de los subgrupos que usaron los
drboles dormideros. HA+I = hembra adulta con infante, MA =
macho adulto, ? = individuo de categoria edad-sexo no determi-
nada, HA = hembra adulta sin infante, HJ = hembra juvenile, HSA
= hembra subadulta.
Tipo de Composici6n Freuenia Frecuencia
subgrupo edad-sexo acumulada
Solitario HA+I 7 (43.75%) 7 (43.75%)
Solitario MA 1 (6.25%) 8 (50.00%)
Dto ?, ? 1 (6.25%) 9 (56.25%)
Dto HA+I, HA+I 1 (6.25%) 10 (62.50%)
Trios ?, ?, ? 1 (6.25%) 11 (68.75%)
Trios ?, MA, HA 1 (6.25%) 12 (75.00%)
Trios HA, HA+I, HJ 2 (15.50%) 14 (87.50%)
Trios MA, HA+I, HSA 2 (15.50%) 16 (100.00%)

adulta con infante, una hembra adulta sin infante y por una
hembra juvenile; los otros dos subgrupos de tres individuos
estuvieron integrados por una hembra adulta con infante,
un macho adulto y una hembra subadulta (Tabla 4).


Al realizar analisis de uso preferencial, various autores han
interpretado al dato con mayor frecuencia de observaci6n,
como el que marca la preferencia en el uso de habitat y
de estrato (de la Torre, 1991; Reyes, 1991). No obstante,
con el fin de verificar si los datos de mayor frecuencia son
realmente preferenciales, es necesario compararlos con los
resultados esperados (Fonseca, 1985) o con la disponibili-
dad de hAbitat en el Area de studio (Jim6nez, 1995; Pozo,
2001). Inclusive, se observa en este studio que las frecuen-
cias con las que se usan los diferentes tipos de bosques para
dormir aparentan ser similares a las frecuencias en las que
6stos se disponen en el sitio de studio (Fig. 2). A pesar
de ello, la prueba Z2 indic6 que no existe dicha semejan-
za. Por otro lado, las frecuencias observadas sobre la situa-
ci6n topogrAfica de los Arboles utilizados como dormideros
mostraron una clara preferencia por los monos arafia para
dormir en los valles; y, a pesar de que los estratos mayo-
res a 25 m son escasos en el sitio de studio (Pozo, 2001),
los animals presentan una alta preferencia para dormir en
ellos. Asf, este articulo demuestra la utilidad de la prueba x2
para probar preferencia de uso de hAbitat.

Las species vegetables identificadas como dormideros en el
PNY fueron Virola pavonis, Cecropia .u/,.lo,',/y//,7, Guarea
sp. y Huberodendron sp.; asf como dos species no identi-
ficadas de la familiar Bombacaceae y una de la familiar Pa-
pilionaceae. Van Roosmalen (1985) report a Buchenavia
capitata, Vataireopsis speciosa, Couratari stellata, Hymenolo-
bium flavum, H. petraeum y Parkia pendula como las es-
pecies mayormente utilizadas como dormideros del mono
arafia en Surinam. Se puede notar que no existe una gran
diferencia entire el ndmero de species de Arboles dormi-
deros reportados para A. belzebuth y para A. paniscus; sin

Neotropical Primates 13(3), December 2005

embargo, hay una gran diferencia entire las families, generos
y species utilizadas como dormideros, lo cual seguramente
depend de la disponibilidad de dichas species en los sitios
de studio.

El 75% de los Arboles utilizados para dormir por los monos
arana en el PNY fueron emergentes. Este dato es similar
a los publicados por otros autores, quienes sostienen que
los monos arana prefieren dormir en Arboles muy altos y
grandes (van Roosmalen, 1985; Chapman, 1989). No obs-
tante, los arboles preferidos como dormideros en el PNY
se muestran mas delgados (DAP = 50.43 cm) que los del
Parque Nacional Santa Rosa en Costa Rica (DAP = 75.1
cm; Chapman, 1989). No obstante, es de notarse que la
mayoria de Arboles utilizados para dormir fueron emer-
gentes y ocuparon los estratos mas altos del bosque. Van
Roosmalen (1985) yVan Roosmalen y Klein (1988) sugie-
ren que el uso de Arboles emergentes, de coronas comple-
tamente separadas de los estratos inferiores del bosque, asf
como la preferencia por una corona ancha, abierta y des-
hojada, parecen ser los criterios mas importantes para los
monos arana al moment de elegir los sitios para dormir.
Los monos arana del PNY en tres ocasiones durmieron en
Arboles cuyas hojas habfan cafdo completamente, pero que
a la mitad de su fuste contaban con una masa vegetal com-
pleja formada por lianas, epifitos, enredaderas y copas de
Arboles vecinos mas bajos. Esta masa vegetal dificultaba la
observaci6n de los subgrupos durmientes desde el suelo y
por ende ayudaria a reducir el riesgo de depredaci6n por
animals terrestres.

La ubicaci6n estrategica de los dormideros de los monos
arana en Costa Rica (A. ..-rr.. I con relaci6n a sus areas
de alimentaci6n, asf como el uso repetido de los arboles
dormideros, permit calificar a estos animals como forra-
jeros de multiples lugares centrales (Chapman et al., 1989).
Para la especie estudiada en el PNY (A. belzebuth) falta
informaci6n que permit concordar con dicho criteria, ya
que s6lo uno de los 15 diferentes Arboles dormideros se us6
en dos ocasiones. Esto se debe, principalmente, a la escasa
informaci6n obtenida en este studio, dada la dificultad
para seguir a los monos arana dfa tras dfa (Chapman et al.,

La categoria hembra adulta con crfa dependiente conform
el 30% del grupo estudiado (Pozo, 2001); esto explica el
mayor porcentaje de tiempo de observaci6n de la mencio-
nada categoria edad-sexo en Arboles dormideros. Ademas,
tanto en A., ..rr .., como en A. belzebuth, las hembras con
crias jovenes tienden a separarse del resto del grupo (Cant,
1995; Pozo, 2001), debido a la dificultad que represent
el acarreo de las mismas, lo que justifica que para este es-
tudio una madre con cria sean considerados como un solo
miembro del grupo (Izawa et al., 1979). El bajo porcentaje
de subgrupos durmientes compuesto por machos adults
solitarios tambidn fue encontrado en Costa Rica (Chap-
man, 1989) y seguramente esta influenciado por la compo-
sici6n social y la tendencia de agregaci6n del grupo, pues

los machos tienden a ser menos solitarios que las hembras
(Cant, 1995).

Los subgrupos durmientes presentaron perfodos de alimen-
taci6n, antes y despuds de utilizar el Arbol dormidero, lo
que coincide con lo reportado para A ... rr .., (Chapman y
Chapman, 1991) y paraA. paniscus (van Roosmalen, 1985).
Cinco de los Arboles dormideros se registraron como com-
ponentes de la dieta del mono arana (Pozo, 2001); aunque
s6lo en una ocasi6n se observ6 que una hembra adulta con
infante dependiente se aliment6 del mismo arbol donde

Finalmente, se concluye que Ateles belzebuth del PNY pre-
fiere dormir en Arboles mayores a 25 m de los bosques
altos asentados en los valles del sitio de studio; la ma-
yoria de species de los Arboles dormideros pertenecen a
las families Bombacaceae y Myristicaceae, alcanzando una
altura promedio de 28.94 my un DAP promedio de 50.43
cm. Los Arboles dormideros son utilizados por subgrupos
conformados por un mAximo de tres individuos que in-
cluyen una alta proporci6n de hembras adults con crfas

Agradecimientos: Los datos de campo fueron tomados gra-
cias al auspicio financiero de la National Science Foun-
dation (Grant # SBR 9222526) y de los fondos del De-
partamento de Anatomfa de la Facultad de Medicina de
la Universidad de Puerto Rico. Debo agradecer al Dr. J.
G. H. Cant por las muy valiosas sugerencias para mi tra-
bajo de campo en el Yasunf. Gracias tambidn al Dr. L.
AlbujaV. de la Escuela Politecnica Nacional de Quito por
avalar mi participaci6n en el Proyecto Primates Ecuador, a
los Drs. Peter S. Rodman y A. Di Fiore de la Universidad
de California en Davis (USA) por permitirme trabajar en
su Area de studio y por compartir juntos el trabajo de
campo, a la Dra. L. Arcos T. de la Pontificia Universidad
Cat6lica del Ecuador y a los trabajadores de la Estaci6n
Cientifica Yasunf por el apoyo logfstico brindado, al Dr.
0. BAez T. de la Universidad Central del Ecuador por su
supervision al analisis de los datos, al Ministerio del Am-
biente del Ecuador por otorgar los permisos de investi-
gaci6n No. 009-IC y 018-IC INEFAN/DANVS/VS, y al
Departamento de Relaciones Comunitarias de la Maxus-
YPF-Repsol por su colaboraci6n en los dialogos con la Na-
cionalidad Huaorani. Quiero expresar mi eterna gratitud a
L. Dew, D. Youlatos y B. Orengo por su ayuda en la toma
de datos. Por 6ltimo quiero agradecer al Dr. D. Youlatos,
al Ing. E. Basante y a los editors de esta revista, por las
sugerencias y cambios que hicieran al primer borrador del
present articulo.

Wilmer E. Pozo R., Escuela de Biologfa, Universidad
Central del Ecuador. Direccidn actual: Escuela Politecnica
del Ejdrcito, Facultad de Ciencias Agropecuarias (IASA),
Laboratorio de Zoologfa, Av. El Progreso s/n, PO Box
231-B, Sangolquf, Ecuador. E-mail: ec>.

Neotropical Primates 13(3), December 2005


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Vanessa Barbisan Fortes
Fabiana Cristina Alves
Juliana Arpini


The conservation of Neotropical primates is compromised
by a lack of information on their distributions (Hirsch et
al., 2002). The black-and-gold howler monkey, Alouatta
caraya (Humboldt, 1812), has a large range-from eastern
Bolivia, western Paraguay and northern Argentina to cen-
tral and southern Brazil, from the states of Piauf and Bahia
to Rio Grande do Sul (Hirsch et al., 1991). It is found
mainly in forests and woodlands in the Cerrado, Pantanal,
Chaco and Pampabiomes (Hirsch et al., 2003), in continu-
ous forest, gallery forest, and flooded forests (Brown and
Zunino, 1994; Eisenberg and Redford, 1999; Zunino et
al., 1996, 2001).

The northern limit of its distribution was recently extended
to 10S, following the discovery of groups in the Chapada

das Mangabeiras, Piauf (Flesher, 2001). Its possible occur-
rence in Uruguay (Department of Artigas, 31S) would
also extend its southern limit. The presence of this species
in the state of Rio Grande do Sul was only recorded in the
beginning of the past decade in Alegrete (Bicca-Marques,
1990), and until recently only a few populations have been
recorded from the Campanha region. Following a broad
survey of all of Rio Grande do Sul (south of Santa Cata-
rina), Codenotti and Silva (2004; Codenotti et al., 2002)
recorded A. caraya throughout its western half, in the phys-
iographic regions of Alto Uruguai (western part), Planalto
Medio (western part), Miss6es, and Campanha.

Methods and Study Area

We found Alouatta caraya during an expedition in
August-September 2003 to the municipality of Guarac-
iaba, state of Santa Catarina, Brazil. Our aim was to find a
study site and habituate a group of brown howler monkeys
(Alouattaguariba clamitans) for an ecological and behavio-
ral study, since we knew that it occurs in the Brazilian pine
forest (mixed umbrophilous forest) found in western Santa
Catarina (Klein, 1978).

Our study area was the district of Ouro Verde, about 30
km west of the city of Guaraciaba. The vegetation there
is typically seasonal semideciduous forest (Klein, 1978).
Brazilian pine (Araucaria angustifolia) is scarce, but found
in some places as this is a transition region. The area is
bisected by the Rio Maria Preta (tributary of the left bank
of the Rio Peperi-guaqu), marking the limit between the
municipalities of Guaraciaba and Sao Jose do Cedro. The
landscape is a mix of pasture and corn and bean planta-
tions, with the forest cover restricted to a few small rem-
nants, mostly on the banks of the Rio Maria Preta and
other small streams.

Results and Discussion

The first group was sighted on 25 August 2003, resting
in a Brazilian pine, in a forest fragment (2630.731'S,
5341.117'W) connected to the riparian forest of the Rio
Maria Preta. There were five of them: one adult male,
two adult females and two juveniles. Another group was
seen on 12 September 2003, in an isolated fragment
close to the first (c. 300 m). A few sparse trees provided
a scanty connection between the two forest fragments.
We were unable to count the entire group in this case,
since they spread out when they saw us. We saw an adult
male and two adult females, but the group was certainly

This is the first record of A. caraya in Santa Catarina. This
is not unexpected, however, following the surveys of Code-
notti and Silva (2004) in Rio Grande do Sul. In some areas
they found A. caraya to be sympatric-overlapping in its
range-with A. guariba, and this might well be the case
in Santa Catarina. A. guariba has been confirmed for the

Neotropical Primates 13(3), December 2005 3

Figure 1. The new locality for the black-and-gold howler monkey (Alouatta caraya), municipality of Guaraciaba, and the nearest localities
where the species has been recorded previously.

municipality of Sao Miguel do Oeste, near Guaraciaba (see
Hirsch et al., 2003).

Black-horned capuchin monkeys (Cebus nigritus) also occur
in Santa Catarina, and are sympatric with A. caraya in
the district of Ouro Verde, as they are in Tobuna (Brown
and Zunino, 1994) and Foz do Iguaqu (Parera and Bosso,
1992). We did not see them in the same forest patches,
however. The capuchin group we sighted was in an isolated
fragment (2631.015'S, 5339.970'W) about 1 km away.

The nearest municipalities in southern Brazil where black-
and-gold howlers have been recorded previously are Foz do
Iguaqu, Parana (Hirsch et al., 2003), and Cruz Alta, Rio
Grande do Sul (Codenotti et al., 2002; Codenotti and
Silva, 2004), about 160 km north and 225 km south from
Guaraciaba, respectively. Black howlers also occur at Puerto
Iguazi, Paraguay, along the Brazilian frontier (Parera and
Bosso, 1992). In Argentina, the closest record is in the mu-
nicipality ofTobuna, Misiones (Brown and Zunino, 1994),
about 50 km west from our locality (Fig. 1).

The discovery of A. caraya at this locality is not unexpected,
since it is within the known potential limit of its geographi-
cal distribution (Fig. 2). It emphasizes, however, the lack

Figure 2. Geographic distribution of black-and-gold and brown
howler monkeys (adapted from Hirsch et al, 2003). The new
locality is indicated by the arrow.

of primatological studies in the region, and the urgency
of conducting more detailed and intensive studies there in
order to achieve a better knowledge of the occurrence and
distribution of primates for their conservation in the state
of Santa Catarina.


Neotropical Primates 13(3), December 2005

Acknowledgments: We thank Jose Simionni for permission
to work in his property. We are also grateful to Jdlio Cesar
Bicca-Marques for commenting on the manuscript.

Vanessa Barbisan Fortes, Centro de Ciencias Agro-Am-
bientais e de Alimentos, Universidade Comunitaria Re-
gional de Chapec6, Rua Senador Atflio Fontana, 591E,
Chapec6 89809-000, Santa Catarina, Brazil, e-mail
, Fabiana Cristina Alves,
Curso de P6s-Graduaqao Stricto Sensu Mestrado em
Ciencias Ambientais, Universidade Comunitaria Regional
de Chapec6, and Juliana Arpini, Curso de P6s-Graduaqao
Lato Sensu em Diagn6stico Ambiental e Recuperaqao de
Areas Degradadas, Universidade Comunitaria Regional de


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of georeferenced occurrence localities of Neotropical pri-
mates. Department of Zoology / UFMG, Belo Horizonte.
htm>. Accessed 20 October 2003.
Klein, R. M. 1978. Mapa F'-.. ..- 'F.. de Santa Catarina.
Herbario Barbosa Rodrigues, Itajaf, Santa Catarina.

Parera, A. and Bosso, A. 1992. Presencia actual del mono
aullador Alouatta caraya en el extremo norte de la Pro-
vincia de Misiones, Argentina. Bol. Primatol. Lat. 3(1):
Zunino, G. E., Bravo, S., Ferreira, F M. and Reisenman,
C. 1996. Characteristics of two types of habitat and the
status of the howler monkey (Alouatta caraya) in north-
ern Argentina. Neotrop. Primates 4(2): 48-50.
Zunino, G. E., Gonzilez, V., Kowalewski, M. M. and
Bravo, S. P. 2001. Alouatta caraya. Relations among
habitat, density and social organization. Prim. Rep. 61:



Eric Bairrdo Ruivo
J. Bryan C.m i//
Alba Lucia Morales-jiminez

Considered one of the richest countries in diversity of
mammals (471 species), Colombia is regarded as the second
richest in terms of primate diversity in the Neotropics, and
the fourth in the world with approximately 39 species, six
of which are endemic (Rylands et al., 1997; Alberico et al.,
2000; Cowlishaw and Dunbar, 2000).

Primates are important for tropical forests. They are key
components in the evaluation of strategies for biodiver-
sity conservation for specific areas (Rylands et al., 1997).
Colombian primates are under great threat from hunting,
forest loss and other human activities, resulting in the des-
ignation of one species as Critically Endangered (Ateles hy-
bridus), one as Endangered (Saguinus oedipus) and five as
Vulnerable (Ateles belzebuth, Aotus lemurinus, Saguinus leu-
copus, ( .... ornatus, Lagothrix lugens) (IUCN, 2005).

The silvery-brown tamarin or titf gris, Saguinus leucopus, is
a Colombian endemic (Defler, 2003). It geographic range
is the smallest of the genus, the main reason for its status
as Vulnerable (Defler, 2003: 112). Saguinus leucopus can
be found in one regional reserve (Canon del Rio Alicante)
where, however, protection is inadequate because this re-
serve is being exploited by the communities in the vicin-
ity (A. L. Morales-Jimenez, pers. obs.). Only a few studies
have been conducted on this species, and these have fo-
cused on population density estimates, habitat use (Calle,
1992; Vargas and Solano, 1996; Poveda, 2000; Poveda and
SAnchez-Palomino, 2004; Valle, 2004), behaviour (Rueda,
2003; Del Valle, 2004) and reintroduction of pet trade ani-
mals into the wild (Garcia, 1997). There are few studies on
these animals in captivity (Alveario et al., 1985).

In Colombia, S. leucopus is maintained in at least seven
zoos, but all of them have problems with its reproduc-
tion and survival. At least 50 zoo animals have died in the
last five years. As of January 2006 no zoo had successfully

Neotropical Primates 13(3), December 2005

bred and parent-reared these animals. The single litter of
surviving young at that time were hand-reared, and most
captive-born animals die before they are one year old (A.
L. Morales-Jiminez, unpublished). Despite this history,
there are no studies of the causes of mortality in Colom-
bian zoos, but it is certainly associated with housing, stress,
inadequate nutrition and husbandry, inappropriate man-
agement of social groups and blood and intestinal para-
sites. This species is also suffering from the illegal pet trade.
There are currently (March 2006) around 100 individuals
in four rescue centres that need to be re-located, and at
least one new individual is confiscated each week (Orjuela
and Caicedo, pers. comm.). Rehabilitation and release pro-
grammes are in their infancy and the success of these so far
is not clear.

All these problems indicate that the species is under very
high pressures, and that urgent in situ and ex situ conser-
vation measures are needed to avoid its further decline,
or even extinction, in the short term. For this reason, a
conservation programme was organised in 2005 under
the umbrella of the EAZA Callitrichid Taxon Advisory
Group (TAG). It was coordinated by Lisbon Zoo (Por-
tugal), and resulted from a consortium of 17 European
zoos and local organizations, including nine Colombian
zoos, regional conservation authorities (Corantioquia and
Cornare), NGOs, Colombian universities and researchers.
The programme is co-ordinated locally by Alba Lucia Mo-
rales, Researcher at Fundaci6n BioDiversa, Colombia, that
aims to establish long term in situ and ex situ conservation
measures to protect S. leucopus. A combination of ex situ,
in situ and education projects is necessary to protect this
threatened species.

Goals of the Program

* Assess the current distribution and density of the sil-
very-brown tamarin in Colombia, in order to establish
priorities and potential areas for conservation and re-
search. Collection of local information on this species
from specimens in museum collections, literature re-
views and interviews with researchers that have worked
in Colombia. This information will be included in a
Geographical Information System (GIS) database, and
GAP analysis will be used to predict the species' po-
tential distribution. Maps will be generated showing
the historical distribution of the species and its current
range, overlaid with maps of Colombian ecosystems.

* Improve husbandry of silvery-brown tamarins under
human care in Colombia through the distribution of
EAZA husbandry guidelines, and the organisation of
workshops on captive care of callitrichids.

* Analyze information on deaths in captivity and deter-
mine causes of mortality. Zoos in ACOPAZOA (Aso-
ciaci6n Colombiana de Parques Zool6gicos y Acuari-
os) will implement a necropsy protocol for this species

to provide uniform data for comparison and analysis.
Behavioral studies will be conducted to acquire more
information about pair bonding, stress, infanticide,
aggression and nutrition, which are key factors in the
survival of this species in captivity. The programme
will provide funding support for the analyses necessary
to identify the causes and patterns of death in all cal-
litrichids under human care.

* Establish a captive-breeding programme for the silvery-
brown tamarin in Colombia (including a studbook)
and build appropriate enclosures in participating Co-
lombian zoos (three cages per zoo) where pairs will be
housed for reproduction. Support rehabilitation cen-
tres by improving their housing, husbandry and diet
and by trying to relocate animals in appropriate places,
including future reintroduction programs, or ex situ
programs if animals cannot be released.

* Continue the reintroduction of confiscated silvery-
brown tamarins into their natural habitat. CORNARE
(Corporaci6n Aut6noma Regional del Rio Negro
Nare) is the leader on this project and the silvery-
brown tamarin is their flagship species. Releases will
be conducted according to IUCN guidelines, and re-
leased animals will be monitored with radio transmit-
ters in follow-up studies.

* Evaluate the pet trade in silvery-brown tamarins and
work on solutions to end it. This project will identify
the main regions involved in the tamarin pet trade in
order to focus efforts on education and identify alter-
natives for people.

* Assess the possible existence of two subspecies of
Saguinus leucopus.

* Use education as an important tool to diminish habi-
tat destruction and the pet trade. A Festival will be
organised in order to increase the awareness of local
populations concerning this conservation issue. An ex-
hibition cage will be built in La Pintada to meet edu-
cation goals for schools in the region, to house surplus
silvery-brown tamarins not suitable for release or the
breeding program. The program will look for alterna-
tive income for local populations to help mitigate the
current exploitation of natural resources.

* Export captive-born animals from the Colombian
Zoos Breeding Program to establish the European
Breeding Program (EEP) for this species.


The Saguinus leucopus conservation project is already a start-
ing point for the conservation of this endemic callitrichid
species in Colombia, and offers solutions for animals in the
pet trade and support to zoos and rescue centers in order

Neotropical Primates 13(3), December 2005

to create an ex situ conservation program. It also supports
research and conservation efforts in Colombia and organ-
izes educational campaigns to reduce the volume of the pet
trade. This project already has many national and interna-
tional partners, and is supported by the central government
of Colombia as well as local conservation authorities such
as CORNARE (Corporaci6n Aut6noma Regional del Rio
Negro Nare), CORANTIOQUIA (Coorporaci6n Au-
t6noma Regional de Antioquia) and ACOPAZOA (Asoci-
aci6n Colombiana de Parques Zool6gicos y Acuarios).

Some important results have already been achieved since
the program started in 2005:

1. Animals housed in poor conditions in rescue centers
were relocated to adequate facilities in Colombian
2. Husbandry of this species was improved with the dis-
tribution of the EAZA Husbandry Guidelines for the
Callitrichidae (Carroll, 2002) translated into Spanish
for all Colombian zoos and rescue centers.
3. A workshop on callitrichid husbandry was organised in
January 2006. The programme supported all expenses
of this workshop, including housing and meals for all
4. Model breeding cages were built during the workshop
to be used as a template for all other breeding cages,
and two infants have already been born in these dem-
onstration cages.
5. Maps showing the present distribution of S. leucopus
are in preparation.
6. Educational poster on S. leucopus is in production and
will be sent to all Colombian zoos.
7. Breeding cages are under construction in six Colom-
bian zoos.
8. Quarantine, rehabilitation and reintroduction cages
were designed for rescue centers, including a manage-
ment protocol for S. leucopus arriving at rescue cen-
ters. This conservation program will help CORNARE
in designing and building their new rescue center at
Los Olivos.


The committed involvement of so many entities in Co-
lombia that have joined efforts to save the silvery-brown
tamarin is already a sign of success for this program. It
is most likely the first time in Colombian history that so
many different people and institutions-from private zoos
to NGOs, central and regional conservation authorities,
and private and public universities-are working together
towards a national conservation goal.

Finally, this serves as a pilot project for other species
of Saguinus, and will contribute to the management of
other threatened Colombian taxa such as S. oedipus and
S. ...* .., These species are underrepresented in Colom-
bian zoos, and it is a priority to maintain ex situ viable

populations in Colombia to contribute to their conserva-
tion. All the experience acquired from this project will help
Colombian zoos to start ex situ conservation projects for
other species.

Acknowledgments: The authors would like to express their
gratitude to the following institutions that have contribu-
ted to this conservation program: Apenheul (The Nether-
lands), Zooparc de Beauval (France), Belfast Zoo (UK),
Bristol Zoological Gardens (UK), CEPA (France), CERZA
(France), Chester Zoo (UK), Parc Zoologique de Dou&-
la-Fontaine (France), Durrell Wildlife Conservation Trust
(British Isles), Faunia (Spain), Zoo de La Palmyre (France),
Jardim Zool6gico (Portugal), London Zoo (UK), Parc Zo-
ologique et Botanique de Mulhouse (France), South Lakes
Wild Animal Park (UK), La Vallke des Singes (France),
Zodiac Zoos (The Netherlands), and in Colombia Fun-
daci6n BioDiversa Colombia, Fundaci6n Ecolombia,
Santa Cruz Zoo, Piscillago Zoo, Santafe Zoo, CAFAM,
Matecafia Zoo, Baranquilla Zoo and Cali Zoo.

Eric Bairrao Ruivo, Jardim Zool6gico de Lisboa, Estrada
de Benfica 158-160, 1500 Lisboa, Portugal, e-mail: zoolisboa.pt>, J. Bryan Carroll, Bristol Zoo Gardens, Bris-
tol, Clifton and West of England Zoological Society, Clift-
on, Bristol BS8 3HA, England, UK, e-mail: bristolzoo.org.uk> and Alba Lucia Morales-Jimenez,
Coordinadora, Programa de Conservaci6n del Titf Gris,
Fundaci6n BioDiversa Colombia, e-mail: fundacionbiodiversa.org>.


Alberico, M., Cadena, A., Hernandez-Camacho, J. and
Mufioz Saba, Y. 2000. Mamfferos (Synapsida: Theria) de
Colombia. Biota Colombiana 1(1): 43-75.
Alveario, M. C., Belcher, A., Caldwell, C., Henry, R. T.
and Epple, G. 1985. Caesarean delivery and handrear-
ing of Saguinus leucopus triplets in the laboratory- a case
report. Primate Rep. 13: 57-68.
Calle, Z. 1992. Informe de actividades y resultados: Censo
preliminary y recomendaciones para el manejo de una
poblaci6n natural de Saguinus leucopus en la zona de in-
fluencia del proyecto hydroelkctrico la Miel II. Informe
no publicado.
Carroll, J. B. (ed.). 2002. EAZA Husbandry Guidelines for
the Callitrichidae. Clifton and West of England Zoologi-
cal Society, Bristol, UK.
Cowlishaw, G. and Dunbar, R. 2000. Primate Conservation
Biology. The University of Chicago Press, Chicago.
Defler, T. R. 2003. Primates de Colombia. Conservaci6n
International, Serie de Gufas Tropicales de Campo. Con-
servaci6n Internacional Colombia, Bogoti.
Del Valle, C. M. 2004. Estudio del comportamiento social
de dos grupos de Saguinus leucopus en el bosque y la zona
urbana de Mariquita, Tolima, Colombia. Tesis de pre-
grado, Universidad Nacional de Colombia, Bogoti.

Neotropical Primates 13(3), December 2005

Garcia, R. 1997. Reintroducci6n de un grupo de titf gris
cautivo (Saguinus leucopus Orden: Primates, Familia: Cal-
lithrichidae) a un ambiente natural. Tesis de pre-grado,
Universidad de Antioquia, Medellin.
IUCN. 2005. 2005 IUCN Red List of Threatened Species.
IUCN Species Survival Commission. IUCN, Gland,
Switzerland, and Cambridge, UK. Website: www.iucnredlist.org>.
Poveda, K. 2000. Uso de habitat de dos grupos de titf de
pies blancos, Saguinus leucopus, en Mariquita, Colombia.
Tesis de pre-grado, Universidad Nacional de Colombia,
Poveda, K. and SAnchez-Palomino, P. 2004. Habitat use
by the white-footed tamarin, Saguinus leucopus: A com-
parison between a forest-dwelling group and an urban
group in Mariquita, Colombia. Neotrop. Primates 12(1):
Rueda, L. H. 2003. Comunicaci6n vocal de dos grupos
de titi gris (Saguinus leucopus) en Mariquita, Colombia.
Tesis de pre-grado, Universidad Nacional de Colombia,
Rylands, A. B., Mittermeier, R. A. and Rodrfguez-Luna, E.
1997. Conservation of Neotropical primates: Threatened
species and an analysis of primate diversity by country
and region. Folia Primatol. 66(1): 134-160.
Sanderson, E. W., Redford, K. H., Chetkiewicz, C. L. B.,
Medellin, R. A., Rabinowitz, A. R., Robinson, J. G. and
Taber, A. B. 2002. Planning to save a species: The jaguar
as a model. Conserv. Biol. 16(1): 58-72.
Valle, H. 2004. Estimaci6n del tamafio poblacional
del titf gris Saguinus leucopus Gunther, 1877 en tres
zonas del municipio de Mariquita, Departamento del
Tolima. Tesis de pre-grado, Universidad del Tolima,
Vargas, N. and Solano, C. 1996. Evaluation of the condi-
tion of two populations of Saguinus leucopus Gunther
1817 [sic], in order to determine potential conservation
areas into [sic] the Middle Magdalena, Colombia. Ab-
stracts: XVIth Congress of the International Primatologi-
cal Society and XIXth Conference of the American Society
of Primatologists, p.373. 11-16 August 1996, Madison,
Wisconsin, USA.


Hilda Tresz

Prior to 2002 the Phoenix Zoo had identified the exist-
ing otter exhibit as deficient, both in terms of husbandry
standards and the quality of public experience. Based upon
visitor feedback, it was determined that this area of the zoo
needed a reliable and more intense animal experience. The
management was looking for a new type of exhibit that
would give our guests a different perspective on animals.
The availability of an appropriate and preferably cohesive

group of monkeys was suggested and researched before the
final commitment was made to the improvement.

We contacted Dr. Ken Gold about his experience as curator
at the Apenheul Zoo in the Netherlands, and he was brought
in to consult on the feasibility of a walk-through primate
exhibit. Based on his advice, the zoo decided to build an
exhibit-to be named Monkey Village-housing squirrel
monkeys and saki monkeys together. The zoo already had
the saki monkeys in its collection, and while the exhibit
was under construction the management began searching
for a group of squirrel monkeys. This group would prefer-
ably have more females than males, a common proportion
both in the wild and in captivity.

Because the New World Primate TAG does not collabora-
tively manage common squirrel monkeys, they are not often
bred in the zoo environment. Since there were no groups of
captive squirrel monkeys that would be available within the
necessary timeframe, we chose to obtain wild-caught mon-
keys. Wild-caught monkeys would be less used to people
and more likely to maintain a larger flight distance from
our guests, an important factor for liability issues. Working
with the Bucksire Corporation, the Phoenix Zoo located
and acquired 2.10 squirrel monkeys from Guyana. They
arrived in pairs on April 29, 2003 from Bucksire Corpora-
tion's facility in Pennsylvania, and were immediately placed
under quarantine in the Phoenix Zoo's Animal Care Center
(ACC). In the ACC the animals were housed the same way
they arrived, in pairs. All animals were tattooed on their
inner left thigh and were shaved for identification.

After the 30-day quarantine, the squirrel monkeys were
transferred into the Upper Education cages (an off-exhibit
area, not open to visitors) for temporary housing. This
consisted of six cages which permitted visual and tac-
tile contact, but which did not allow the animals to have
direct access to each other. This was the perfect setting to
maintain the animals in pairs until the introductions could
begin. After consulting several times with Leo Hulsker, Pri-
mate Supervisor at the Apenheul Zoo, and with Dr. Law-
rence Williams, Assistant Professor at the Primate Research
Laboratory at the University of South Alabama, we decided
that since none of these primates had seen the new exhibit
before, did not know each other well and had not have a
chance to form subgroups yet, the best course would be to
introduce all the animals together at the same time.

Since the animals were wild-caught, it was necessary to
establish a feeding routine quickly, while they would be
accustomed to seeing people but still not overly friendly
toward them. While the animals were housed in the Upper
Education cages, we began a new behavioral management
program, which focused on teaching the animals to recog-
nize feeding time through classical conditioning. Keepers
wore orange-colored jackets while provisioning the mon-
keys, and always approached from the front of the section,
to allow the animals to see the orange jackets clearly.

Neotropical Primates 13(3), December 2005

y 7/..

Figure 1. All squirrel monkeys were uniquely shaved and tattooed
for purposes of identification. Photo by Josh Crabtree.

Figure 2. Primary trainer Tracy Fleshman, wearing an orange-
colored vest during feeding conditioning. Photo by Hilda Tresz.

Figure 3. Common squirrel monkey exhibit on the Discovery
Trail. Photo by Hilda Tresz.

At this point we were faced with a dilemma: the monkeys
needed to be moved out from the small cages into a more
natural exhibit, but Monkey Village was still under con-
struction. The keepers started renovating another exhibit,
this one open to the public, where the squirrel monkeys
could be held temporarily until Monkey Village could be
built. The keepers put an enormous effort into remodeling
the exhibit: they changed all the branches, logs and ropes,
painted all the walls with murals, provided extra hiding
places and escape routes and redesigned the night house.
By the end of June 2003 everything was ready for the

The introduction took place on July 1, 2003 at 7:00 am in
the temporary Squirrel Monkey Exhibit on the Discovery
Trail. We decided that the best approach would be to leave
the animals alone to work out their differences and to es-
tablish a social hierarchy; we would not intervene unless
a severe injury occurred. An observation program was de-
veloped to document the introduction; due to a shortage
of volunteers, we scheduled a combination of instructors,
keepers, interns and volunteers to make observations for at
least two weeks, including documentation on video.

The introduction was a complete success. The monkeys
were released through their night house area to the exhibit.
The animals spent their time cautiously looking around,
testing the mesh and branches and foraging from their feed
pans. Mating behaviors were observed and mostly were ini-
tiated by the dominant male. At first the two males spent
a considerable amount of time huddling together, but by
the second day they were seen joining some of the females.
Aggressive chasing and grabbing was minimal among both

On the second day we released our male ocelot back
into his own exhibit, directly across from the squirrel
monkeys-he had been confined out of sight during their
introduction-and the squirrel monkeys came together in
a single group when he appeared. Our hot weather also
aided us. On most mornings, the temperature rose above
90F after 8:30 am, slowing all activities down; at those
temperatures, the squirrel monkeys spent most of their
time drinking and huddling in the trees, either by them-
selves or in groups of up to six individuals. All the animals
acclimated very fast to their new environment and started
to form bonds within a couple of days.

After the animals settled down, a second training pro-
gram started. For medical and husbandry reasons the
animals had to be taught to enter the night house every
day. Wooden branches were provided to connect the night
house entrance with the exhibit for easy access, and food
was placed on it to lure the animals inside. They received
novel food items only inside the night house, to encourage
them to enter and explore. Keepers routinely entered onto
exhibit, rang a bell and distributed food items as a reward;
they gradually reduced the distance between the animals

. ty0. ,.* 0*-..r

Neotropical Primates 13(3), December 2005

and the night house, and eventually the squirrel monkeys
were only fed within.

In October the monkeys received their physical exams
and were transferred into the Monkey Village holding
area. We repeated the same night-house training program
before they were introduced to the visitors. Monkey Vil-
lage opened to the public on the weekend of November
13-14, 2004, and has been one of the zoo's most popular
exhibits ever since.

Acknowledgements: The Phoenix Zoo would like to give
special thanks to Leo Hulsker, Primate Supervisor at the
Apenheul Zoo, and to Dr. Lawrence Williams, Assistant
Professor at the Primate Research Laboratory at the Univer-
sity of South Alabama, for their advice and collaboration.

Hilda Tresz, Behavioral Management Coordinator, The
Phoenix Zoo, 455 N. Galvin Parkway, Phoenix, Arizona
85008, USA. E-mail: .


Patricia Mie Matsuo
Vanessa Boucinha


The Atlantic Forest is one of Brazil's most threatened eco-
systems, ranking among the world's top five biodiversity
"hotspots" (Mittermeier et al., 1999). Today less than 8%
of the original forest remains (Fundagao SOS Mata Atlan-
tica / INPE, 1998). Among the countless species affected
by this extreme habitat fragmentation is the golden lion
tamarin (Leontopithecus rosalia), an Endangered primate
endemic to the coastal lowland forest of the state of Rio
de Janeiro.

The range of L. rosalia is now restricted to a few forest rem-
nants in the region. According to Kierulff and Proc6pio
de Oliveira (1996), the species occurs in only seven mu-
nicipalities: Armacao dos Bdzios, Cabo Frio, Casimiro de
Abreu, Rio Bonito, Rio das Ostras, Saquarema and Silva
Jardim (Fig. 1). Together these municipalities cover an
area of 2,916,631 km2; their total human population is
approximately 327,000, served by 247 municipal schools
(IBGE, 2000). The local economy is based on agriculture,
livestock, commerce and tourism.

Two biological reserves protect the golden lion tamarin:
Pogo das Antas Biological Reserve and Uniao Biological
Reserve, both administered by IBAMA, the Brazilian Insti-
tute of the Environment and Renewable Natural Resourc-
es. Pogo das Antas, located in Silva Jardim, is the largest
remaining fragment of golden lion tamarin habitat.

Figure 1. The occurrence of the golden lion tamarin (Leontopi-
thecus rosalia) in the state of Rio de Janeiro, Brazil.

The Golden Lion Tamarin Association (Associafao Mico-
Leao-Dourado AMLD), a Brazilian non-governmental
organization, coordinates a multidisciplinary program to
conserve a viable population of golden lion tamarins in
their natural habitat. AMLD's strategy includes monitor-
ing all known populations of golden lion tamarins; work-
ing with landowners to protect smaller forest fragments for
reintroduction of zoo-born family groups; and the imple-
mentation of forest corridors connecting these fragments.
Both research and community involvement are essential to
guarantee the long-term conservation of threatened ecosys-
tems and species (Dietz and Nagagata, 1995; Padua et al.,

The AMLD recognizes that support and involvement
from local communities will be essential for the success-
ful implementation of any conservation strategy. We use
environmental education to stimulate the local population
to become more aware and active in all aspects of forest
conservation. The AMLD coordinates environmental edu-
cation efforts in Pogo das Antas Biological Reserve and the
surrounding communities.


The long-term training program started in June 2003
and ended in December 2004 with the participation of
25 teachers from 14 schools in Silva Jardim. The train-
ing program began with the basics of the Atlantic Forest:
physical characteristics of the forest, native flora and fauna,
threats and conservation actions. The activities included
presentations by researchers that conduct research in the
area, field projects in the forest (Fig. 2), and suggestions
on how teachers could carry out similar projects with their
students in forest fragments in their region.

After each topic was explored, each teacher planned his or
her own project to integrate these concepts and commu-
nity conservation actions into his or her curriculum. We
encouraged the teachers to plan activities according to the
characteristics of their school, community, students' level,
and their teaching interest. As the program developed, the

/ "A / A -i

"--, s ,' f. eAi-
-t caps

Neotropical Primates 13(3), December 2005

figure 2. field activity on the soil types ot the Atlantic forest.

teachers presented the activities they carried out in their
schools and the results they obtained, and incorporated the
feedback they received into subsequent activities.


Out of the original 25 participants, 17 teachers from
ten schools completed the entire program, consisting of
ten workshops and one seminar. These teachers devel-
oped and presented to their students more than 80 study
units on the themes of the workshops-native plants
and animals, threats, water resources, and conservation
actions-integrating subjects such as art, biology, math-
ematics and Portuguese.

The teachers worked on many conservation-themed ac-
tivities with their students, including drawing, posters,
poetry, music, games, writing projects, interviews, exhib-
its, history projects, and activities in the field. More than
90% of the teachers began conducting field exercises in
nearby forests after the second workshop, helping to im-
prove their students' awareness and appreciation of the
local environment. Similarly, these activities helped en-
hance the teachers' appreciation of the importance of con-
serving the Atlantic Forest, as well as the local problems
that affect the forest and animals such as the golden lion

The core group of teachers participated in all decisions re-
lating to the project. They also collaborated in the produc-
tion of a 2004 calendar produced by the AMLD, intended
to highlight the training program; the calendar incorpo-
rated 15 drawings about the Atlantic Forest created by the
teachers' students. In addition, the teachers helped with the
organization of the First Environmental Education Work-
shop in Silva Jardim, a one-day seminar featuring posters,
presentations and mini-courses. The teachers assisted in
the planning of the workshop and presented classes to the
workshop participants, who included 180 educators from
60 institutions from the seven municipalities where the
golden lion tamarin is found. It was an important event in

the region to promote a better understanding of the role of
education for the Atlantic Forest and for the conservation
of golden lion tamarins.

The training program provided an opportunity for the
teachers to try new ideas and teaching methodologies, to
evaluate the results, and to share their experiences with the
wider community. As a result of this first training program,
the Public Departments of Education and Environment of
the municipality of Silva Jardim signed an agreement with
the AMLD to support the teacher training program and
provide financial resources. In addition, the Department
of the Environment started its own teacher training project
in 2003, based on the official environmental curriculum
recommended by the Brazilian Ministry of Education. As
this curriculum does not specifically focus on the Atlantic
Forest, the Department of the Environment invited the
AMLD to participate in these courses, by presenting talks
on the local Atlantic Forest habitat and field activities at the
Poco das Antas Biological Reserve. This initiative by Silva
Jardim is a valuable complement to our efforts to build the
capacity of local teachers of Silva Jardim, as-owing to fi-
nancial limitations-the AMLD teacher training project is
only able to train 25 teachers per year.


Support and involvement from the local communities
will always be essential for the successful implementation
of conservation strategies for the Atlantic Forest and the
golden lion tamarin. The AMLD teacher training program
has demonstrated its ability to improve awareness of this
threatened ecosystem and its flagship primate species, and
is important to achieving sustained public support for con-
servation in the region.

The participation of the teachers in all project decisions
is essential to establish a cooperative relationship and re-
sponsibility for the project. When each teacher can plan
the content and methods used in his or her own project,
he or she feels more responsible and invests more time and
resources into it.

The partnership between the AMLD and the Public De-
partments of Education and Environment of the munici-
pality of Silva Jardim has been crucial to guaranteeing the
long-term success of our education project in that mu-
nicipality. This is the model we intend to follow when
establishing partnerships aimed at long-term implemen-
tation of environmental education in the schools of the
seven municipalities where golden lion tamarins still

Acknowledgements: We thank all the teachers who partic-
ipated in this project, the researchers that gave talks on
their studies, the participants in other AMLD programs,
the Disney Wildlife Conservation Fund, the Philadelphia
Zoological Park, the American Society of Primatologists,

Neotropical Primates 13(3), December 2005

the Copenhagen Zoo and the Departments of Educa-
tion and the Environment of the municipality of Silva

Patricia Mie Matsuo and Vanessa Boucinha, Associa-
gao Mico-Leao-Dourado, Caixa Postal 109.968, Casimi-
ro de Abreu, RJ, 28.860-970, Brazil. E-mail: micoleao.org.br>.


Dietz, L. A. and Nagagata, E. 1995. Golden Lion Tamarin
Conservation Program: A community educational effort
for forest conservation in Rio de Janeiro State, Brazil. In:
Conserving i, j. International Education and Communi-
cation Approaches, S. Jacobson (ed.), pp.64-68. Columbia
University Press, New York.

Fundacao SOS Mata Atlantica / INPE. 1998. Atlas da
Evolufao dos Remanescentes Florestais e Ecossistemas Associa-
dos no Dominio da Mata Atldntica no Periodo 1990-1995.
Fundagao SOS Mata Atlantica e Instituto Nacional de
Pesquisas Espaciais (INPE), Sao Paulo.

IBGE. 2000. Censo D. .. 'F.. 2000. Available at www.ibge.gov.br>.

Kierulff, M. C. M. and Proc6pio de Oliveira, P.
1996. Re-assessing the status and conservation of the
golden lion tamarin Leontopithecus rosalia in the wild.
Dodo, Journal of the '1 --j. Preservation Trust 32:

Mittermeier, R. A., Myers, N., Gil, R. P. and Mittermeier,
C. G. 1999. Hotspots: Earth's 1:; ... Richest and Most
Endangered Terrestrial Ecoregions. Conservation Interna-
tional and CEMEX, Cidade de Mexico.

Padua, S. M., Dietz, L. A., Souza, M. G. and Santos, G. R.
2002. In situ conservation education and the lion tamarins.
In: Lion Tamarins Biology and Conservation, D. G. Klei-
man and A. B. Rylands (eds.), pp.271-282. Smithsonian
Institution Press, Washington, DC.


In December 2005, Eliana Ruth Steinberg defended her
undergraduate thesis (tesis de licenciatura) analyzing
sex chromosomes and sperm morphology in the squirrel
monkey, Saimiri boliviensis. It was presented at the School
of Exact and Natural Sciences at the University of Buenos
Aires (FCEyN UBA), Argentina. Her supervisor was
Marta Dolores Mudry, Associate Professor in the Depart-
ment of Ecology, Genetics and Evolution. This research
was funded by MDM CONICET PIP 2450, UBACyT

X031 and ERS Sigma Xi Grant 3040277. The following is
a summary of her thesis.

Accurate systematic information is essential to mammal
conservation, both in captivity and in the wild, and taxo-
nomic, phylogenetic and biogeographic analyses are in-
dispensable tools. The geographic origin of many animals
held in zoos and conservation breeding centers is often un-
known, obscured by the exchange of individuals between
institutions. The challenge, then, is to generate accurate
identifications for animals with uncertain provenance.

Phenotypic and morphometric characters have been tradi-
tionally employed for systematic diagnosis, but the wide
variety of phenotypes and polymorphisms may cause con-
fusion when designing programs for ex situ maintenance.
This may result in the unintentional assembly of mixed
groups, with the possible consequence of loss of variabil-
ity due to endogamic depression. In this context, genetic
tools such as karyotyping and DNA analysis take on a new
importance, and have been employed in the last decade
in many breeding centers. Mitotic parameters in particu-
lar have been widely used in the past decade to accurately
typify individuals. However, only meiotic data allow a con-
firmation of sex determination, and are thus essential for
guiding breeding efforts.

Sperm from different species of mammals may clearly differ
in their morphology and dimensions. This is important for
the successful implementation of biotechnological tech-
niques, such as cryopreservation of gametes and assisted
reproduction, in ex situ conservation programs in zoos and
in breeding and research facilities. In this thesis, cytoge-
netics and sperm morphology were used for the first time
for taxonomic diagnosis in the Ceboidea, using the squirrel
monkey, Saimirisp., as an experimental model.

Species and subspecies of squirrel monkeys may be distin-
guished by their coat coloration, but this is often difficult
for the untrained eye and may be ambiguous in backcrossed
animals. However, species of Saimiri differ in their karyo-
types by pericentric inversions, resulting in differences in
the ratio of acrocentrics to non-acrocentrics. This allows a
clear identification of species and hybrids both in captivity
and in the wild. This study examined male squirrel mon-
keys held at the Corrientes Biological Station (EBCo) and
females from the Buenos Aires Zoo and the C6rdoba Zoo,
all in Argentina.

This study found a diploid number of 2N = 44, XX/XY,
in all specimens analyzed, with a fundamental number
FN = 75. The karyotype included five metacentric chro-
mosome pairs, 10 submetacentric pairs and six acrocentric
pairs. The X chromosome is submetacentric and the Y is a
small acrocentric. In the meiotic analysis, the presence of
22 bivalents with a distinctive XY bivalent was observed
in late diakinesis/Met I. The G banding patterns agreed
with those previously published for S. boliviensis boliviensis.

Neotropical Primates 13(3), December 2005

As a result of this analysis, the animals held at the Buenos
Aires and C6rdoba Zoos, previously assigned to Saimiri
sciureus, are now recognized as S. b. boliviensis. In addition,
the morphometric characterization and comparison with
other cebids demonstrated that the spermatozoa of Saimiri
b. boliviensis are distinguished by a larger midpiece. The
different variables analyzed here support genetic charac-
terization in this and other New World primate genera in
the "total evidence" framework accepted for modern taxo-
nomic studies.

Lic. Eliana Ruth Steinberg, Grupo de Investigaci6n en
Biologfa Evolutiva gibeE), Departamento de Ecologfa,
Genetica y Evoluci6n, Facultad de Ciencias Exactas y Nat-
urales, UBA CONICET, Ciudad Universitaria, Pabel-
16n II, 4to. Piso, Lab 46 (5411), Buenos Aires, Argentina.
E-mail: .


Steinberg, E. R. 2005. Cytogenetics and sperm morphol-
ogy as tools of systematic value: The example of Saimiri
boliviensis (Primates: Platyrrhini). Undergraduate thesis,
University of Buenos Aires, Argentina.


O Institute de Desenvolvimento Sustentavel Mamiraud,
uma unidade de pesquisas do Ministerio da Ciencia e
Tecnologia (IDSM-OS/MCT), lancou a revista eletronica
Uakari. Uakari foi concebida para a publicaqao de resul-
tados de pesquisas originals em conservacao da biodiversi-
dade e uso sustentavel e participativo dos recursos naturais
do biota amazonico, e se dedica especialmente na publica-
qao dos resultados das pesquisas desenvolvidas nas Reser-
vas de Desenvolvimento Sustentavel Mamiraua e Amana,
ou mesmo em outros sftios amaz6nicos corn o apoio ou
patrocfnio do IDSM. Deste modo, Uakari public textos
de ciencias biol6gicas e ambientais em geral, e tambdm de
ciencias humans relacionadas a questao da conservacao da
biodiversidade amaz6nica.

Os trabalhos publicados em Uakari devem se enquadrar
numa das duas categories seguintes: 'Artigos cientificos' e
'Notas curtas.' (Na 6ltima categoria tambdm se admitem
as listas de levantamentos taxonomicos, e as series hist6ri-
cas de dados ambientais ou sociais ligados a conservacao da
biodiversidade.) Uma versao fisica, em papel, tambdm sera
produzida, em baixa tiragem, exclusivamente para envio e
dep6sito em algumas bibliotecas de referencia.

O primeiro ndmero (Ano 1, Ndmero 1, Novembro de
2005), inclui os seguintes artigos: Frugivoria e dispersao de
sementes por peixes na Reserva de Desenvolvimento Sus-
tentavel Aman L. Lopes de Souza; Participacao comuni-
taria na preservacao de praias para reproducao de quelonios
na Reserva de Desenvolvimento Sustentavel Mamiraud,

Amazonas, Brasil A. F Terin; Comportamento reprodu-
tivo das mulheres ribeirinhas do Amand E. A. F Moura;
O manejo sustentavel das areas alagaveis daAmaz6nia Cen-
tral e as comunidades de herbaceas aquaticas M. T. F Pie-
dade, J. Schoengart e W. J. Junk; Richness and abundance
of gall-forming insects in the Mamiraua vdrzea, a flooded
Amazonian forest G. R. Juliao, E. M. Venticinque e G.
W. Fernandes; Um ensaio sobre a adaptacao de Pygocentrus
nattereri a variaqao sazonal das aguas do Lago Mamiraua
- Reserva do Desenvolvimento Sustentavel Mamiraua -
M. Camargo e H. Queiroz; Ritmo de atividade diaria de
Osteoglossum bicirrhosum (Peixes: Osteoglossiformes) em
quatro lagos da Reserva de Desenvolvimento Sustentavel
Mamiraua, AM R. Chaves, M. Camargo, H. Queiroz e
A. Hercos. Para maiores informafoes: br/uakari>.


On 20 October 2005, Brazil's Ministry of the Environ-
ment announced the creation of two protected areas in the
state of Santa Catarina. The Araucarias National Park of
12,481 ha and the Mata Preta Ecological Station of 6,563
ha are both of great importance in protecting remnant for-
ests of the Parana or Brazilian pine, Araucaria angustifolia
(Araucariaceae). The Araucaria pine forest of southern and
southeastern Brazil, a distinctive formation of the Brazil-
ian Atlantic forest, once covered more than 20 million ha,
but-devastated by the timber industry and clear-cut-
ting-is today reduced to remnants covering a mere 2%
of that area. The Brazilian pine has been on the Brazilian
threatened species list since 1992. Source: SBS Sociedade
Brasileira de Silvicultura, website: , ac-
cessed 25 October 2005.


The National Institute on Aging (NAI) and the Wisconsin
National Primate Research Center are proud to announce
the release of the internet Primate Aging Database (iPAD),
a new tool for researchers interested in studying biomarkers
of aging in nonhuman primates.

The iPAD is a multi-centered, relational database of bio-
logical variables in aging, captive nonhuman primates.
iPAD provides an invaluable veterinary and clinical re-
source capable of generating normative values from over
half a million data points drawn from 18 different species,
including representatives of Aotus, C .-'. (... Le-
ontopithecus, Saguinus and Saimiri. These data were drawn
from routine screenings of healthy animals at regular inter-
vals throughout their lifetimes, and include blood chem-
istry, hematology, and body weight. Users of the database
are able to access individual data points at the species-, bio-
marker- or site-specific level, and are able to export and
manipulate the data at will.

Neotropical Primates 13(3), December 2005

The iPAD database has already resulted in several papers
published in peer-reviewed journals, and its usefulness
increases as new datasets are contributed. iPAD is now
expanding the number of species and measurements in
the database, and augmenting the number of data points
for existing species. Researchers interested in becoming
a part of this collaborative effort and contributing data
from healthy, non-experimental primates are encouraged
to contact the System Administrator, Wendy Newton, at
for more information. The
iPAD database is available online at wisc.edu>.


The InternationalJournal of Biological Sciences is an open-
access general biological journal which publishes peer-
reviewed scientific papers in all areas of the biological
sciences. Types of articles include regular research papers,
reviews, short research communications, viewpoints and
commentaries. In addition to the mainstream areas of bi-
ology, the Journal encourages articles from emerging fields
or cross-disciplinary with mathematics, physics, chemis-
try, engineering, and computer science. The Journal will
publish special issues that focus on topics of common in-
terest from time to time.

The Journal aims at rapid publication of high-quality re-
search results while maintaining a rigorous peer-review
process. The Journal is committed to open access to its
online version in order to maximize the distribution of
articles. Full texts of published articles appear in PubMed
Central, the U.S. National Institutes of Health (NIH) dig-
ital archive of biomedical journal literature, and abstracts
are indexed in PubMed.

Prospective authors should use the Journal's online sub-
mission system to facilitate rapid review and turnaround
time. Only PDF or Word files will be accepted. The entire
manuscript must be submitted as a single PDF or Word
file. Figures and tables should be embedded in the PDF or
Word file of the manuscript where they will appear, and
not separated as individual files. Submission implies that
the work is not previously published or under considera-
tion for publication elsewhere, that the manuscript has
been approved by all co-authors, and that the authors agree
to all terms of the copyright and publishing agreement
of the Journal. The manuscript's first page should include
the title of the article, the author's name(s), affiliations)
with complete addresses, corresponding author's email
address, phone/fax number, an abstract of not more than
250 words, and a set of up to 6 keywords after the ab-
stract. The author may suggest two to three potential ref-
erees who work in a closely related area and who can make
expert evaluation of the manuscript. Detailed information
may be found at .


Researchers at the Universidad Veracruzana (UV), Uni-
versidad de Buenos Aires (UBA) and the University of
Michigan (UM) are conducting a collaborative project on
the genetics of Neotropical primates at the northern and
southern edges of their distributions. The main aim of this
project is to combine expertise in ecological, genetic and
behavioral approaches to understand the taxonomy and
evolution of primates in these regions.

This complementary effort started in 2001, when Marta
D. Mudry visited field sites in Mexico with members of
the UV primate research group, collecting samples from
Alouatta palliata and Ateles .'. r.'. as part of a cytoge-
netic project on atelid taxonomy. Dr. Mudry described the
karyotypes of these species and identified the presence of
different patterns in sex chromosomes from the samples
obtained during this trip (Mudry et al., 2001; Nieves and
Mudry, 2001; Nieves, 2003; Nieves et al., 2005; Solari and
Rahn, 2005). At the same time, Liliana Cortes-Ortiz was
using molecular data to examine the phylogenetic relation-
ships and biogeography of the genus Alouatta (Cortes-Ortiz
et al., 2003), as well as characterizing the hybridization be-
tween Mexican howler monkey species and studying the
phylogeography of A. palliata in Mesoamerica (Cortes-
Ortiz, 2003). In addition, Ernesto Rodrfguez-Luna, Do-
mingo Canales-Espinosa, and Francisco Garcia-Orduna
had initiated a long-term study to understand the current
distributions of Mexican primates, as well as the sympatry
of A. palliata and A. pigra in the state of Tabasco, Mexico
(Garcfa-Ordufa et al., 1999, 2005; Rodrfguez-Luna et al.,

For over 25 years, researchers at UV have studied the ecol-
ogy, physiology and behavior of Mexican primates, includ-
ing A. palliata mexicana (see Appendix I), A. pigra (e.g.,
Gardca-Ordufa et al., 1999, 2005; Rodrfguez-Luna et al.,
2001) and Ateles ..rr..i (see Appendix II), and have de-
veloped and implemented strategies for their conservation
(see Appendix III). Over approximately the same period,
researchers of GIBE (Grupo de Investigaci6n en Biologfa
Evolutiva) of UBA have analyzed the cytogenetics of sev-
eral Neotropical primates, including Alouatta caraya, Aotus
azarae, Ateles spp., ( .- jacchus, Cebus jc/l/a, and
Saimiri boliviensis (see Appendix IV for a complete listing
by species). The GIBE team has used heterochromatic pat-
terns and chromosome rearrangements to elucidate the tax-
onomy of New World primates (see Appendix IV). More
recently, researchers at GIBE have also used biochemical
electrophoresis (Szapkievich et al., 1998; Szapkievich,
2000; Szapkievich and Mudry, 2003) and other molecular
techniques to characterize populations and to explore the
phylogenetic relationships of some of these taxa (Ascunce
et al., 2002, 2003a, 2003b). Finally, Liliana Cortes-Ortiz,

Neotropical Primates 13(3), December 2005

formerly part of the UV group and now affiliated with UM,
is continuing her studies on the systematics and phylogeog-
raphy of Mesoamerican primates, the genetic characteriza-
tion of a hybrid zone between A. palliata and A. pigra, the
evolution of reproductive isolation in howler monkeys, and
the application of molecular approaches to the understand-
ing of primate behavior.

These researchers from UV, UBA and UM are now joining
efforts to study the taxonomy and evolution of Neotropical
primates using molecular, cytogenetic, ecological and be-
havioral approaches. Through this collaboration, research-
ers are currently investigating the heterochromatic pattern
and sex chromosome system of A. pigra- the only species
of Alouatta whose karyotype has never been studied. This
cytogenetic work will be complemented by morphological
and molecular characterizations of each sampled individual
to verify their identification to species.

These molecular and cytogenetic studies will improve our
understanding of the origin and evolution of the excep-
tionally diverse chromosomal arrangements that occur in
howler monkeys (see for example, Ma et al., 1975; Lima
and Seuanez, 1991; Consigliere et al., 1996; Rahn et al.,
1996; Mudry et al., 2001; Oliveira et al., 2002; Rahn and
Solari, 2005). In addition, the cytogenetic study of A. pigra
will provide a foundation for the comparison of karyotypes
of hybrid and backcrossed A. palliata I A. pigra individuals
against the typical karyotype of each parental species.

We expect that this multidisciplinary approach will con-
tribute to the understanding of primate evolution in the
Neotropics, as well as establishing a basis for primate con-
servation in this region. Furthermore, this collaborative
project can be a model for combining efforts, expertise, and
resources to study primates and other organisms in Latin

Liliana Cort6s-Ortiz, Museum of Zoology and Dept. of
Ecology and Evolutionary Biology, University of Michigan,
1109 Geddes Ave., Ann Arbor, MI 48103, USA, e-mail:
, Domingo Canales-Espinosa, Er-
nesto Rodriguez-Luna, Francisco Garcia-Orduna, Insti-
tuto de Neuroetologfa, Universidad Veracruzana, A.P. 566,
C.P. 91000 Xalapa, Veracruz, Mexico, Mariela Nieves,
Eliana Steinberg and Marta D. Mudry, Grupo de Inves-
tigaci6n en Biologfa Evolutiva gibeE), Dpto. de Ecologfa,
Genetica y Evoluci6n, Facultad de Ciencias Exactas y Nat-
urales, Universidad de Buenos Aires, Ciudad Universitaria,
Pabell6n II, 4to Piso, Lab. 46-47 (EHA1428), Buenos
Aires, Argentina.


Ascunce, M. S., Hasson, E. and Mudry, M. D. 2002. De-
scription of the cytochrome c oxidase subunit II gene in
some genera of New World monkeys (Primates, Platyr-
rhini). Genetica 114: 253-267.

Ascunce, M. S., Oklander, L. and Mudry M. D. 2003a.
Amplification of mitochondrial COII gene from DNA
extracted from hair samples in some species of New
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Ascunce, M. S., Hasson, E. and Mudry, M. D. 2003b.
COII: A useful tool for inferring phylogenetic relation-
ships among New World monkeys (Primates, Platyrrhi-
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Boscarol, G., Piazza, S., Pizzigalli, C., Falcone, I., Gamba,
M., Martinoli, L., Fiore, R., Rodriguez-Luna, E. and
Giacoma, C. 2004. Vocal repertoire of Alouatta palliata
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and Garcia, F 1987. Undernourishment and parasitism
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Appendix I: Universidad Veracruzana studies on
Alouatta palliata mexicana.
Boscarol et al., 2004; Canales-Espinosa, 2000; Canales-
Espinosa et al., 1987, 1992, 2002; Canales-Espinosa and
Garcia-Ordufa, 1993, 2001, 2005; Canales-Espinosa and
Hermida-Lagunes, 1998; Carrera-Sinchez et al., 2003;
Cortes-Ortiz, 1998; Cortes-Ortiz et al., 1993, 1994;
Cortes-Ortiz and Martinez-Morales, 1996; Crist6bal-
Azkarate et al., 2005; Dias and Rodriguez-Luna, 2003,
2005; Domingo-Balcells et al., 2003; Garcfa-Orduia and
Canales-Espinosa, 1995; Jones and Cortes-Ortiz, 1998;
Rodriguez-Luna, 1997, 2000; Rodriguez-Luna et al., 1995,
2003; Serio-Silva and Rodriguez-Luna, 1994, 1995.

Appendix II: Universidad Veracruzana studies on Ateles
Cortes-Ortiz and Rodriguez-Luna, 1997; Garcia-Orduna
et al., 1993, 2002; Hernindez-Salazar et al., 2001, 2003;
Laska et al., 1996, 2000a, 2000b, 2003; Rodas et al., 2002;
Rodriguez-Luna, 1991; Rodriguez-Luna et al., 1996; Silva-
L6pez et al., 1986, 1988.

Appendix III: Universidad Veracruzana studies on pri-
mate conservation.
Canales-Espinosa, 1997; Cortes-Ortiz et al., 1996; Garcia-
Ordufa et al., 1987; Rodriguez-Luna et al., 1987a, 1987b,
1993a, 1993b, 1996a, 1996b, 1996c; Rodriguez-Luna and
Cortes-Ortiz, 1994, 1995a, 1995b; Rodriguez-Luna and
Dominguez-Dominguez, 2002; Rylands and Rodriguez-
Luna, 2000.

Neotropical Primates 13(3), December 2005

Appendix IV: Universidad de Buenos Aires cytogenetic
studies on Neotropical primates.
Alouatta caraya: Fundia et al., 2000; Mudry, 1983; Mudry
et al., 1992, 1994, 1998; Aotus azarae: Mudry, 1983;
Mudry et al., 1982c, 1984, 1990; Mudry and Galliari,
1985; Ateles spp.: Nieves, 2003; Nieves et al., 2003, 2005;
( .- jacchus: Mudry etal., 1981, 1982a, 1990; Cebus
a,'c//a: Fundia et al., 1987; Mantec6n et al., 1984; Mar-
tinez, 2003; Martinez et al., 2002, 2004; Mudry, 1983,
1990; Mudry et al., 1984, 1985, 1990; Mudry and Sla-
vutsky, 1987; Mudry and Vinuesa, 1988; PonsA Fontanals
et al., 1995; and Saimiri boliviensis: Fundia et al., 2000;
Garcia et al., 1995; Mudry et al., 1982b, 1990; Slavutsky
and Mudry, 1990; Steinberg et al., 2003, 2004, 2005.


Commensalism and Conflict: The Human-Primate In-
terface, edited by James D. Paterson and Janette Wallis.
2005. American Society of Primatologists. 504pp. ISBN:
0965830136 (hardback, $60.00). Contents: Foreword L.
D. Wolfe; Preface J. D. Paterson; Perceptions of pests:
Human attitudes to primates, conflict and consequences
for conservation P. C. Lee and N. E. C. Priston; By-
product mutualism: Conservation implications amongst
monkeys, figs, humans, and their domesticants in Hon-
duras F. Burton and A. Carroll; People, crops, and pri-
mates: A conflict of interests C. M Hill; Human social
issues, disease, and sympatric apes in the Central African
Republic A. A. Lilly; Residents and immigrants: Reac-
tions and perceptions of crop raiding in Masindi District,
Uganda J. Paterson; Primates in the forest: Sakalava
ethnoprimatology and synecological relations with black
lemurs at Ambato Massif, Madagascar I. C. Colquhoun;
Tanala synecological relations with lemurs in southeastern
Madagascar P. C. Wright, B. R. Andrianamihaja and S.
Raharimiandra; Tourist impacts on the behavior of black
howling monkeys (Alouatta pigra) at Lamanai, Belize A.
Treves and K. Brandon; Monkey forests and human land-
scapes: Is extensive sympatry sustainable for Homo sapiens
and Macacafascicularis on Bali? -A. Fuentes, M. Southern
and K. Gde Suaryana; Characterizing primate pet owner-
ship in Sulawesi: Implications for disease transmission L.
Jones-Engel, M. Schillaci, G. Engel, U. Paputungan and
J. Froehlich; Farmer attitudes toward the conservation of
"pest" monkeys: The view from Nepal M. K. Chalise and
R. L. Johnson; Rhesus commensalism in India: Problems
and prospects C. H. Southwick, I. Malik and M. Farooq
Siddiqi; City monkeys (Macaca mulatta): A study of
human attitudes- A. Srivastava and F. Begum; Coexistence
of bonnet monkeys (Macaca radiata radiata Geoffroy) with
planters in the cardamom (Elettaria cardamum Maton) and
coffee (Coffea arabica L.) plantations of Karnataka, South

India: Hospitable or hostile? A. K. Chakravarthy and N.
E. Thyagaraj; Habitat destruction, population compression
and overbrowsing by the Zanzibar red colobus monkey
(Procolobus kirkii) K. S. Siex; Round up the usual sus-
pects conflict between monkeys and farmers in East and
West Africa -J. Chism; The population status of the ursine
colobus (Colobus vellerosus) at Boabeng-Fiema, Ghana T.
Saj, J. Teichroeb and P. Sicotte; Tibetan macaques, visitors,
and local people at Mt. Emei: Problems and countermeas-
ures Qi-Kun Zhao; Recent expansion of the range of
Japanese macaques, and associated management problems
- K. Watanabe and Y. Muroyama; Applied conditioned
taste aversion and the management and conservation of
crop-raiding primates D. L. Forthman, S. C. Strum and
G. M. Muchemi; Managing monkeys and mangos M.
Baker and A. Schutt. Online ordering only; more infor-
mation available at volume4.html>.

TheLaboratory Primate, edited by SoniaWolfe-Coote. 2005.
Academic Press, New York. 650pp. ISBN 0120802619
(hardback, $200.00). A volume in the Handbook of Ex-
perimental Animals series, The Laboratory Primate details
the past and present use of primates in biomedical research,
and the husbandry, nutritional requirements, behaviour,
and breeding of each of the commonly used species. Practi-
cal information on regulatory requirements, not available
in other texts, is also covered. Sections on experimental
models cover the major areas of biomedical research, in-
cluding AIDS, cancer, neurobiology and gene therapy;
assisted reproductive technology, tissue typing, and mini-
mum group sizes for infectious disease/vaccine studies are
also included. Several chapters deal with the husbandry and
biomedical applications of New World primates. Contents:
Part 1: Definition of the primate model. 1.1. The taxon-
omy of primates in the laboratory context C. Groves;
1.2. Similarities of non human primates to humans G.
M. Miller and B. K. Madras; 1.3. General anatomy L.
R. Godfrey; 1.4. Pathology: 1.4.1. Non infectious diseases
- A. D. Lewis and L. M. A. Colgin; 1.4.2. Common viral
infections N. W. Lerche; 1.4.3. Modelling parasitic dis-
eases in non human primates: Malaria, Chagas disease and
filariasis M. T. Philipp and J. E. Purcell; 1.5. Reproduc-
tion: Definition of a primate model of female fertility A.
Einspanier and M. A. Gore; 1.6. Male reproduction and
fertilization R. M. Harrison and H. M. Kubisch; 1.7.
Primate natural history and social behaviour: Implications
for laboratory housing C. K. Lutz and M. A. Novak. Part
2: Primate Management. 2.1. New World Primates: 2.1.1.
Husbandry and management: Marmosets and tamarins S.
Rensing and A.-K. Oerke; 2.2. Old World Primates: 2.2.1.
Practical care and management of macaques K. Terao;
2.2.2. Vervet monkey breeding J. Seier; 2.3. Nutrition
and nutritional diseases S. M. Lewis, C. E. Hotchkiss and
D. E. Ullrey; 2.4. Environmental enrichment and refine-
ment of handling procedures V. Reinhardt; 2.5. Develop-
ment of specific pathogen free (SPF) non human primate
colonies K. Mansfield; 2.6. Medical care J. Mahoney.

Neotropical Primates 13(3), December 2005

Part 3: Current Uses in Biomedical Research. 3.1. Factors
affecting the choice of species H. Weber; 3.2. Techniques
and Procedures: 3.2.1. Anaesthesia- S. Unwin; 3.2.2. Rigid
endoscopy J. W. Fanton; 3.2.3. Ultrasound imaging in
rhesus (Macaca mulatta) and long-tailed (Macaca fascicula-
ris) macaques: Reproductive and research applications A.
F Tarantal; 3.2.4. Functional magnetic resonances imaging
in conscious marmoset monkeys: Methods and applications
in neuroscience research C. F Ferris and C. T. Snowdon;
3.2.5. Radiographic imaging of non human primates C.
R. Valverde and K. Christe; 3.2.6. Imaging: Positron Emis-
sion Tomography (PET) S. Chefer. 3.3. Current Uses in
Biomedical Research: 3.3.1. The use of the primate model
in research W. R. Morton, K. B. Kyes, R. C. Kyes, D. R.
Swindler and K. E. Swindler; 3.3.2. Chronic diseases B.
Hart, M. Losem, H. P. M. Brok and M. H. de Baets; 3.3.3.
Practical approaches to pharmacological studies in non
human primates F. H. Koegler and M. A. Cowley; 3.3.4.
Non human primate models of human aging X. Tigno, J.
M. Erwin and B. C. Hansen; 3.3.5. Primate models of neu-
rological disease C. A. Szabo; 3.3.6. Genetics: A survey
of non human primate genetics, genetic management and
applications to biomedical research J. Rogers; 3.3.7. The
respiratory system and its use in research C. G. Plopper
and J. R. Harkema; 3.3.8. Reproduction Male G. van
der Horst; 3.3.9. Reproduction Female W. R. Duke-
low; 3.3.10. The baboon as an appropriate model for study
of multifactoral aspects of human endometriosis J. M.
Mwenda, C. M. Kyama, D. C. Cahi, S. Debrock and T.
M. D. Hooghe; 3.3.11. Virology research P. Barry, M.
Marthas, N. Lerche, M. B. McChesney and C. J. Miller;
3.3.12. Parasitic diseases of non human primates J. E.
Purcell and M. T. Philipp. Available fom: Academic Press
/ Elsevier Inc., 30 Corporate Drive, Suite 400, Burlington,
MA 01803, USA, Tel.: 1-781-221-2212, Fax: 1-781-221-
1615. Website: .

Parenting for Primates, by Harriet J. Smith. 2005. Har-
vard University Press, Cambridge, Massachusetts. 394pp.
ISBN 0674019385 (hardback, $29.95). In this natural
history of primate parenting, Smith compares parenting
by nonhuman and human primates. In a narrative rich
with vivid anecdotes derived from interviews with pri-
matologists, from her own experience breeding cotton-
top tamarin monkeys for over thirty years, and from her
clinical psychology practice, Smith describes the thousand
and one ways that primate mothers, fathers, grandparents,
siblings, and even babysitters care for their offspring, from
infancy through young adulthood. Smith learned the hard
way that hand-raised cottontop tamarins often mature
into incompetent parents. Her observation of inadequate
parenting by cottontops, plus her clinical work with trou-
bled human families, sparked her interest in the process of
how primates become "good-enough" parents. The story of
how she trained her tamarins to become adequate parents
lays the foundation for discussions about the crucial role of
early experience on parenting in primates, and how certain
types of experiences, such as anxiety and social isolation,

can trigger neglectful or abusive parenting. Smith reveals
diverse strategies for parenting by primates, but she also
identifies parenting behaviors crucial to the survival and
development of primate youngsters that have stood the
test of time. Available from: Harvard University Press, 79
Garden Street, Cambridge, MA, 02138 USA, Tel.: 1-800-
405-1619, Fax 1-800-406-9145. Website: hup.harvard.edu>.

Primate Cytogenetics, edited by Stefan Miiller. S. Karger,
Basel. 2005. 268pp. ISBN 3805578601 $111.00 (hard-
back). Primate molecular genetics, cytogenetics and genom-
ics currently form a highly dynamic field of research, largely
due to the insight that many aspects of human genome
functioning can be better understood in light of the con-
servation of, and changes in, genome organization during
the course of evolution. Studies of our closest phylogenom-
ic relatives, the nonhuman primates, can provide detailed
information on the molecular mechanisms that shape the
human genome. Including review articles and original
investigations, this single-topic issue of Cytogenetic and
Genome Research bundles various different perspectives and
provides a broad overview of the present knowledge about
molecular cytogenetics, genome organization and evolution
of primates. It will be of great interest to researchers and
geneticists in the fields of primatology, anthropology, mo-
lecular phylogeny, evolution, human genetics and genome
research. Of particular interest is its emphasis on the genet-
ics of New World primates, with papers involving Alouatta,
Ateles, Brachyteles, Cebuella and ( .-'. Contents: Preface.
Part I: Comparative Genomics and Molecular Evolution.
Conservation genomics: Applying whole genome studies
to species conservation efforts 0. A. Ryder; Evolution of
hominoids and the search for a genetic basis for creating hu-
manness N. Saitou; The dynamic nature and evolutionary
history of subtelomeric and pericentromeric regions S. K.
Mewborn, C. Lese Martin & D. H. Ledbetter; Primate phy-
logeny: Molecular evidence from retroposons J. Schmitz,
C. Roos & H. Zischler; The primates of the Neotropics:
Genomes and chromosomes H. N. Seuanez, C. R. Bon-
vicino & M. A. M. Moreira; Evolutionary implications of
pericentromeric gene expression in humans J. M. Mudge
& M. S. Jackson; Tandem insertions of Alu elements M.
El-Sawy & P Deininger; Identity by descent and DNA se-
quence variation of human SINE and LINE elements A.
H. Salem, D. A. Ray & M. A. Batzer; Molecular evolution
of the human chromosome 15 pericentromeric region D.
P. Locke, Z. Jiang, L. M. Pertz, D. Misceo, N. Archidi-
acono & E. E. Eichler; Nucleotide sequence comparison of
a chromosome rearrangement on human chromosome 12
and the corresponding ape chromosomes M. K. Shimada,
C.-G. Kim, T. Kitano, R. E. Ferrell, Y. Kohara & N. Saitou;
Breakpoint analysis of the pericentric inversion between
chimpanzee chromosome 10 and the homologous chromo-
some 12 in humans H. Kehrer-Sawatzki, C. A. Sandig,
V. Goidts & H. Hameister; Genomic structure and paralo-
gous regions of the inversion breakpoint occurring between
human chromosome 3pl2.3 and orangutan chromosome

Neotropical Primates 13(3), December 2005

2 Y. Yue, B. Grossmann, E. Tsend-Ayush, E Griitzner,
M. A. Ferguson-Smith, E Yang & T. Haaf; Cytochrome b
polymorphisms and population structure of two species of
Alouatta (Primates) EF. E Nascimento, C. R. Bonvicino, E
C. D. da Silva, M. P. C. Schneider & H. N. Seuanez. Part
II: Comparative Molecular Cytogenetics and Chromosome
Evolution. The impact of chromosome sorting and paint-
ing on the comparative analysis of primate genomes M.
A. Ferguson-Smith, F. Yang, W Rens & P C. M. O'Brien;
Origins of primate chromosomes as delineated by Zoo-
FISH and alignments of human and mouse draft genome
sequences L. Froenicke; Fluorescence in situ hybridization
to chromosomes as a tool to understand human and pri-
mate genome evolution J. Wienberg; Evolutionary con-
served chromosomal segments in the human karyotype are
bounded by unstable chromosome bands A. Ruiz-Herre-
ra, E Garcia, L. Mora, J. Egozcue, M. Ponsa & M. Garcia;
Reciprocal painting between humans, De Brazza's and patas
monkeys reveals a major bifurcation in the Cercopithecini
phylogenetic tree R. Stanyon, R. Bruening, G. Stone, A.
Shearin & E Bigoni; Phylogenetic inferences of Atelinae
(Platyrrhini) based on multi-directional chromosome paint-
ing in Brachyteles arachnoides, Ateles paniscus paniscus and
Ateles b. marginatus- E. H. C. de Oliveira, M. Neusser, J.
C. Pieczarka, C. Nagamachi, I. J. Sbalqueiro & S. Muller;
Investigation of marmoset hybrids (Cebuella pygmaea x Cal-
lithrix jacchus) and related Callitrichinae (Platyrrhini) by
cross-species chromosome painting and comparative ge-
nomic hybridization M. Neusser, M. Munch, G. Anzen-
berger & S. Muller; Application of molecular cytogenetics
for chromosomal evolution of the Lemuriformes (Prosim-
ians) S. Warter, M. Hauwy, B. Dutrillaux & Y. Rumpler;
Evolutionary breakpoint analysis on Y chromosomes of
higher primates provides insight into human Y evolution -
R. Wimmer, S. Kirsch, G. A. Rappold & W Schempp; The
evolution of the azoospermia factor region AZFa in higher
primates R. Wimmer, S. Kirsch, G. A. Rappold & W.
Schempp; New insights into the evolution of chromosome
1 -A. Weise, H. Starke, K. Mrasek, U. Claussen &T. Liehr;
Panels of somatic cell hybrids specific for chimpanzee, go-
rilla, orangutan, and baboon R. Marzella, C. Carrozzo, P.
Chiarappa, V. Miolla & M. Rocchi; Comparative mapping
of human claudin-1 (CLDN1) in great apes I. Nanda, E
Kramer, B. H. E Weber, W. Schempp & M. Schmid; Evolu-
tionary breakpoints are co-localized with fragile sites and in-
trachromosomal telomeric sequences in primates A. Ruiz-
Herrera, E Garcia, E. Giulotto, C. Attolini, J. Egozcue, M.
Ponsa & M. Garcia. Part III: Primate Meiosis and Nuclear
Architecture. Chimpanzee chromosomes: Retrotranspos-
able compound repeat DNA organization (RCRO) and its
influence on meiotic prophase and crossing-over H. Hirai,
K. Matsubayashi, K. Kumazaki, A. Kato, N. Maeda &
H.-S. Kim; Inter- and intra-specific gene-density-correlated
radial chromosome territory arrangements are conserved in
Old World monkeys H. Tanabe, K. Kiipper, T. Ishida, M.
Neusser & H. Mizusawa; Fine structure and meiotic behav-
iour of the male multiple sex chromosomes in the genus
Alouatta- A. J. Solari & M. I. Rahn.


Volume 67(1) (September 2005) of the American Journal of
P -..'. was a special issue, dedicated to "Advances in
Field Behavioral Endocrinology," edited by Karen B. Strier
and Toni E. Ziegler. Contents: Introduction: Advances in
field-based studies of primate behavioral ecology K. B.
Strier & T. E. Ziegler, pp. 1-4; Effects of reproductive and
social variables on fecal glucocorticoid levels in a sample
of adult male ring-tailed lemurs (Lemur catta) at the Beza
Mahafaly Reserve, Madagascar L. Gould, T. E. Zeigler
& D. J. Wittwer, p.5-23; Social and reproductive factors
affecting cortisol levels in wild female golden lion tama-
rins (Leontopithecus rosalia) K. L. Bales, J. A. French, C.
M. Hostetler & J. M. Dietz, pp.25-35; Behavioral strate-
gies and hormonal profiles of dominant and subordinate
common marmoset (( .-' jacchus) females in wild
monogamous groups M. B. C. de Sousa, A. C. S. da
R. Albuquerque, F da S. Albuquerque, A. Araijo, M. E.
Yamamoto & M. de F Arruda, pp.37-50; Behavioral in-
dicators of ovarian phase in white-faced capuchins (Cebus
capucinus) S. D. Carnegie, L. M. Fedigan & T. E. Zei-
gler, pp.51-68; Variation in the resumption of cycling and
conception by fecal androgen and estradiol levels in female
northern muriquis (Brachyteles hypoxanthus) K. B. Strier
& T. E. Ziegler, pp.69-81; Coming of age: Steroid hor-
mones of wild immature baboons (Papio cynocephalus) L.
R. Gesquiere etal., pp.83-100; Female testosterone, domi-
nance rank, and aggression in an Ethiopian population of
hybrid baboons J. C. Beehner, J. E. Phillips-Conroy &
P. L. Whitten, pp.101-119; Radioimmunoassay of estrone
conjugates from urine dried on filter paper C. D. Knott,
pp.121-135; Reproductive endocrinology of wild female
chimpanzees (Pan troglodytes schweinfurthii): Methodo-
logical considerations and the role of hormones in sex and
conception M. E. Thompson, pp. 137-158; Fecal steroid
research in the field and laboratory: Improved methods for
storage, transport, processing, and analysis T. E. Ziegler
& D. J. Wittwer, pp.158-174.


Volume 67(4) (December 2005) of the American Journal
of .-. *'. ". dedicated to "Recent Advances in Color
Vision Research." It was edited by Hannah M. Buchanan-
Smith, and includes eight articles along with an introduc-
tion by the editor. Contents: Introduction: Recent ad-
vances in color vision research H. M. Buchanan-Smith,
pp.393-398; Comparative use of color vision for fru-
givory by sympatric species of platyrrhines K. E. Stoner,
P Riba-Hernandez & P W. Lucas, pp.399-409; Sugar
concentration of fruits and their detection via color in
the Central American spider monkey (Ateles .. rr., i P.
Riba-HernAndez, K. E. Stoner & P E. Lucas, pp.411-423;

Neotropical Primates 13(3), December 2005

Advantage of dichromats over trichromats in discrimina-
tion of color-camouflaged stimuli in nonhuman primates
- A. Saito, A. Mikami, S. Kawamura, Y. Ueno, C. Hira-
matsu, K. A. Widayati, B. Suryobroto, M. Teramoto, Y.
Mori, K. Nagano, K. Fujita, H. Kuroshima & T. Haseg-
awa, pp.425-436; Influence of stimuli size on color dis-
crimination in capuchin monkeys U. R. Gomes, D. M.
A. Pessoa, E. Suganuma, C. Tomaz &V. F. Pessoa, pp.437-
446; Color vision polymorphism in wild capuchins (Cebus
capucinus) and spider monkeys (Ateles ..-rr .., I in Costa
Rica C. Hiramatsu, T. Tsutsui, Y. Matsumoto, F. Aureli,
L. M. Fedigan & S. Kawamura, pp.447-461; Color vision
pigment frequencies in wild tamarins (Saguinus spp.) A.
K. Surridge, S. S. Suarez, H. M. Buchanan-Smith, A. C.
Smith & N. I. Mundy, pp.463-470; Demonstration of a
genotype-phenotype correlation in the polymorphic color
vision of a non-callitrichine New World monkey, capuch-
in (Cebus .i.i//.) A. Saito, S. Kawamura, A. Mikami, Y.
Ueno, C. Hiramatsu, K. Koida, K. Fujita, H. Kuroshima
& T. Hasegawa, pp.471-485; Color vision in marmosets
and tamarins: Behavioral evidence D. M. A. Pessoa, C.
Tomaz and V. E Pessoa, pp.487-495.


Escobedo-Morales, L. A. and Mandujano, S. 2005. As-
sessing local extinction risk in howler monkeys groups
(Alouattapalliata mexicana) in a highly fragmented land-
scape in Los Tuxtlas, Veracruz. ASP Bulletin 29(3): 8-9.
Nieves, M., Ascunce, M. S., Rahn, M. I. and Mudry, M.
D. 2005. Phylogenetic relationships among some Ateles
species: The use of chromosomic and molecular charac-
ters. Primates 46(3): 155-164.
Osorio, D., Smith, A. C., Vorobyev, M. and Buchanan-
Smith, H. M. 2004. Detection of fruit and the selection
of primate visual pigments for color vision. Am. Nat.
164(6): 696-708.
Partridge, J. 2005. The European studbook for black
howler monkeys an update. IZN- Int. Zoo News 52(5):
Ribeiro, S. and Bicca-Marques, J. C. 2005. Caracteristi-
cas da paisagem e sua relagao com a ocorrencia de bu-
gios-ruivos (Alouatta guariba clamitans Cabrera, 1940;
Primates, Atelidae) em fragments florestais no Vale do
Taquari, RS. Natureza e Conservafao 3(2): 65-78. [Eng-
lish version, pp.168-181.]
Snarr, K. A. 2005. Seismic activity response as observed
in mantled howlers (Alouatta palliata), Cuero y Salado
Wildlife Refuge, Honduras. Primates46(4): 281-285.
Villalobos, E, Valerio, A. A. and Retana, A. P. 2004. A phy-
logeny of howler monkeys (Cebidae: Alouatta) based on
mitochondrial, chromosomal and morphological data.
Rev. Biol. Trop. 52(3): 665-675.
Zaldivar, M. E., Rocha, 0., Glander, K. E., Aguilar, G.,
Huertas, A. S., Sanchez, R. and Wong, G. 2004. Distri-
bution, ecology, life history, genetic variation, and risk of

extinction of nonhuman primates from Costa Rica. Rev.
Biol. Trop. 52(3): 679-693.

Nagao, K., Takenaka, N., Hirai, M. and Kawamura, S.
2005. Coupling and decoupling of evolutionary mode
between X- and Y-chromosomal red-green opsin genes in
owl monkeys. Gene (Amsterdam) 352: 82-91.
Rotundo, M., Fernandez-Duque, E. and Dixson, A. E
2005. Infant development and parental care in free-
ranging Aotus azarai azarai in Argentina. Int. J. Primatol.
26(6): 1459-1473.
Santos, S. N. dos, Reis, J. W. L. dos, Silva-Filho, M. da,
Kremers, J. and Silveira, L. C. L. 2005. Horizontal cell
morphology in nocturnal and diurnal primates: A com-
parison between owl-monkey (Aotus) and capuchin
monkey (Cebus). VisualNeurosci. 22(4): 405-415.
Shmuel, A., Korman, M., Sterkin, A., Harel, M., Ullman,
S., Malach, R. and Grinvald, A. 2005. Retinotopic axis
specificity and selective clustering of feedback projections
from V2 to V1 in the owl monkey. J. Neurosci. 25(8):
Spirig, R., Peduzzi, E., Patarroyo, M. E., Pluschke, G. and
Daubenberger, C. A. 2005. Structural and functional
characterisation of the Toll like receptor 9 of Aotus nan-
cymaae, a non-human primate model for malaria vaccine
development. Immunogenetics 57(3-4): 283-288.

Ramos-Fernandez, G. 2005. Vocal communication in a
fission-fusion society: Do spider monkeys stay in touch
with close associates? Int. J. Primatol. 26(5): 1077-1092.
Riba-Hernandez, P., Stoner, K. E. and Lucas, P. W. 2005.
Sugar concentration of fruits and their detection via color
in the central American spider monkey (Ateles..-rr ,
Am. J. Primatol. 67(4): 411-423.
Russo, S. E. 2005. Linking seed fate to natural dispersal
patterns: Factors affecting predation and scatter-hoard-
ing of Virola ./lof/'1y//.I cc'd, in Peru.j. Trop. Ecol. 21(3):
Russo, S. E., Campbell, C. J., Dew, J. L., Stevenson, P. R.
and Suarez, S. A. 2005. A multi-forest comparison of di-
etary preferences and seed dispersal by Ateles spp. Int. J.
Primatol. 26(5): 1017-1037.
Schaffner, C. M. and Aureli, F. 2005. Embraces and groom-
ing in captive spider monkeys. Int. J. Primatol. 26(5):
Schmitt, D., Rose, M. D., Turnquist, J. E. and Lemelin,
P. 2005. Role of the prehensile tail during ateline loco-
motion: Experimental and osteological evidence. Am. J.
Phys. Anthropol. 126(4): 435-446.
Tardona, D. R. 2005. Influence of waterborne observers
on group spread among previously captive spider mon-
keys (Ateles ..Trr .. Animal Keepers' Forum 32(9):
Wallace, R. B. 2005. Seasonal variations in diet and forag-
ing behavior of Ateles chamek in a southern Amazonian
tropical forest. Int. J. Primatol. 26(5): 1053-1075.

Neotropical Primates 13(3), December 2005

Nascimento Borges, B. do and Harada, M. L. 2004. Diver-
gent evolution and purifying selection of the H (FUT1)
gene in New World monkeys (Primates, Platyrrhini). Ge-
netics Molec. Biol. 27(3): 342-349.
Strier, K. B. and Ziegler, T. E. 2005. Variation in the re-
sumption of cycling and conception by fecal androgen
and estradiol levels in female northern muriquis (Brach-
yteles hypoxanthus). Am. J. Primatol. 67(1): 69-81.
Talebi, M., Bastos, A. and Lee, P. C. 2005. Diet of southern
muriquis in continuous Brazilian Atlantic forest. Int. J.
Primatol. 26(5): 1175-1187.

O'Brien, J. K., Stojanov, T., Heffernan, S. J., Hollinshead,
F K., Vogelnest, L., Maxwell, W. M. C. and Evans, G.
2005. Flow cytometric sorting of non-human primate
sperm nuclei. Theriogenology 63(1): 246-259.
Pereira, M. M., Silva, J. J. P. da, Pinto, M. A., Silva, M. E
da, Machado, M. P., Lenzi, H. L. and Marchevsky, R. S.
2005. Experimental leptospirosis in marmoset monkeys
(C .- jacchus): a new model for studies of severe pul-
monary leptospirosis. Am. J. Trop. Med. Hygiene 72(1):
Philibert, B., Beitel, R. E., Nagarajan, S. S., Bonham, B.
H., Schreiner, C. E. and Cheung, S. W. 2005. Functional
organization and hemispheric comparison of primary au-
ditory cortex in the common marmoset (C .- jac-
chus). J. Comp. Neurol. 487(4): 391-406.
Pryce, C. R., Feldon, J., Fuchs, E., Knuesel, I., Oertle, T.,
Sengstag, C., Spengler, M., Weber, E., Weston, A. and
Jongen-Relo, A. 2005. Postnatal ontogeny of hippocam-
pal expression of the mineralocorticoid and glucocorti-
coid receptors in the common marmoset monkey. Euro-
peanJ. Neurosci. 21(6): 1521-1535.
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of dominant and subordinate common marmoset (Cal-
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Walker, S. E. 2005. Leaping behavior of Pithecia pithecia
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Stevenson, P. R., Pineda, M. and Samper, T. 2005. Influ-
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Notcher, R., Pilon, S., Singer, T. and Whiteley, A. 2004.
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Pantucek, R., Sedlacek, I., Petras, P., Koukalova, D.,
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Vokurkova, J., Ruzickova, V., Doskar, J., Swings, J. and
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Miller, C. T., Iguina, C. G. and Hauser, M. D. 2005.
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Serum concentrations of intact parathormone and

Neotropical Primates 13(3), December 2005

ionized calcium in vitamin D deficient emperor tama-
rins (Saguinus imperator): Response to environmental
modifications. In: Proceedings: American Association
of Zoo Veterinarians, American Association of 11 -.--
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sociation of Zoo Veterinarians, Place of publication not


Primate Society of Great Britain (PSGB) Winter Meet-
ing 2005, 9 December 2005. Flett Theatre, The Natural
History Museum, London. The theme is "Primate Evo-
lution and the Environment." Guest speakers include R.
D. Martin (The Field Museum, Chicago), Erik Seiffert
(Oxford University), Peter Andrews (The Natural History
Museum), Jussi Eronen and Mikael Fortelius (University
of Helsinki), Susan Ant6n (New York University), Sarah
Elton (University of Hull), Christophe Soligo (The Natural
History Museum), Jonathan Kingdon (Oxford University),
Urs Thalmann (University of Zirich) and Laurie Godfrey
(University of Massachusetts). Organised by: Christophe
Soligo, The Natural History Museum, e-mail: nhm.ac.uk>. See website: Winter2005.html>.

V Gbttinger Freilandtage "Primate Diversity Past,
Present and Future," 13-16 December 2005. University
of Gbttingen and German Primate Center, Gbttingen,
Germany. Organized by Peter M. Kappeler. Confirmed
invited speakers: Diversity in the past- Extinct primate
communities John F1, I,> (State University of New
York, Stony Brook). Diversity today. Diversity of Mala-
gasy primates Anne Yoder (Yale University); Diversity of
American primates Anthony B. Rylands (Conservation
International); Diversity of Asian primates Jatna Supri-
atna (Conservation International Indonesia); Diversity of
African primates John F Oates (Hunter College New
York); Primate biogeography- Shawn Lehman (University
of Toronto); Speciation and taxonomy Colin P. Groves
(Australian National University); Human diversity Mark
Stoneking (Max Planck Institute, Leipzig). Preserving Di-
versity for Tomorrow: Diversity and conservation hotspots
- Russell A. Mittermeier (Conservation International); Ex-
tinction biology Carlos Peres (University of East Anglia);
Conservation genetics George Amato (Wildlife Conser-
vation Society); Conservation genetics Michael Bruford
(Cardiff University); Reintroductions Carel P. van Schaik
(University of Zirich). Comparative Perspectives: Specia-
tion in birds Trevor Price (University of Chicago); Bird

taxonomy and conservation Robert Zink (University of
Minnesota). Contact: Prof. Dr. Peter M. Kappeler, Deut-
sches Primatenzentrum (DPZ), Kellnerweg 4, D-37077
Gbttingen, Tel/Fax: +49-551-3851-284/291, e-mail:
, website: sociobiology/GFT2005/index.htm>.


Ecology in an Era of Globalization: Challenges and Op-
portunities for Environmental Scientists in the Ameri-
cas, 8-12 January 2006, Merida, Mexico. This conference
will be held at the Fiesta Americana Hotel in Merida and
is co-hosted by the Universidad Aut6noma de Yucatan and
the Centro de Investigaciones Cientificas de Yucatan. Ab-
stracts should address one of the meeting's three subthemes:
invasive species, human migration, and production. The
invasive species subtheme includes such topics as disper-
sal of invasive plant and animal species, emerging diseases,
and resistance of local ecosystems to invasive species and
disease. The human migration subtheme includes the envi-
ronmental effects of international and local emigration and
immigration on recipient and source areas. Potential topics
include infrastructure development needs and impacts, ef-
fects on land cover, and land-use impacts. The production
subtheme focuses on ecosystem transformations, including
land-use change required to produce goods and services for
human use. Potential topics include the effects of changes
in forest and agricultural policy on economies, biodiversity,
and ecosystems throughout the Americas, in terrestrial,
marine, and freshwater systems. We particularly welcome
reports of projects that are interdisciplinary and that con-
sider the need to communicate with broad audiences. For
more information or to submit an abstract, visit www.esa.org/mexico>. Deadline for abstract submissions:
16 September 2005.

75' Annual Meeting of the American Association for
Physical Anthropology, 5-12 March 2006, Anchorage,
Alaska, USA. For program information, please contact
the Program Chair, Lyle W. Konigsberg, Department of
Anthropology, University of Tennessee, Knoxville, TN
37996-0720, USA, Tel: (865) 974-4408, fax: (865) 974-
2686, e-mail . Local Arrangements
Committee Chair: Christine Hanson, Department of An-
thropology, University of Alaska Anchorage, Anchorage,
AK 99508, USA, tel: 907-786-6839, fax: 907-786-6850,
e-mail . Website at physanth.org/annmeet>.

Primate Society of Great Britain (PSGB) Spring Meet-
ing 2006, 27-28 March 2006, University of Stirling, Stir-
ling, Scotland. The theme is "Primate Mentality and Well-
being." On the afternoon of 27 March invited speakers will
address the relationship between cognition and welfare in
primates. Other topics are welcomed for posters and oral
sessions. There will be a prize for the best postgraduate
presentation and poster. A provisional programme and

Neotropical Primates 13(3), December 2005

instructions for presenters can be found on the meeting
web site at: PSGB2006.php>. For more information please contact: Dr
Sarah Vick (PSGB), Psychology Department, University of
Stirling, FK9 4LA, Scotland. E-mail address for enquiries:

21st Congress of the International Primatological Soci-
ety, 25-30 June 2006, Imperial Resort Beach Hotel, En-
tebbe, Uganda. Theme: "Primate Conservation in Action."
Preliminary contact details: Dr. William Olupot, Chair,
Organizing Committee, IPS 2006 Congress, P. 0. Box
21669, Kampala, Uganda, Tel: 077598134, 077947397,
041501020, e-mail .

29th Annual Meeting of the American Society of Prima-
tologists (ASP), 16-19 August 2006, San Antonio, Texas.
Sponsored by Southwest National Primate Research Center.
Tentative deadline dates are 5 December 2005 to notify
program chair of intent to offer a symposium or workshop;
9 January 2006 to send symposia and workshop abstracts
with confirmed list of participants to program chair; and
6 February 2006 for all final abstracts for symposia, oral,
and poster presenters. See the ASP website for updates and
further information: html>.

1t European Congress of Conservation Biology, 22-26
August 2006, Eger, Hungary. The European Section of the
Society for Conservation Biology is determined to promote
the development and use of science for the conservation of
European species and ecosystems, and to make sure that
conservation policy is firmly underpinned by the best avail-
able scientific evidence. This keystone congress will bring
together a wide array of academics, policymakers, students,
NGO representatives, and biodiversity managers from
throughout Europe and beyond. For more information,
see the Congress website at or
contact Andras Bildi, Chair of the Local Organising Com-
mittee, at .

VII Congreso Internacional sobre Manejo de Fauna
Silvestre en la Amazonia y Am6rica Latina, del 3 al 7
de septiembre de 2006, Ilhdus, Bahia, Brasil. El VII Con-
greso Internacional sobre Manejo de Fauna Silvestre en la
Amazonfa y Am&rica Latina enfocara su atenci6n en los
studios y programs de manejo que actualmente estin
siendo ejecutados en la Amazonfa y en Latinoam&rica,
con el prop6sito de evaluar los resultados alcanzados y las
limitaciones encontradas en la conducci6n de los mismos.
Una de sus principles metas sera expandir el enfoque del
event a los mas amplios aspects del manejo de fauna en
toda Latinoam&rica. El VII Congreso Internacional sobre
Manejo de Fauna Silvestre en la Amazonfa y Amdrica
Latina incluira conferencias magistrales, mesas redondas,
secciones tematicas con presentaciones orales libres, exposi-
ciones en posters, simposios, workshops, cursos durante
y posteriores al congress, y excursions pos-congreso. Las

areas tematicas que se abordaran en este event seran: con-
servaci6n in situ y Areas naturales protegidas, conservaci6n
ex situ de fauna silvestre, preservaci6n y recuperaci6n de
habitats, metodologfas aplicadas para el manejo de fauna
silvestre con comunidades, criterios para el uso sustentable
de fauna silvestre, indicadores de sustentabilidad, etologfa
aplicada al manej o, medicine veterinaria de la conservaci6n,
fisiologfa y ecologfa, producci6n en criaderos, comercio,
polftica y legislaci6n de fauna silvestre. Estamos recibiendo
propuestas para mini-cursos, workshops y simposios hasta
el 31/12/05. Apreciaremos el apoyo de diversas instituci-
ones. Existen posibilidades de instalaci6n de stands institu-
cionales para difusi6n y ventas. El plazo para el envfo de
resdmenes es hasta el 30/04/06. Para mayor informaci6n:

Primate Society of Great Britain (PSGB) Winter Meet-
ing 2006 5 December 2006, Cambridge, UK. The
focus of the winter meeting will be on Primate Conserva-
tion Genetics; it will take place in the West Road Concert
Hall at the University of Cambridge from 9:00 19:00
on 5 December 2006. This will be a unique opportunity
to hear about current exciting research on molecular ge-
netic studies of great apes, Old and New World monkeys
and strepsirrhines. Information about the meeting, includ-
ing titles and abstracts of presentations, can be found at
. Registration at the door (PSGB
student members: 10; student non-members: 15; PSGB
members: 15; non-members: 25). The meeting is organ-
ised by Prof. Mike Bruford ()
and Dr. Leslie Knapp (), who can be
contacted for further information.


6th Zoos & Aquariums Committing to Conservation
Conference, 26-31 January 2007, Houston, Texas. ZACC
is a bi-annual event that promotes the role of zoos and
aquariums in supporting conservation activities world-
wide, both at their institutions and in the field. Confer-
ence participants include representatives from zoological
institutions, international conservation organizations, local
non-governmental organizations, government agencies,
funding agencies and, most importantly, field biologists
and conservationists. Presentations at the 2007 ZACC will
highlight both ongoing projects and new initiatives that
offer opportunities for institutional support. There will be
a major focus on field-based initiatives that have already es-
tablished links to zoos and aquariums, as well as promising
candidates for such partnerships. In addition, the program
will feature presentations related to the organization, man-
agement, and support of zoo-based and aquarium-based
conservation programs. The full conference registration fee
($195) will include icebreaker event, all sessions, breaks,
lunches, conference proceedings, zoo day transport, zoo
day lunch and dinner. All funds raised above conference
costs will be allocated to the conservation fund for this
conference. The deadline for submitting paper and poster

Neotropical Primates 13(3), December 2005

abstracts is 1 September 2006. Abstracts submitted elec-
tronically should be addressed to org> and to . Abstracts submit-
ted as hard copy should be addressed to: 2007 ZACC Con-
ference, Attn: Bill Konstant, Director of Conservation and
Science, Houston Zoo, 1513 North MacGregor, Houston,
Texas 77030, USA. For more information, see the confer-
ence website at .

XII Congresso Brasileiro de Primatologia, 22 a 27 de
julho de 2007, Belo Horizonte, Minas Gerais, Brasil. 0
local escolhido d a PUC-BH, de excelente infra-estrutura
e localizada no bairro Coraqao Eucarfstico, em Belo Hori-
zonte (MG), o que facilitara o acesso de quem vemn de
fora. 0 tema escolhido pela diretoria foi "Prioridades de
pesquisa para o estudo de primatas neotropicais" e, por-
tanto, solicitamos a colaboraqao e participacao de todos os
interessados na construcao da programaqao deste event
tao important para todos n6s. Estaremos recebendo pro-
postas de mesas-redondas, palestras, mini-cursos e events
paralelos, tudo voltado ao tema citado acima, ate dia 30 de
abril pr6ximo, onde colocaremos a disposicao no site da
SBPr ou na lista de discusses dos primat6logos ( br.groups.yahoo.com/group/primatologia>) para votaq6es
finals e conclusao de nossos trabalhos ainda neste semestre.
Informao6es adicionais: ologia/> ou entrar em contato com Prof. Dr. Fabiano R. de
Melo, Presidente da Sociedade Brasileira de Primatologia,
Coordenador do curso de Ciencias Biol6gicas, UEMG/
FAFILE, campus de Carangola, Praca dos Estudantes, 23,
Santa Emflia, Carangola 36800-000, Minas Gerais, Brasil,
(32) 3741-1969 / (32) 8845-2904.


The journal/newsletter aims to provide a basis for conservation
information relating to the primates of the Neotropics. We
welcome texts on any aspect of primate conservation, including
articles, thesis abstracts, news items, recent events, recent publica-
tions, primatological society information and suchlike.


Please send all English and Portuguese contributions to:
John M. Aguiar, Conservation International, Center for Applied
Biodiversity Science, 2011 Crystal Drive, Suite 500, Arlington,
VA 22202, Tel: 703 341-2400, Fax: 703 979-0953, e-mail:
, and all Spanish contributions to:
Ernesto Rodrfguez-Luna, Instituto de Neuroetologfa, Universi-
dad Veracruzana, Apartado Postal 566, Xalapa 91000, Veracruz,
Mexico, Tel: 281 8-77-30, Fax: 281 8-77-30, 8-63-52, e-mail:


Manuscripts may 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 (English,
Portuguese) or (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 should
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), Biogeogra-
phy, 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 English, 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 descriptions, for example). Full articles
will be sent out for peer-review.

Short articles. These are usually reviewed only by the editors.
A broader range of topics is 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 conservation
and research programs (who, what, where, when, why, etc.) and
most particularly information on geographical distributions,
locality records, and protected areas and the primates which
occur in them. Texts should not exceed 10 pages in length
(1.5 spaced, including the references).

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

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

References. Examples of house style may be found throughout this
journal. Please refer to these examples when listing references:

Journal article
Stallings, J. D. and Mittermeier, R. A. 1983. The black-tailed
marmoset (C( .- argentata melanura) recorded from Paraguay.
Am. J. Primatol. 4: 159-163.

Chapter in book
Brockelman, W. Y. and All, R. 1987. Methods 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.

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

Wallace, R. B. 1998. The behavioral ecology of black spider
monkeys in north-eastern Bolivia. Doctoral thesis, University of
Liverpool, Liverpool, UK.

Muckenhirn, N. A., Mortensen, B. K., Vessey, S., Fraser,
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 Applied
Biodiversity Science, 2011 Crystal Drive, Suite 500,
Arlington, Virginia 22202, USA.

r mg~irt~~

Neotropical Primates
A Journal and Newsletter of the IUCN/SSC Primate Specialist Group
Vol. 13(3), December 2005


Short Articles

Population Survey of the Azuero Howler Monkey (Alouatta palliata trabeata) in Herrera Province,
Republic of Panama
Pedro G M ndez-C arvajal ........................ .........................................................................................................................1.......
Estudos Preliminares da Presenca de Sagiiis no Municipio de Bauru, Sao Paulo, Brasil
Hugo Medeiros Garrido de Paula, Renata Souza Tdvora, Marcos Vinicius de Almeida, Larissa Sbeghen Pelegrini,
Graziela ValenCa da Silva, Rosdngela Lopes Zaganini e Anderson Lucindo......................................................................................6...
Boa constrictor Predation on a Titi monkey, Callicebus discolor
Diego E Cisneros-Heredia, AndrAs Ledn-Reyes and Sylvia Seger..................................................................... ....................11
Uma Avaliaqao da Dieta, da Area de Vida e das Estimativas Populacionais de Cebus nigritus (Goldfuss, 1809)
em um Fragmento Florestal no Norte do Estado do Parana
Gabriela Ludwig, Lucas M. Aguiar e Vlamirj Rocha.......................................................................................12
Coprophagy in Captive Brown Capuchin Monkeys (Cebus apella)
H elissandra M attjie Prates and Julio C sar Bicca-M arques............................................................. ................. ....................... 18
Rehabilitaci6n y Reproducci6n de Alouatta caraya Fuera de su Area de Distribuci6n Natural
Gabriela Bruno, Aldo M. Giudice, Mariela Nieves y Marta D. Mudry..........................................................................................21
Male Tenure and Reproductive Success in Single-male vs. Multi-male Groups of Free-Ranging Howling Monkeys
in Costa Rica
M argaret R. Clarke and K enneth E. G lander ............................................... ............. ........... ............................................................ 23
Caracterizaci6n de los Dormideros Usados por Ateles belzebuth en el Parque Nacional Yasuni, Ecuador
W ilm er E. Pozo R. ......................................... ........................................................................ 27

N ew s ........ ..................................................................... ....................................................................................................... 34

R recent Publications........................................ ............................ .........................................51

M eetin g s ....................................................................... ....................................................................................................... 5 8

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