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Title: Neotropical primates
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Permanent Link: http://ufdc.ufl.edu/UF00098814/00047
 Material Information
Title: Neotropical primates a newsletter of the Neotropical Section of the IUCNSSC Primate Specialist Group
Abbreviated Title: Neotrop. primates
Physical Description: v. : ill. ; 27 cm.
Language: English
Creator: IUCN/SSC Primate Specialist Group -- Neotropical Section
IUCN/SSC Primate Specialist Group -- Neotropical Section
Conservation International
Center for Applied Biodiversity Science
Publisher: Conservation International
Place of Publication: Belo Horizonte Minas Gerais Brazil
Belo Horizonte Minas Gerais Brazil
Publication Date: December 2004
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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Bibliographic ID: UF00098814
Volume ID: VID00047
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: oclc - 28561619
lccn - 96648813
issn - 1413-4705


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    Back Matter
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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
1919 M St. NW, Suite 600, Washington, DC 20036, USA

ISSN 1413-4703 Abbreviation: Neotrop. Primates
DOI: 10.1896/ci.cabs.2004.np.12.3

Anthony B. Rylands, Center for Applied Biodiversity Science, Conservation International, i 1...... DC
Ernesto .iI._. -T ... UniversidadVeracruzana, Xalapa, Mexico

Assistant Editors
John M. Aguiar, Center for Applied Biodiversity Science, Conservation International, i- ...... DC
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 Germany
Russell A. Mittermeier, Conservation International, i- ...... DC
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 -T .
Vice Chair for Asia Ardith A. Eudey
Vice Chair for Africa Thomas M. Butynski
Vice Chair for Madagascar Jorg U. Ganzhorn

Design and Layout: Kim Meek, Center for Applied Biodiversity Science,
Conservation International, -i ......1 DC

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

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

Front Cover: Callithrixflaviceps from Minas Gerais, Brazil. 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, the Houston Zoological Gardens Conservation Program, General Manager F.I .... 1513 North MacGregor, Houston, Texas 77030, USA,
and the Los Angeles Zoo, Director John R. Lewis, 5333 Zoo Drive, Los Angeles, California 90027, USA.



PL aWd nRuK OwtDenbL

Neotropical Primates 12(3), December 2004


Luciane L. de Souza
Helder L. Queiroz
Jos Mdrcio Ayrest

The mottled-face tamarin, Saguinus inustus(Schwarz, 1951),
has never been studied in the wild, and its distribution,
ecology and behavior are virtually unknown. It occurs
in southeastern Colombia, between the Rfos Caqueti,
Yarf, and Guayabero/Guaviare, and extends into adjacent
territory in Brazil between the Rios JapurA and Negro
(Hernandez-Camacho and Cooper, 1976; Hershkovitz,
1977; Hernandez-Camacho and Defler, 1991; Defler, 2004).
How far its range extends east between the Rios JapurA and
Negro is not known. A. B. Rylands (unpubl., reported in
Barnett et al., 2002) observed a group on the north bank
of the Lago Amana, north of the mouth of the Rio JapurA
in 1980. Surveys by Rylands (1992) and Neri and Borges
(1998) resulted in inconclusive reports of the species along
the lower Rio Jad and the Rio Uninf respectively, both
in the Jad National Park (Barnett et al., 2002). Iwanaga
(2004) was the first to confirm its presence in the park, on
the upper Rio Jad in the west at the site called Monteiro
(0236'22"S, 6321'27"W). Iwanaga (2004) also noted
that they were reported to be common by local people
on the upper Rio Uninf. Here we record the presence of
S. inustus in the Amand Sustainable Development Reserve
in the southwestern part of the Rios Negro and JapurA
interfluvial region.

The Amand Sustainable Development Reserve (SDR) of
2,350,000 hais part of one of the largest blocks ofcontinuous
protected forests in the world, linking the MamirauA SDR

of 1,124,000 ha to its southwest and the Jad National Park
of 2,378,410 ha to the east, in the north-central part of
the state of Amazonas (Fig. 1). The vegetation there is
predominantly tall terra firma forest interspersed with
flooded forests (vdrzea and igapd) and some small areas of
white sand forest or campinarana (Ayres et al., 1997).

In June of 2001, we were participating in the first
expedition of a faunal survey in the vicinity of the head
of the Lago Amana, near the village of Comunidade
Boa Esperanca (CBE) in the municipality of Maraa
(02o28'12"S, 6444 2- '".) (Figure 1). There we found
a dead adult male mottled-face tamarin, S. inustus, killed
by local people from the Comunidade (Fig. 2). It was wild
(not a pet), one of a group occupying secondary forest scrub
behind the village. The skin is preserved in the reference
collection of the Instituto de Desenvolvimento Sustentavel
MamirauA (IDSM), Tefd. The skin of its face was almost
completely unpigmented and the hair melanic, as described
by Hershkovitz (1977) features considered typical for this
species (Fig. 3). The fur of the rest of the body was entirely
black, with the exception of the mantle, which was dark
brown. It weighed approximately 500 g and had a total
length (head to tail tip) of 700 mm.

During our surveys, we saw groups of S. inustus foraging in
secondary (around abandoned cultivated plots) and primary
forest in the vicinity of the CBE, and also in the terra firma
forests by the JuA Grande Igarapd (creek) (L. Souza, pers.
obs.). Mottled-face tamarins are not hunted due to their
small size, but they are occasionally captured by the local
people to be raised as pets (Fig. 4). Other primates known
in this area and recorded during the survey were the squirrel
monkey (Saimiri sciureus), capuchin monkeys (Cebus ,J''I//.
and Cebus albiJfons), the collared titi ((. .... torquatus),
the red howler monkey (Alouatta seniculus) and the black-
headed uakari (GCcajao melanocephalus). The Amand SDR
is within the geographic range of the noisy night monkey,
Aotus vociferans, as indicated by Hershkovitz (1983), but
none were seen by our survey team. Three other primates

Figure 1. The Amana State Sustainable Development Reserve in the state of Amazonas (AM), central Amazon.

Neotropical Primates 12(3), December 2004

Figure 2. Adult male of Saguinus inustus killed by local inhabitants
of the Amana State Sustainable Development Reserve, Amazonas.

Figure 3. The mottled face of Saguinus inustus.

Figure 4. A juvenile pet S. inustus.

that may occur in the area but have not
there to date are members of the genera
and Lagothrix.

been recorded
Pithecia, Ateles

This note records the presence of groups of S. inustus in the
Amana Sustainable Development Reserve on the lower Rio
Japuri, more than 300 km west of the locality recorded by
Iwanaga (2004) in the Jad National Park. Additional field
surveys are needed to determine the northern limits of its
range in Brazil and the more exact delimitation of its range
further east towards the lower Rio Negro.

Acknowledgments: It was the late Jose Mircio Ayres who
made this study of the fauna and flora of the Amana
State Sustainable Development Reserve possible. We also
thank the inhabitants of the Comunidade Boa Esperanga,
who provided information regarding this species and
others in the reserve, and the Mamiraud Institute for
Sustainable Development (IDSM-OS/MCT) for financial
and logistical support of the field expeditions, and especially
to Josivaldo F. Modesto, coordinator of operations of

Luciane L. de Souza, Postgraduate Program in Zoology,
Universidade Federal do Para/ Museu Paraense Emflio
Goeldi, Caixa Postal 399, Belem 66040-170, Pard, Brazil,
e-mail: , Helder L. de Queiroz,
Director Cientifico, Instituto de Desenvolvimento Sustentivel
Mamiraud, Avenida Brasil 195, Jurui, Caixa Postal 38, Tefe
69470-000, Amazonas, Brazil, e-mail: org.br>, and Jose Mircio Ayres, erstwhile researcher at the
Museu Paraense Emiflio Goeldi, Belem, founder of the
Institute de Desenvolvimento Sustentivel Mamiraud, Tefe
(deceased March, 2003).


Ayres, J. M., Silva, V. F. and Nelson, B. 1997. Proposta de
Criaqdo Reserva de Desenvolvimento Sustentdvel Amand.
Report. Institute de Protegao Ambiental do Estado do
Amazonas (IPAAM), Manaus.
Barnett, A. A., Borges, S. H., de Castilho, C. V., Neri, E
M. and Shapley, R. L. 2002. Primates of the Jad National
Park, Amazonas, Brazil. Neotrop. Primates 10(2): 65-70.
Defler, T. R. 2004. Primates of Colombia. Conservation
International Tropical Field Guide Series, Conservation
International, Bogoti.
Hern~ndez-Camacho, J. and Cooper, R. W. 1976. The non-
human primates of Colombia. In: Neotropical Primates:
Field Studies and Conservation, R. W. Thorington Jr. and
P. G. Heltne (eds.), pp.35-69. National Academy of
Sciences, Washington, DC.
Hernindez-Camacho, J. and Defler, T. R. 1991. Algunos
aspects de la conservaci6n de primates no-humanos en
Colombia. In: La Primatologia en Latinoamerica, C. J.
Saavedra, R. A. Mittermeier and I. B. Santos (eds.), pp.67-
100. World Wildlife Fund, Washington, DC.
Hershkovitz, P. 1977. Living New World Monkeys
(I'P/.! ,,, With an Introduction to Primates. Vol. I. The
University of Chicago Press, Chicago.
Hershkovitz, P. 1983. Two new species of night monkeys,
genus Aotus (Cebidae, Platyrrhini): A preliminary report
on Aotus taxonomy. Am. J. Primatol. 4: 209-243.
Iwanaga, S. 2004. Levantamento de mamfferos diurnos de
medio e grande porte no Parque Nacional do Jad: Resul-
tados preliminares. In: Janelas para a Biodiversidade no
Parque Nacional do Jau Uma Estrategia para o Estudo da
Biodiversidade na Amazdnia, S. H. Borges, S. Iwanaga,
C. C. Durigan and M. R. Pinheiro (eds.), pp.195-207.
Fundagao Vit6ria Amaz6nica, Manaus.

Neotropical Primates 12(3), December 2004


Erwin Palacios
Adriana Rodriguez
Claudia Castillo


Saguinus is the most diverse of the Neotropical primate
genera, with 13-15 species and 33 recognized forms
(Hershkovitz, 1977; Rylands et al., 2000). There have been
detailed studies of the feeding ecology, social organization,
and behavioral ecology of the majority of the extant
Saguinus species. S. leucopus is notable for the deficiency
of information about it (Snowdon and Soini, 1988; Calle,
1992; Vargas and Solano, 1996; Poveda, 2000, and Cuartas-
Calle, 2001), and perhaps the least known is S. inustus. The
latter occurs in southeastern Colombia west of the Andes,
between the Rio Mesay and the frontier with Brazil, and
between the Rios Guayabero-Guaviare and CaquetA. There
is still no accurate delimitation of the eastern and western
boundaries of its geographical range in Colombia (Defler,
2003). S. inustus also occurs in western Brazil, between the
Rios Negro and JapurA and the Colombian border. Here

we present preliminary data on some aspects of the ecology
of this species on the lower Rio Caqueta, Colombian
Amazonia and briefly discuss the importance of conducting
further research on its ecology in the region.

Subjects, Study Site and Data Collection

Mottled-face tamarins are small: head-body = 208-259
mm (n = 10) with a tail slightly longer, between 330 and
410 mm (n = 10) (Hershkovitz, 1977). On the lower Rio
Caqueta they are known as "hueviblanco" because the
males have naked external genitalia and a white scrotal sac.
Otherwise black, they have white patches of skin on each
side of the muzzle, which makes them easily recognizable
from a distance.

The study was carried out in the interfluvial forests adjacent
to the lower Rios Caqueta and Apaporis, Colombian
Amazonia. Observations were made in the vicinity of
Comeyafi (117'S, 6934'W), a 19,000-ha indigenous
reserve on the left margin of the Rio Caqueta in the state
of Amazonas (Fig. 1). We first met with the community
in order to tell them the purpose of conducting the study
and to select people to participate in the fieldwork. With
the help of field assistants we found a group of five animals
frequently seen near the community. Existing trails in
the area were used to search for and follow them, and

Table 1. Fruits eaten by a group of S. inustus on the lower Rio Caqueti, Colombian Amazonia.
Species Family Yucuna name Part eaten
Mendoncia ovata Acanthaceae Pijiture camure Pulp
Tapirira guianensis Anacardiaceae Uayapala or ingna uala Pulp
Rollinia mucosa Annonaceae Cahayi Pulp
Couma macrocarpa Apocynaceae Yuuchi Pulp
Lacmellea cf. arborescens Apocynaceae Gemacacu Pulp
Pourouma cecropiifolia Cecropiaceae Caami Pulp
Pourouma tomentosa Cecropiaceae Cahamuld or Maprimutula Pulp
Buchenavia cf. viridiflora' Combretaceae Cumela Pulp
Inga edulis Leguminosae Guiro o Yucurupi Aril
Inga leptocarpa Leguminosae Guiro "de rastrojo" Aril
Inga pilosula Leguminosae Guiro Aril
Inga thibaudiana Leguminosae Guiro de rastrojo Aril
Inga yasuniana Leguminosae Guiro Aril
Inga sp. 1 Leguminosae Guiro Aril
Inga sp.2 Leguminosae Guiro Aril
Inga sp.3 Leguminosae Guiro Aril
Mouriri cf. acutiflora Melastomataceae Yukurt or Yauhimapula Pericarp and pulp
Abuta grandifolia Menispermaceae Pulp
Pseudolmedia laevis Moraceae Amasi Pulp
Pouteria guianensis Sapotaceae Imad Pulp
Pouteria sp. 1 Sapotaceae Uiyunumala Pulp
Pouteria sp.2 Sapotaceae Jarapila Pulp
Unknown Quiinaceae? Maya pijulare Aril
A group of 11 animals were seen eating this fruit near the Lomalinda Indigenous Community (12 km west towards Comeyafu).

Neotropical Primates 12(3), December 2004

additional trails were cut to cover the group's known range.
We observed the group from March to June 2003, which
included the end of the dry season (early rainy season) and
the rainy season. For a period of 18 days we attempted to
follow the group for the entire day, but this was achieved
on only five days because of the lack of a more extensive
trail grid. Daily ranges were drawn on a map scale 1:1000
and were measured using the distances between consecutive
group positions recorded during the day. Home range size
was calculated using the convex polygon method, which
although possibly overestimating the range (Albernaz,
1997), was the most appropriate because the quadrate
method requires a comprehensive trail grid. Trees used as
feeding sources were marked with colored flagging tape and
later revisited to obtain botanic specimens for identification.
Information on feeding by S. inustus was limited to the
animal and plant species we saw them eat.

Results and Discussion

Group size, use of space and daily ranges
Group size varied from three to six (mean 4.4, n = 5, sd
= 1.14) at our study site, but larger groups were observed
elsewhere. A group of 11 and another of nine were seen
22 km to the west (E. Palacios, pers. obs.). Including these
two groups we have a mean group size of 6.0 (n = 7; sd =
2.7). Defler (2003) reported group sizes of three, seven, and
eight individuals based on sightings in the same area. These
figures are similar to those of other Saguinus species (Freese,
1975; Soini, 1987; Sussman and Kinzey, 1984; Janson and
Terborgh, 1985; Kostrub, 1997; Peres, 2000). We never
observed temporary associations between groups.

Our study group used an area of 35 ha, which included
terra firma forest and flooded forest. The former included
areas of primary and secondary forest (locally called
rastrojo), and clearings abandoned after being farmed.
Rastrojo alto was the local name for high secondary forest,
and rastrojo bajo for low secondary forest. Flooded forest
includes areas that suffer occasional flash floods (of one
to a few days from overflowing creeks) as well as vdrzea
(seasonally flooded for three to five months). A little more
than half (54%) of the group's range was secondary forest,
and of that mostly (93%) rastrojo bajo. Primary forest took
up 34% of the home range, but we believe that this forest
type would have come to comprise a larger portion of
the group's home range if we had observed the group for
longer. Only about 1% of the range was flooded forest.
Approximately 12% of the area included in the forest
matrix used by the mottled-face tamarins was occupied by
the Indian's cultivation plots.

Mean daily range length was 961 m (range 750-1100 m;
sd = 137; n = 5 complete days). Although the home range
size is similar to S. .... S. fuscicollis, S. imperator
and S. labiatus (Izawa, 1978; Terborgh, 1983; Kessler,
1995; Veracini, 2000), it is notably smaller than the ranges
for S. mystax and S. fuscicollis reported by Peres (2000).
This undoubtedly is a reflection of the short period of

study S. inustus range would be larger if recorded over an
entire year.

The mottled-face tamarins were seen to eat the fruits of
23 plant species from 12 families and 13 genera (Table 1).
These plant species were spread through terra firma (primary
and secondary forest) and flooded forest, and all, except for
the liana Mendoncia ovata, were trees.

Some of these fruits were typically found in the secondary
forest. For instance, Mendoncia ovata was commonly
seen growing in the low and high rastrojo, as was Inga
thibaudiana. Other species, such as the two Pourouma spp.,
grow near large forest gaps and along the borders between
the primary and secondary forest and the cultivated plots.
Buchenavia sp. and two of the Inga species, on the other
hand, were found only in the flooded forest.

As reported for other tamarins, S. inustus was also seen
to eat small spiders, orthopterans, and ant larvae. These
resources were commonly obtained as the tamarins foraged
in the middle and lower levels of the forest, especially in the
low rastrojo, where they were often seen moving about only
0.5 m above the forest floor.

Inter-specific associations
Twice we saw mottled-face tamarins interacting with
groups of ( .... torquatus. They were observed feeding
together in a Pourouma cecropiifolia tree; titis and tamarins
shared different levels of the tree crown, and no agonistic
behaviors were seen. No encounters with other primates
were recorded; continued hunting in the forests around
Comeyafi and the neighboring community has extirpated
the larger to middle-sized primate species such as woolly
monkeys, Lagothrix lagothricha, and the tufted capuchin,
Cebus aj/,i/ (E. Palacios, unpubl. data). Although squirrel
monkeys, Saimiri sciureus, and red howlers, Alouatta
seniculus, are still present in the area, we never observed
them during our time there.

Conservation Aspects

S. inustus is ranked as of Least Concern under the IUCN
categorization (2001). It seems to be common around
Indian villages along the lower Rios Caqueta and Apaporis
interfluvium. In general, one sees more tamarins in habitats
that have been disturbed by human activities. Peres (1999),
for example, found that densities of S. fuscicollis, S. mystax,
and S. imperator were higher in areas subject to moderate
to heavy hunting pressure than in those where hunting was
minimal or non-existent. Density comparisons of the same
set of species, and including S. ..rr .., in protected and
unprotected areas have shown that these species are more
abundant and comprise a greater proportion of the total
primate density in the latter (Freese et al., 1977; Soini,
1987). Nevertheless, some unprotected sites show densities
as low or lower than those in protected sites (Muckenhirn
etal., 1975; Freese et al., 1982). Plots cultivated by Indians,

Neotropical Primates 12(3), December 2004

peasant crops, and small-scale logging have transformed
many areas of the mottled-face tamarin's natural habitat.
Slash-and-burn is the prevalent agricultural practice in
the region, but can be considered of low impact in forest
conversion only when human population densities are
very low. Rural populations are growing considerably,
concentrating their numbers in certain areas and making
increasing demands on forest resources. The interfluvium
between the Rfos Caqueta and Apaporis, east to the
mouth of the Rio Miriti, is a case in point, where a very
large proportion of the region's indigenous population is
concentrated (1.24 people/km2) and where numbers will
continue to increase through immigration.

Much (60-70%) of the range of S. inustusin Colombia falls
within the boundaries of indigenous reserves (Defler, 2003),
along with smaller areas on the lower Rio Caqueta settled by
peasant families. It will be important to continue studying
these tamarins and monitoring their densities along with
current trends of forest clearing and other human activities,
so that we can attain a better understanding of the ecology
and conservation status of the species. These actions will, we
hope, both continue our learning about the ecology of this
interesting primate, and provide a mechanism to involve
local people in joint conservation measures in their lands.

Acknowledgments: Conservation International provided
funding through the Primate Action Fund, and we thank
particularly RussellA. Mittermeier and William R. Konstant
for their support. We also thank Rodolfo Yucuna, chief of
the Comeyafi Yucuna Indigenous Community, and field
assistants from the community who helped us during the
study. Dairon Cardenas from Instituto Sinchi provided
valuable help with the identification of plants.

Erwin Palacios, Adriana Rodriguez and Claudia
Castillo, Conservation International Colombia, Bogota
DC, Colombia, e-mails: ,
; .


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Bogota, DC.
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censo preliminary y recomendaciones para el manejo de
una poblaci6n natural de Saguinus leucopus en la zona
de influencia del Proyecto Hidroeldctrico La Miel II.
Unpublished manuscript.
Cuartas-Calle, C. A. 2001. Partial distribution of tamarins
(Saguinus leucopus, Callitrichidae) in Departamento de
Antioquia, Colombia. Neotrop. Primates 9: 107-111.
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of the Callitrichidae, D. G. Kleiman (ed.), pp.121-130.
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Freese, C. H., Heltne, P. G., Castro, N. and Whitesides,
G. 1982. Patterns and determinants of monkey densities
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Primatol. 3: 53-59.
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19: 241-274.
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el bosque lluvioso, con referencia a la Estaci6n Biol6gica
de Cocha Cashu, Parque Nacional de Manu. Capftulo
15, Centro de Datos para la Conservaci6n, Universidad
Nacional Agraria, La Molina, Peru.
Kessler, P. 1995. Revierverhalten, nahrungsstrategie und
habitatpraferenzen des Rothandtamarins (Saguinus
midas midas) in Franzosish-Guayana. Diplomarbeit An-
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Salvador Mandujano
Luis Arturo Escobedo-Morales
Rodolfo Palacios-Silva


Individual dispersal is part of a reproductive strategy that
balances the costs and benefits for an individual as it
chooses to stay with or leave its natal group (Jones, 1995).
Dispersal in Alouatta has been documented in a number
of studies (Agoramoorthy and Rudran, 1993; Clarke and
Glander, 2004; Glander, 1992; Moore, 1992). However,
habitat fragmentation may significantly limit the options
available for an individual to move between social groups
(Swart and Lawes, 1996). In many cases, this interference
with the ability to disperse forces primates to live in small
and isolated fragments, which in turn may cause changes
in foraging and activity patterns, social organization, and
physiological conditions, leading to inbreeding that can
diminish genetic variability (Clarke et al., 2002; Gongalves
et al., 2003). The ability to disperse across fragmented
landscapes will depend on the characteristics of each species
(Bicca-Marques, 2003; Jones, 1999) as well as the spatial
configuration of the landscape in question (Fahrig, 2003).
Tischendorf et al. (2003) defined a specialist disperser as
having a low probability of crossing boundaries from habitat
to matrix, a high risk of mortality while in the matrix, and
fast movement and high inter-step movement correlation
(i.e., small turning angles between consecutive movement
steps, in matrix). In contrast, generalist dispersers have
a higher probability of leaving habitat, lower dispersal
mortality and less directed movement paths while traveling
through the matrix (i.e., larger turning angles between
consecutive movement steps).

Howler monkeys are arboreal quadrupeds and are observed
only occasionally to leave the trees and walk along the
ground (Glander, 1992). For example, A. pigra individuals
walk among the naturally patchy vegetation in the Petenes
of Yucatan (A. Estrada, pers. comm.) and Tabascan

swamps (J. C. Serio-Silva, pers. comm.). The same is true
for red howlers, A. seniculus, travelling among clumps of
trees in the Venezuelan llanos. Data on travel patterns in
continuous forest suggest that A. palliata uses routes that
minimize travel times from feeding to resting trees (Garber
and Jelink, 2004). Fedigan et al. (1998) mention that the
formation of new A. palliata groups in Santa Rosa, Costa
Rica, occurred as a result of large groups splitting, and the
dispersal of lone individuals in search of females. Glander
(1992) reported an average travel distance of 700 m for
A. palliata at Hacienda La Pacifica, Costa Rica. Individu-
als had to cross open areas to reach a new group; in some
cases, these movements occurred in several stages between
"stepping stone" fragments. In particular, dispersal success
declines with a decrease in habitat and increased fragmen-
tation of the landscape, but the rate of this decline acceler-
ates once the amount of remaining habitat falls below 10-
20% (King and With, 2002). Therefore, one might expect
that howler monkeys do not travel among fragments ran-
domly, and that the spatial configuration of habitat patches
and the nature of the surrounding matrix is critical to suc-
cessful dispersal.

The tropical rainforest in Los Tuxtlas in the Mexican state of
Veracruz has been largely deforested: 75% of native habitat
has been lost, 20% now survives only in isolated fragments,
and barely 5% is comprised of widespread contiguous
rainforest at high elevations (>800 m) (Estrada and Coates-
Estrada, 1996). Alouatta palliata, the mantled howler
monkey native to Veracruz, now survives in archipelagos
of forest fragments that vary in size, age, and degree of
isolation. Their existence in these scattered forest remnants
is precarious both ecologically and demographically, which
compounds the dilemma of dispersal (Estrada and Coates-
Estrada, 1996; Mandujano et al., in press). Here we present
data on the movements of howler monkeys in this region's
highly altered landscape and develop a preliminary model
of the probability of interchange between fragments.


This research was conducted in the Sierra Santa Marta
in the south of Los Tuxtlas, Veracruz, Mexico (1822'N,
9445'W). We defined the study area as the land-
scape between the Rfos Tecuanapa and Pilapa, covering
4,960 ha, of which only 11% is suitable primate habitat
(Fig. 1). Elevation ranges from sea level to 900 m. Corn
crops and livestock pasture make up the matrix that sur-
rounds the 92 remaining fragments, most of which are lo-
cated in riparian zones along rivers and streams, often on
slopes steeper than 30. Some fragments are on hilltops,
while others lie in permanently flooded areas. Of these
fragments, 81% are smaller than 5 ha, and only five (8%)
are between 10 and 75 ha. The mean distance between
fragments and the higher elevation continuous forest was
3,625 m, while the mean distance from one fragment to the
next was 111 m. The mean distance from any fragment to
the nearest town was 880 m.

Neotropical Primates 12(3), December 2004

We carried out fieldwork from January 2001 to July 2003.
Three times a year we conducted a census of all fragments
identified in the study area noting the presence or absence of
Alouatta groups in each. Two to five people spent 4-5 hours
sampling in small fragments (<10 ha) and 1-2 days in larger
fragments (>10 ha). We compiled a catalog of individuals
according to their identifying features and facial shapes,
plus scars and coloration patterns of the back and tail.
Although animal dispersal is broadly defined (see Clobert
et al., 2004), for the purpose of this study we consider it be
movement between fragments by individuals of any age or
sex class. To quantify dispersal, we observed the movement
of individuals into adjacent fragments and noted whether
or not they later returned to their group of origin. We also
recorded the number of solitary individuals inhabiting
fragments at the time, plus the number of individuals
entering empty fragments. Although we report here all the
movements we observed, we recognize that there were other
arrivals and departures that we were unable to observe.

We divided up all observations of the movements of
individuals between adjacent fragments into the following
distance classes: 0-100 m, 101-200 m, 201-400 m, and
401-800 m. and then calculated the proportion of the
movements in each. We observed no movements greater
than 800 m (see Results). We then fitted these data to

negative exponential, half-normal, and linear inverse models
through the least squares method using the STATISTICA
Program (StatSoft, Inc., 1998). These and other dispersal
models have been theoretically and empirically used for
investigating patterns in a number of animal and plant
species (e.g., Urban and Keitt, 2001; Mennechez et al.,
2003; Skalski and Gilliam, 2003).

In these three models, the dispersal probability decreased
as isolation distance increased between fragments; however,
the probability varied among models. For the negative
exponential model, the dispersal probability p, was
estimated as

,, = exp ( x d,),

for the half-normal model, the dispersal probability was

,, = exp (0 x d ),

while for the linear inverse model, the dispersal probability

P = 1 (- j

In all cases, 0 is a distance-decay coefficient (0 <0.0) that
determines the slope of the relationship (Urban and Keitt,

Figure 1. The landscape of the study area in Los Tuxtlas showing the fragments (numbered) occupied by Alouatta palliata (in black) and
those that were empty (in white). See Table 1 for information on each of the occupied fragments.

* Fragment occupied

Z Fragment unoccupi

Neotropical Primates 12(3), December 2004

2001). To fit the observed data, we made two assumptions.
First, if two different primate groups inhabit the same
fragment, the probability of dispersal between them is equal
to 1.0 because there is no isolation. Second, if two groups
inhabit different fragments with an isolation distance equal
or greater than 1000 m, the probability of dispersal was
equal to 0.0. The 1000 m limit was chosen considering the
very few observations of howler monkeys traveling between
groups inhabiting fragments at distances of this magnitude
(see Glander, 1992).


Overall counts ranged from 71 to 76 howler monkeys
inhabiting 19 fragments -17.5% of the 92 fragments
in the study area. Three groups of howlers (of 6, 5 and
3 individuals, respectively) lived in fragment F19 (Fig. 1
and Table 1), while, at the other extreme, one group used
four fragments. Eight fragments were each inhabited by a
separate group, while five fragments were each inhabited by
a separate solitary male. The mean isolation distance of any
one occupied fragment to any other was 2.71 0.75 km.
The mean distance of any one group to the closest fragment
was 0.33 0.39 km, while the mean distance of the howler
groups to continuous forest was 6.18 +2.31 km.

The following are some examples of movements that we

1. A male moved from his group in fragment F6 into
fragment F8, 78 m away; he remained for almost a
year, then returned to F6.

2. Another adult male left fragment F6 to inhabit frag-
ment F7, 79 m away. He remained in this fragment for
less than six months before returning to F6.
3. The entire group inhabiting F6 moved to F5, 120 m
away, and stayed there for less than one month before
returning to their original fragment.
4. At the beginning of the study, a solitary male was
found inhabiting F48. According to the field assistant
and local landowners, this individual had moved from
F19, 656 m away.
5. There were no howlers in F37 when it was first sur-
veyed, but one male who had not been recorded in
the next nearest groups, F41 and F17, subsequently
took up residence. We suspect that he moved from an
unstudied group located 80 m away.
6. Another male appeared in F37, probably from the
nearest fragment 80 m away as well.
7. An older male arrived in F37 and then moved on to
F38. Again, we suspect that this individual was from
an unstudied group located 80 m away. Local people
supported this supposition.
8. One group of six individuals initially occupied F32.
This group later divided into two, each with three
individuals; one group moved out and took up resi-
dence in the nearest fragment, F33, 47 m away.
9. Fragment F4 was empty at the beginning of the study;
six to eight months later, individuals were continu-
ously observed there. It is probable that they moved
from fragments F2 or F3, located 41 m and 92 m
away, respectively.
10. An adult male moved 171 m from F5 to F102.

Table 1. Characteristics of the study area. Annual numbers and group composition of howler monkeys and of forest fragments characteristics
in the study area. Fragments are labeled in Figure 1.
Isolation distance (m) to nearest Number of individuals
Fragment Size (ha)
Fragment Town Continuous forest 2001 2002 2003
1 11 96 1438 6704 3 4 3
2 9.3 34 2125 6169 5 5 5
3 4.7 34 2542 5900 7 8 7
15 11.8 115 4 3675 10 15 14
17 57.2 18 307 3364 5 5 6
191 29.9 196 562 3197 11 14 11
32 5.3 24 1988 4426 6 5 6
33 3.67 12 2186 4817 6 3 4
36 75.5 75 81 144 1 1 1
38 5 23 192 1184 0 0 1
41 6.5 57 625 2850 5 5 5
48 13 15 557 2660 1 1 1
5,6,7,82 14.6 43 1941 5634 10 7 8
1013 71.0 75 438 206 ? ? 2
1023 1.4 171 1970 6624 ? ? 1
iFragment 19 inhabited by 3 groups.
I 5, 6, 7 and 8 were used by one group; therefore the size is the sum of each fragment, and isolation is the mean distance.
I . 101 and 102 were sampled only once at the end of 2003; thus there is no precise data for previous years.

Neotropical Primates 12(3), December 2004

Distance of movement ranged from 15 to 656 m, but 70%
were under 100 m (Fig. 2). Data fit the negative exponential
(r2 = 0.90, F= 28.6, df = 1, 3, PP= 0.01) and half-normal
(r2 = 0.89, F= 24.6, df = 1, 3, PP= 0.02) models (Fig. 2).
In contrast, data did not fit the linear inverse model (r2 =
0.47, F= 2.7, df= 1, 3, P= 0.20). The 0 coefficients were
-0.007 and -0.000035 for the negative exponential and
half-normal models, respectively.


The landscape of our study area is characterized by consider-
able destruction and fragmentation of the natural habitats;
the remaining forest occurs predominantly in small frag-
ments with a lack of corridors between the fragments (e.g.,
riparian vegetation, live fences), and a homogeneous matrix
consisting mainly of pastures and seasonal agriculture. As a
result, Alouatta .- i, scarce, and the remaining individ-
uals inhabit only a few fragments (19 of the 92 fragments
we investigated), which are isolated from one another and
from continuous forest (Rodriguez-Toledo et al., 2003;
Mandujano et al., in press). As such, the probability that
animals will disperse from one group to another is sharply
limited by isolation distances. Considering Tischendorf et
al.'s (2003) definition of specialist and generalist dispersers,
we initially expected that each of the three dispersal models
in this study (negative exponential, half-normal, and linear
inverse) would represent the hypothetical facility with which
the monkeys could disperse from one fragment to another.
If a generalist species, the howler monkey would have a
greater capacity to move along the ground in the matrix
as it disperses from one fragment to other; in this case, the
data should be adjusted to a linear inverse model. But if the
howler monkey is a habitat specialist and more limited in
its dispersal ability, then field data should be adjusted to a

0 100 200 300 400 500 600 70 800 900 1000
Distance between fragments (m)

Figure 2. Relationship between dispersal probability and isolation
distance. The points represent the proportion of Alouatta
palliata movements at different isolation distances. Note our
assumptions (see methods) that at 0 m the dispersal probability
is 1.00; and at 1000 m the probability is 0.00. Lines represent,
from left to right, the expected probability of movements in the
exponential negative, half-normal, and linear inverse models.
Note the field data fit better to the negative exponential and
half-normal models.

negative exponential or half-normal model. The difference
between negative exponential and half-normal models
is that in the first, the probability of dispersal decreases
exponentially with a small increase in isolation distance;
in the second model, there is an isolation distance where
the dispersal probability is high, and then the probability
decreases slowly. The few data obtained during field work
showed that the most frequent movements were towards
fragments located a very short distance away (<100 m);
beyond this distance the frequency of movements dropped.
Thus, data fit better into the negative exponential or half-
normal models, indicating, therefore, that A. palliata may
be classified as a specialist disperser.

Our analysis suggests that the degree of structural heteroge-
neity in the landscape may be an important factor determin-
ing the possibility of dispersal by primates across it. In het-
erogeneous landscapes, primates may use forest fragments as
well as tree plantations (for example, shade coffee and cacao
agroecoystems) as stepping stones or corridors when moving
from one forest patch to another (Estrada et al., in press).
In highly homogeneous landscapes (i.e., dominated by pas-
ture or other monocultures, with scant tree cover) exchange
of individuals among fragments is more difficult. Using
percolation models, it has been found that dispersal success
declines with increasing fragmentation of the landscape,
with this decline accelerating once the amount of suitable
habitat falls below 10-20% (King and With, 2002). The
connectivity of habitat patches in a landscape, therefore,
depends on the dispersal capacity of the individuals. In the
study landscape, connectivity is currently low (<30%) given
the high rate of habitat destruction and loss, with only 11%
of the original vegetation remaining, and the relatively low
dispersal capacity of howler monkeys through the matrix
(Palacios-Silva and Mandujano, in press).

Habitat connectivity is a central theme in both metapopu-
lation ecology and conservation biology (Bennett, 2004).
As the number of occupied fragments decreases, so too does
the probability of persistence on a regional level decline,
due to a possible imbalance between the extinction rate of
local populations and the colonization rate (Ovaskainen and
Hanski, 2004). Therefore, if the dispersal rate proves lower
than the deforestation rate, the only conservation alterna-
tive on a regional level would involve habitat rehabilitation
in an effort to create corridors and stepping stones, plus
the translocation of individuals and/or groups to other frag-
ments offering better survival conditions. In particular, the
creation of stepping stone fragments could be an alternative
management action that increases connectivity and could
allow movement among primate groups (Mandujano et al.,
in press). Basically, a stepping stone can be any landscape
element that the animal perceives as a transitional step
leading toward a habitat patch (Bennett, 2004). For
primates, a stepping stone can be a group of isolated trees,
live fences that separate strips of land, riparian zones,
corridors, remnants of arboreal vegetation and/or habitat
patches that are substantially smaller than an animal's home
range. Studies showed that for species with limited dispersal

Neotropical Primates 12(3), December 2004

ability and a landscape with isolated habitat, stepping stone
habitat patches greatly increase a species' ability to disperse
(Bennett, 2004; King and With, 2002).

Evidently, more factors than isolation distance contrib-
ute to an individual's dispersal between habitat fragments
(Clobert et al., 2004). For example, in a 30-year study
of A. palliata at La Pacifica, Clarke and Glander (2004)
found that female migration patterns were primarily
associated with environmental variables (habitat, rainfall)
and secondarily with social variables (number of females in
the group, sex ratio); while male migration patterns were
only associated with social variables (i.e., male-female ratio
and male age). In contrast, from a census of 333 howlers
in a recent study in the northern part of Los Tuxtlas,
Cristobal-Azkarate et al. (2004) found 16 solitary males
and only one solitary female. These authors suggest that
females remain in their natal groups, probably reinforced
by habitat fragmentation. Thus, the relationships between
intrinsic factors (social, demographic, and genetic), habitat
factors (quantity, quality, and spatial configuration), and
human factors (deforestation, hunting, and others) that
influence the decision of an individual monkey to stay
or leave its natal group are complex. In consequence, the
dispersal models presented here, based only on isolation
distance between fragments, are preliminary, and field and
experimental data will be required to test their accuracy
and general application.

Acknowledgments: The authors thank the Mateo-Gutierrez
family for their hospitality and invaluable help during
the fieldwork, and also E. M. Rodriguez-Toledo and
A. Gonzalez-Zamora for their assistance in the field. A.
Estrada and G. Echaverria-Lozano reviewed an early
version of this manuscript. The Department of Biodiversity
and Animal Ecology of the Instituto de Ecologia A. C.
provided the support necessary for the completion of this
research. The American Society of Primatologists (ASP) and
Primate Conservation, Inc. provided additional support
for some aspects of this study through their respective
small-grants programs.

Salvador Mandujano, Luis Arturo Escobedo-Morales,
and Rodolfo Palacios-Silva, Instituto de Ecologfa A. C.,
km 2.5 Carret. Ant. Coatepec No. 351, Congregaci6n
del Haya, Xalapa 91070, Veracruz, Mexico. E-mail:


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Maren Huck, Petra Lottker
Eckhard W Heymann, Michael Heistermann


In order to recognize differences in ontogeny between species
and between their life history trajectories, it is necessary to
have an understanding of the timing of milestone events in
behavior and physiology (Pereira and Leigh, 2003). Assessing
when and how individuals become independent from their
mother and other caregivers is necessary to evaluate costs
of varying parental strategies, which in turn allows broader

issues to be addressed, such as the evolution of non-maternal
care in anthropoid primates (Ross and Maclarnon, 2000).
Patterns of ontogeny may also explain aspects of individual
differences in behavior, especially, for example, when same-
sexed adults become competitors. Ventura and Buchanan-
Smith (2003) have stressed that a good understanding
of the species' development and behavior in the wild is
essential to assess and ensure the well-being of maturing
individuals in captivity. Observations are often anecdotal,
but the difficulties and imprecision in obtaining data of this
sort under field conditions can be overcome by repeated
studies of groups of the same or related species.

The ontogeny of immature primates is tied to endocrino-
logical changes. Typically the gonads are only partially
developed at birth. The production of sex hormones such
as testosterone increases during puberty until adult levels
are reached (Plant, 1988). In adult males, testosterone and
other androgens are important in the control of spermato-
genesis (Wickings etal., 1986) and seem to play a role in the
expression of aggressive behavior and in the achievement of
social status, though this latter relationship might be less
pronounced in primates (Dixson, 1980; Bouissou, 1983).
In three callitrichine species (( .-'. jacchus, (
kuhlii, and Saguinus oedipus), however, newborn males
show relatively high testosterone levels that drop after two
to six months and rise slowly again thereafter (Abbott, 1984;
French and Schaffner, 1995; Ginther et al., 2002). Ginther
et al. (2002) speculated that this neonatal elevation might
be related to the sexual maturation of the gonads, endocrine
system, and behavior, but it is not known how representa-
tive this pattern may be of other callitrichine species.

During a field study on moustached tamarins (Saguinus
mystax), we were able to observe some aspects of the
development of six immature individuals (five males and
one female). For some of them we were able to record, for
the first time, the onset and final appearance of a number of
behaviors in particular, the first observations of feeding
on solid food, foraging, social play, and marking behavior,
and the last days they were carried and successfully begged
for food.

We expected that the patterns of testosterone and androgen
in the five male immatures would be similar to those of
the three callitrichines mentioned above. In addition, the
immigration of a new female into one of the groups on 5
December 2001 provided the opportunity to study whether
this caused endocrinological changes in the immature males.
Given that one subadult male was evicted from the group
following the new female's arrival, we also tested whether he
and his twin differed in their testosterone levels, and whether
their values differed from those of adult males in general.
If co-twins differ in their sexual maturation, it might be
supposed that the more mature of the two would be forced
to leave the group in order to remove a potential competitor.
If this is true, it might be expected that the evicted twin had
higher testosterone levels than the remaining one. Different
testosterone levels due to different aggression levels in the

Neotropical Primates 12(3), December 2004

twins are unlikely, since we were able to show previously that
adult moustached tamarins do not differ in their testosterone
levels despite different breeding and "dominance" status
(Huck et al., 2005b; note: aggression occurs too rarely to
establish hierarchical relationships). These results should
be considered preliminary owing to our small sample size,
but they nevertheless present the first data on the behavioral
ontogeny of Saguinus mystax and the first endocrine data for
immature males of any callitrichine in the wild.


The study took place in 2001 (January to December) at
the Estaci6n Biol6gica Quebrada Blanco, in northeastern
Amazonian Peru (for further details of the study site,
see Heymann, 1995). We collected fecal samples and
behavioral data from two habituated groups of S. mystax,
each with two sets of offspring, one born in 2000 and the
other in 2001. Group W contained one adult female, three
adult males, male twins born in February 2000 (WM4 and
WM5), and a single male offspring born in February 2001
(WM6). Group E consisted of three adult females and three
adult males, a single male born in May 2000 (EM4), and
twins one male (EM6) and one female (EF4) born in
January 2001. Based on physical characteristics, Soini and
Soini (1990) classified moustached tamarins as infants from
birth until about three months of age, as juveniles between
4-12 months, as subadults between 13-18 months, and
as adults thereafter. Following these criteria, three of the
immature males (WM4, WM5, EM4) reached adulthood
at the end of the study in December 2001. Two males
(WM6, EM6) and one female (EF4) were born near the
beginning of the investigation, developing from newborns
to relatively independent juveniles over the course of the
year (see Table 1). In Group W the breeding female WF1,
mother of WM6 and (probably) aunt of WM4 and WM5
(relationships were evaluated by microsatellite analysis as
described by Huck et al., 2005a), died near the end of the
study (3 December, 2001). Two days later an unrelated
female (WF3) immigrated to Group W, and one of the male
subadults (WM4) was evicted on 5 December 2001. The
composition of the groups and major demographic events
are described in more detail by Lsttker et al. (2004).

We followed each group daily from the time they left
their sleeping site (about 05:45) until they entered their
sleeping site in the afternoon (about 15:45 h). From
January to December 2001 we accumulated a total of
3004 and 3257 contact hours over 330 and 351 days for
groups W and E, respectively. The average visibility of
individuals measured as the percentage of hourly scans in
which each individual could be seen was 18.0 and 18.6%
for adults and immatures in group W, respectively, and 11.3
and 10.9% for adults and immatures in group E. Following
the protocols for behavior sampling of Martin and Bateson
(1993), we recorded the first observation of an infant
independent of a carrier (being "off"), eating solid food
(without distinguishing between prey or fruit), engaging
in social play, and foraging for prey. We also noted the last
observations of an infant being carried and of successfully
begging for food. Except for the data on food-begging, the
records were taken from the younger litter in the two groups
(i.e., WM6 in group W, and EM6 and EF4 in group E). We
were unable to distinguish the Group E twins, EM6 and
EF4, until they were 81 days old; until then, data of the first
occurrences of either one of the two infants are used. Data
on food-begging were recorded from the older immatures
(WM4 and WM5 in group W, EM4 in group E).

For hormonal analyses, we collected a total of 151 fecal
samples from immature males between January and
December 2001 (see Table 1 for the number of samples
per individual). Immediately after an animal was seen
defecating, the feces were collected and immersed in
96% ethanol. (The protocols for treatment and storage
of the samples are given in Huck et al. [2005b], as are
techniques for hormone extraction and measurement of
immunoreactive testosterone by enzyme immunoassay.)
Hormone concentrations are given in ng/g fecal dry weight
and were log-10-transformed for parametric statistics.

In order to examine changes in testosterone excretion
over time, we calculated Pearson's correlation coefficient
(r) between age and mean monthly testosterone values. As
in ( .-' jacchus, ( kuhlii and Saguinus oedipus,
testosterone levels in newborns drop only after two to six
months and rise slowly again thereafter (cf. Abbott, 1984;

Table 1. Immatures of Saguinus mystax study groups W and E, with mean testosterone values of the males and number of fecal samples
analyzed (N).
Individual Sex Demographic notes Mean testosterone +S.D. N
Group W
WM4 M Born 22 February, 2000; ...... ... I 5 2720.0 3274.9 35
December, 2001 (before 18 mo: 3219.3 4046.6)
WM5 M Born 22 February, 2000 1918.6 3185.1 40
(before 18 mo: 1435.6.3 1160.7)
WM6 M Born 24 February, 2001 470.8 549.4 28
Group E
EM4 M Born May, 2000 990.3 1057.1 34
EM6 M Born 21 January, 2001 814.1 915.0 14
EF4 F Born 21 January, 2001

Neotropical Primates 12(3), December 2004

4.5 -- WM4

4. .........+ EM4 *
----A A A


+ \. p ......... .. A,

2.5 I ++ + + +

+ A
1 1.5

90 120 150 180 210 240 270 300 330 360 390 420 450 480 510 540 570 600 630 660
Age [days]

Figure 1. Log-10-transformed fecal hormone concentrations for all individuals related to relative age (in days). Curves give the distance-
weighted least-squares fit for each male. Only the month, not the exact date of birth was known for EM4, so it was taken to be 15 May.

French and Schaffner, 1995; Ginther et al., 2002). The data
depicted in Figure 1 appear to show a drop in testosterone
levels between months 4 and 5 (around day 140). Therefore
we also calculated the coefficient r for the younger males
WM6 and EM6, excluding the potentially high values
before their 140th day of life, which seems to correspond
to the turning point in testosterone production described
for ( .-' jacchus, ( .'. kuhlii, and Saguinus oedipus
at similar ages (Abbott, 1984; French and Schaffner, 1995;
Ginther et al., 2002). Three samples were collected from
WM6 before he was 140 days old and one for EM6.

Unfortunately we could not obtain enough samples for
statistical analysis after WFI's death (3 December 2001)
and the subsequent emigration of WM4 (5 December
2001); we collected only one sample each for WM4 and
WM6. Instead, we evaluated whether the mean hormone
concentration of these samples collected after the new
female had immigrated lay above the 95% upper boundary
of the group average when WF1 was still alive. In addition,
we compared testosterone levels between the twin brothers
in group W and in relation to mean testosterone levels of
the adult males in the two study groups W and E (values for
adult males appear in Huck et al., 2005b).


Our first and last observations of milestone events in behavior
are listed in Table 2. Nursing was observed only rarely and
was not included. Apart from "last time seen successfully
begging", the data report on the second litters only (WM6 in
group W, and EM6 and EF4 in group E). Table 1 shows the
testosterone concentrations of all samples by individual and
the distance-weighted least-squares fit for each. No immature
showed a significant increase in testosterone levels during the
year, and none of the correlations between age and mean
monthly testosterone values were significant (WM4: r =

0.06, p = 0.86; WM5: r = 0.37, p = 0.23; WM6: r = -0.03, p
= 0.94; EM4: r = 0.21, p = 0.50; EM6: r = 0.11, p = 0.83).

When we calculated the regressions for WM6 and EM6
after their 140k' day of life, only WM6 showed a significant
increase in testosterone values over the following six months
until December 2001 (r = 0.50, p = 0.012, see Fig. 1);
we did not find such a correlation for EM6 for the seven
months after his 140th day of life until December 2001
(r = -0.27, p = 0.37; see Fig. 1).

We had no samples from WM4 after the death of the
breeding female WF1. The samples of WM5 and WM6,
collected after immigration of the new female, both fell above
the 95% boundary (Table 3). WM5 had significantly lower
testosterone values than the adult males of group W, but the
values of WM4 did not differ significantly from adult levels

Table 2. First and last occurrence of behavioral milestone events
in two study groups of S. mystax.
Group W Group E
First day "off" carrier'' Day 17 Day 21
First day seen eating Day 30 Day 33
solid food, Day
First day seen socially Day 57 Both: Day 37
First day seen foraging' Day 103 Day 61
EM6: Day 104-105
Last day carried Day 106 EF4: Day 104-105
EF4: Day 114
WM6: Day 99
First time seen marking Day 154 WF6: Day 99
WF4: Day 114
Last day of successful WM4: Day 325 EM4: 1 year 1
begging' WM5: lyear 8 days month
N = One individual in group W (WM6) and two in group E (EM6 and
Infants in group E (EM6 and EF4) were too small to be distinguished
N = Two individuals in group W (WM4 & WM5) and one in group E

134 Neotropical Primates 12(3), December 2004

Table 3. Mean (untransformed) testosterone values and the upper 95% boundary of juveniles of Group W during the tenure of WF1, and
the value of a sample collected after the immigration of a new female.

Individual Mean (with WF1) 95% boundary Mean of last 3 samples with WFI Sample with new female
WM4 2720.0 10615.6 3017.9
WM5 1471.3 4740.2 1487.7 19361.4
WM6 413.4 1488.9 782.3 2022.1

(mean and SD for adults: 2782.1 3233.3; for WM4 and
WM5: see Table 1. Scheffk post hoc tests: adults vs. WM4:
p = 0.72; adults vs. WM5: p = 0.014). The testosterone
levels of the twin brothers (WM4 and WM5) did not differ
significantly but tended to be lower in WM5 (Scheffk post-
hoc test: p = 0.30; compare fit curve in Fig. 1).


In general, the two study groups showed consistent results
but differed in two ways. First, WM6 began foraging
and showing social play later than his peers from group
E. This might have been due to his having been raised as
a singleton; his sibling was killed by a raptor at the age
of 28 days (Oversluijs VAsquez and Heymann, 2001).
Other studies have shown that infants prefer to play with
same-aged playmates and that singletons play significantly
less than twins (e.g., Cleveland and Snowdon, 1984).
This holds true even when older siblings are available as
potential playmates (Cleveland and Snowdon, 1984).
Without further study we cannot say whether our results
are representative and, if they are, whether the reduced
play retarded the development of motor skills vital to
foraging or whether with only a single infant to care
for-caregivers are more indulgent, resulting in less need
for an infant to forage for itself.

The only slight (non-significant) increase of testosterone
levels with increasing age in all subjects was unexpected.
WM6 showed a testosterone excretion pattern similar
to those described for other callitrichines (Abbott, 1984;
French and Schaffner, 1995), with high levels very early in
life that drop after three months and then rise again after
the seventh month. The similar-aged EM6 did not show
this pattern, but this might be due to the fact that only
one fecal sample was analyzed before his 140th day of life.
There was a strong increase in testosterone concentrations
in the samples of WM5 and the considerably younger
WM6 collected after the death of the related WF1 and the
subsequent immigration of the unrelated female WF3.

With only one fecal sample in each case, this finding must
be regarded with caution, but we suspect that the elevated
levels might be related to the fact that incest avoidance no
longer had a role to play (see also Baker et al., 1999, for
(C .- ... In addition, WM4, who was later evicted,
seems to have been more precocious than his brother.
Although the twins in group W showed no differences in
time budgets (G-test with William's correction, G6 = 11.6,
p = n.s.; unpubl. data), their endocrine levels were different

(although non-significantly), and WM4 may have posed
more of a threat to the adult males after the arrival of the
unrelated female WF3. The testosterone values of WM5
were significantly lower than adult levels, while those of his
brother were not. WM4 had apparently reached adult levels
before the onset of the study when he was 10.5 months
old. In contrast, his twin WM5 reached adult levels only at
the end of the study at nearly 23 months old (cf. Ginther
et al., 2002, for similar ranges). Age and morphology alone
are thus insufficient predictors of sexual maturity, since
testosterone levels, which are likely to be related to sperm
production and fertility, differ widely during maturation
even between twins.

Acknowledgments: We thank our field assistants for their
invaluable help in collecting samples. Fieldwork was carried
out under a letter of understanding between the Universidad
Nacional de la Amazonfa Peruana (UNAP), Iquitos, and
the Deutsches Primatenzentrum, G6ttingen, and complied
with Peruvian laws. Financial support was provided by the
DFG (HE 1870/10-1,2).

Maren Huck, Department of Biology & Environmen-
tal Science, John Maynard Smith Building, University
of Sussex, Brighton BN1 9QG, Great Britain, e-mail:
, Petra Ldttker, Abteilung
V, 1- ilr, ...l...1. -.i1 & Soziobiologie, Deutsches Primaten-
zentrum, Kellnerweg 4, 37077 Gbttingen, Germany, e-mail:
and Institut fur Neuro- und Ver-
haltensbiologie, Abteilung Verhaltensbiologie, N. I .rFtilI,. -J
Wilhelms-Universitat, Mtinster, Germany, Eckhard W
Heymann, Abteilung L1, ilr, r..l..1..l.. .1 & Soziobiologie,
Deutsches Primatenzentrum, Kellnerweg 4, 37077 Gdttin-
gen, Germany, e-mail: and Michael
Heistermann, Abteilung Reproduktionsbiologie, Deutsches
Primatenzentrum, Kellnerweg 4, 37077 Gbttingen, Germa-
ny, e-mail: .


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social relationships in higher mammals. Hormone Res. 18:

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Cleveland, J. and Snowdon, C. T. 1984. Social development
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Jodo M. D. Miranda, Itibere R Bernardi,
Rodrigo F Moro-Rios, Lucas M. Aguiar,
Gabriela Ludwig, Fernando C. Passos


The social group of the brown howler monkey, Alouatta
guariba clamitans Cabrera, 1940 is typically small (2-12
individuals), with one or two adult males, and more females
than males (Silva Jr., 1981; Neville et al., 1988; Mendes,
1989; Steinmetz, 2000, 2001; Treves, 2001; Miranda and
Passos, 2005). In primates where males outnumber females,
males compete for both females and food, which normally
results in groups with dominant males (Clutton-Brock and
Harvey, 1977; Dunbar, 1988; Mittermeier et al., 1999). As
in all howler monkeys, A. g. clamitans is sexually dimorphic;
adult males are larger than females and are dominant to
them (von Ihering, 1914; Altmann, 1959; Neville et
al., 1988; Bonvicino, 1989; Mendes, 1989; Chiarello,
1995; Mittermeier et al., 1999; Treves, 2001). Although
howler monkeys are the most studied of the Neotropical
primates, most of the research has been concentrated on
a few species, in particular A. palliata and A. seniculus,
and only more recently A. caraya, A. pigra and A. guariba
(Neville et al., 1988; Bicca-Marques, 2003). Field studies
on the composition, social structure and hierarchy of A. g.
clamitans are rare (Miranda, 2004; Jardim, 2005; Miranda
and Passos, 2005), and long-term investigation will be
necessary to obtain more detailed information on their
social patterns. Here we describe a particular case of an A. g.
clamitans group in which the dominant animal was an adult
female-in effect, a matriarchal group.


Study area
The study area is a forest fragment of approximately 700
ha in the ChAcara Payquere, a component of the Area
de Protecao Ambiental da Escarpa Devoniana, in the
municipality of Balsa Nova, Parana, Brazil (25o29'52"S,
49O39'24"W). This remnant forest is within the Floresta
Ombr6fila Mista (Araucaria Pine Forest biome) and consists
mainly of disturbed primary forest, broken by patches of
secondary forest.

Observations of the behavior and ecology of brown howler
monkeys in the study area have been continuous since
February 2002 (Miranda, 2004; Miranda and Passos, 2004,
2005; Miranda et al., 2005; Miranda et al., in press). The
study group, the Forninho Group, was the best known in the
area while its members remained together. Our observations
here were recorded ad libitum (Altmann, 1974). We used
the system of age-sex classification proposed by Mendes
(1989) and used by Miranda and Passos (2005).

Neotropical Primates 12(3), December 2004


In February 2002, the Forninho Group included two adult
males (AM), three adult females (AF), and a juvenile I (JI).
After four births through two years of study (two births in
2003 and two in 2004), the group consisted of two AM, a
sub-adult male (SAM), three AF, two juveniles II (JII), one
male and one female, and two JI (both males). In January
2004 the dominant adult male (AM1) disappeared, which
apparently disrupted the social structure of the group. This
resulted in the Forninho Group splitting into two: the
Carrano Group with one AM, one AF and one JII (female),
and the Vava Group with one SAM, two AF (AF 1 and AF2),
one JII (male), two JI (males) and one infant (INF) born
one month after the disappearance and presumed death of
AMI (Table 1). Another birth occurred in July of 2004,
this time to an AF2 in the Vava Group. Miranda and Passos
(2005) give the details of the dynamics of this group.

AF1 of the Vava Group was the mother of the SAM, as well
as the JII, one of the JI and the older INF. AF1 may also
have been the mother of AF2, but we could not confirm
this because AF2 was already an adult at the beginning of
our study. If AF1 was indeed the mother of AF2, then every
member of the Vava Group would have been the progeny of
this single adult female.

In confrontations with neighboring groups, AF1 vocalized
the most (bark and howl sensu Oliveira, 2002) and
challenged the dominant AM of Carrano Group, forcing
him away from the confrontation. This female behaved in
the same way toward humans when one of the investigators
accidentally startled the animals, alarming the group and
provoking AF 1 to come their defense. On two occasions,
AF1 challenged the subadult male over food. The male
retreated both times.


The breakup of the original Forninho Group may have been
prompted by AM2, who was possibly the son of AF1, and,
as such, leaving the group to avoid endogamy. Following

Table 1. Composition of rhe Forninho social group and of the re-
sulting (Vavi and Carrano) groups which were formed after it split
up, coincident with the disappearance of the adult male AM1.
Before splitting January2004 After splitting
Forninho Group Disappearance of Vavi Group
3 AF 1 SAM
2 JI 1 INF (born Feb 2004)
Total = 10 Total = 7
Carrano Group
Total = 3

his departure, AM2 became the nucleus of the new Carrano
Group. However, given the occurrence of infanticide in
howler monkeys (Clarke, 1983; Zunino etal., 1986; Rumiz,
1990; Clarke et al., 1994; Galetti et al., 1994; Calegaro-
Marques and Bicca-Marques, 1996; Palacios, 2000; Aguiar
et al., in press), it is also possible that AF1 may have forced
AM2 out of the group, since AF1 was pregnant by AM1,
who had been the dominant male until his disappearance.
AF 1, therefore, may have been trying to protect her infant
from possible infanticide by AM2. Mendes (1989) observed
a similar case in which an AM drove out the dominant male
of his study group. Although successful in his ouster of the
dominant male, the AM in Mendes' study group was forced
out by the two adult females of the group, primarily by the
adult female who was pregnant. This behavior on the part
of the female ended only when her infant disappeared, and
she finally accepted the presence of the new male. Mendes
(1989) interpreted these events as an attempt by the pregnant
female to avoid infanticide.

The AF1 was followed by her offspring with AM1 who
had disappeared-INF, JI, JII and SAM, and possibly
AF2-which gave rise to a group dominated by a female
rather than a male. AF1 remained dominant despite
the presence of the subadult male; she was the one who
defended the group against neighboring howlers and the
researchers who accidentally startled her group. Hirano
(2003), studying A. guariba, and Calegaro-Marques and
Bicca-Marques (1997), studying A. caraya, have reported
that adult females are dominant to subadult males.

In howler monkeys, individuals of both sexes emigrate,
forming new groups or simply integrating with another
group (Neville etal., 1988; Mendes, 1989; Bonvicino, 1989;
Calegaro-Marques and Bicca-Marques, 1996; Giudice, 1997;
Brockett et al., 2000; Ostro et al., 2001; Jardim, 2005).
Bonvicino (1989) and Mendes (1989) recorded only subadult
males as solitary individuals. We saw solitary adults of both
sexes in our study area, showing that females emigrate as well
as males (Miranda, 2004; Miranda and Passos, 2005). The
subadult male of the Vava Group could have remained in the
group to become the dominant adult, or else moved out to
avoid endogamy. Groups without adult males may be more
susceptible to predation, as well as to infanticide by strange
adult males or to an upset in group hierarchy (Zunino et
al., 1986; Dunbar, 1988; Clarke et al., 1994; Galetti et al.,
1994; Palacios, 2000). We would expect that sooner or later
an adult male, either the resident subadult or an immigrant,
would take over as the dominant individual in the group.

Groups of A. g. clamitans are usually dominated by males,
but they may be adaptable enough to change this pattern
in certain cases. We believe that this is evident in the case
of our original focal group, the Forninho Group, in which
an adult female left in order to avoid imminent infanticide,
establishing, at least temporarily, a female-dominated group.

Acknowledgments: We thank Edner L. Rosa for permission
to work on his property and Kaue Cachuba Abreu for his

Neotropical Primates 12(3), December 2004

help with the fieldwork. We also appreciate the assistance of
Dr. Albert Leyva in preparing the manuscript.

Joao M. D. Miranda, Postgraduate in Zoology UFPR,
Rua Rio Guapord 1275, Bairro Alto, Curitiba, 82840-
320 Parand, Brazil, e-mail: ,
Itiber6 P. Bernardi, Graduate in Biology URI do Alto
Uruguai e das Miss6es, Rua Guerino Cerutti 161, ap. 702,
Centro, 98400-000 Frederico Westphalen, Rio Grande do
Sul, Rodrigo E Moro-Rios, Graduate in Biology UFPR,
Rua Wilson Valdfvia Domingues 203, casa 6, Bairro Jardim
das Am&ricas, Curitiba 81540-170, Parand, Lucas M. Aguiar
and Gabriela Ludwig, Postgraduate in Zoology UFPR,
Rua Bronislau Ostoja Roguski 649, Casa 3, Bairro Jardim
das Amdricas, Curitiba 81540-080, Parand, and Fernando
C. Passos, Laborat6rio de Biodiversidade, Conservacao e
Ecologia de Animais Silvestres, Departamento de Zoologia
- UFPR, Caixa Postal 19020, Curitiba, Parand, Brazil.


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Neotropical Primates 12(3), December 2004

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190. Sociedade Brasileira de Primatologia, Brasilia.


Fldvia Koch
Jilio Cesar Bicca-Marques

Primates are distinguished from other groups of animals by
having large brains, enhanced manipulative abilities, and
more complex cognitive skills (Garber, 2004). These traits
allow nonhuman primates to perform complex behaviors
such as tool use, a behavior previously considered to be lim-
ited to humans (Panger, 1998). Most instances of manipu-
lating objects as tools have been recorded in apes (Beck,
1975; Goodall, 1964). There are some records for Old
World monkeys, including baboons (Papio) and macaques
(Macaca) (Van Lawick-Goodall et al., 1973; Tomasello and
Call, 1997; Westergaard, 1992); in New World monkeys,
tool use has been observed in capuchin monkeys (Cebus)
(Beck, 1972, 1975; Chevalier-Skolnikoff, 1989; Fragaszy et
al., 2004; Ottoni and Mannu, 2001; Phillips, 1998; Struh-
saker, 1977; Vauclair and Anderson, 1994; Visalberghi,
1990; Westergaard, 1988).

Beck (1975) defined tool use as "the manipulation of an
unattached environmental object, the tool (not part of the
user's body), to alter more efficiently the form or position
of a separate object, when the user holds or carries the tool
in toto during or just prior to use and is responsible for the
critical connection between tool and incentive" (p.414).
Urbani and Garber (2002), however, warned that several

reports of tool use cited in the scientific literature are better
classified as "proto tool-use or object manipulation." True
tool use involves the detachment and manipulation of
both the object of change and the agent of change (the
tool), whereas in proto tool-use, only the object of change
is detached and manipulated (Panger, 1998; Parker and
Gibson, 1977).

Here we report a case of substrate manipulation by a
brown howler monkey (Alouatta guariba clamitans Ca-
brera, 1940). It was recorded by E Koch during a study of
the ecology and behavior of a group of brown howlers at
the Morro da Extrema (30o12'S, 51004'W), Porto Alegre,
Rio Grande do Sul, Brazil. On 16 October 2002, around
15:00 h, the sky became overcast and the wind picked
up, signaling an approaching rainstorm. The study group
began moving away. At 15:20 h, the group came to a gap
in the canopy of about 2 m. All successfully leaped across
the gap to the next tree except an infant (in the process
of becoming independent from its mother). The branches
were blowing about vigorously because of the high winds,
and the infant stopped and vocalized (cried) while hold-
ing onto the end of the branch. The group members did
not return to help the infant, which made no attempt to
jump but continued vocalizing loudly, until eventually its
mother went back to rescue it. In order to help her infant,
the mother manipulated a nearby branch (without detach-
ing it) of the tree she was in until it was positioned close
to the infant. The infant immediately used this branch as
a bridge to traverse the gap. Once safely across, it quickly
climbed onto its mother's back. Given the configuration
of the arboreal canopy, the only way for group members
to cross the gap was by leaping from one tree to the other.
A similar situation involving the same mother-infant pair
was observed on a second occasion when there was a strong
wind but a clear sky.

This note reports the observation of a complex behav-
ior performed by a howler monkey to solve a problem
commonly faced by arboreal primates. This is the first
record of the manipulation of an object to help an infant
howler monkey travel across a gap in the canopy. Previous
reports indicate that adult howler monkeys may use their
bodies to form a "bridge" in order to help immatures cross
such gaps. According to the definition proposed by Beck
(1975), this behavior cannot be considered as true tool use
because the animal (mother) did not detach the branch
used as a bridge from the tree. This use of the substrate
as an object, however, can be classified as proto-tool-use
or object substrate manipulation (sensu Parker and
Gibson, 1977).

Acknowledgments: We thank Paul A. Garber and Anthony
B. Rylands for critical comments on an earlier version of
this manuscript, Denise Brutto for permission to work at
the study site, and the Fundacao de Amparo a Pesquisa do
Estado do Rio Grande do Sul/FAPERGS for financial sup-
port (Research Grant nr. 01/1136.4 to JCB-M, and a Re-
search Fellowship to FK).

Neotropical Primates 12(3), December 2004

Flivia Koch, Instituto de Biociencias, Universidade Federal
do Rio Grande do Sul, Porto Alegre, Rio Grande do Sul,
and Jilio C6sar Bicca-Marques, Laborat6rio de Primato-
logia, Faculdade de Biociencias, Pontificia Universidade
Cat61lica do Rio Grande do Sul, Av. Ipiranga 6681, Predio
12A, Porto Alegre 90619-900, Rio Grande do Sul, Brazil,
e-mail: .


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and Psychology in Primates, R. Tuttle (ed.), pp.413-447.
Mouton, The Hague.
Chevalier-Skolnikoff, S. 1989. Spontaneous tool use and
sensorimotor intelligence in Cebus compared with other
monkeys and apes. Behav. Brain Sci. 12: 561-627.
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Oliveira, M. G. 2004. Wild capuchin monkeys (Cebus
libidinosus) use anvils and stone pounding tools. Am. J.
Primatol. 64: 359-366.
Garber, PA. 2004. New perspectives in primate cognitive
ecology. Am. J. Primatol. 62: 133-137.
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201: 1264-1266.
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tufted capuchins (Cebus ..I/i/) spontaneously use tools
to crack open nuts. Int. J. Primatol. 22: 347-358.
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capuchins (Cebus capucinus) in Costa Rica. Am. J. Phys.
Anthropol. 106: 311-321.
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tool use and sensorimotor intelligence as feeding
adaptations in Cebus monkeys and great apes. J. Hum.
Evol. 6: 623-641.
Phillips, K. A. 1998. Tool use in wild capuchin monkeys.
Am. J. Primatol. 46: 259-261.
Struhsaker, T. T. 1977. Palm-nut smashing by Cebus a.
..I/i / in Colombia. Biotropica 9: 124-126.
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Oxford University Press, Oxford.
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1973. Tool use in free-living baboons in Gombe National
Park, Tanzania. Nature, Lond. 241(19): 212-213.
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tool use, and the social context in human and non-human
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Westergaard, G. C. 1988. Lion-tailed macaques (Macaca
silenus) manufacture and use tools. J. Comp. Psychol. 102:
Westergaard, G. C. 1992. Object manipulation and the use
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Comp. Psychol. 106: 398-403.


Fabiano R. Melo, Adriano G. Chiarello
Michel B. Faria, Pedro A. Oliveira
Rafael L. A. Freitas, Fernando S. Lima
Daniel S. Ferraz


O muriqui-do-norte (Brachyteles hypoxanthus) e conside-
rado uma das 25 esp&cies de primatas mais ameagadas do
planet. Tern sua ocorr&ncia restrita a Mata Atlantica e suas
populag6es se encontram ameagadas pela destruigao e frag-
mentagao do habitat e tambem pela atividade de caga (Mit-
termeier et al., 2005). Essa situacao e considerada ainda
mais critical se levarmos em conta que a area de distribui-
gao geogrifica original da esp&cie se encontra localizada na
regiao Leste do Brasil, onde as ao6es antr6picas foram mais
several (Mittermeier et al., 1989). Como se nao bastasse,
o muriqui e um dos mamfferos mais cacados nesta regiao,
conforme relatos de caga recentes para o muriqui-do-sul
(Brachyteles arachnoides, em Mittermeier et al., 1982; 1987;
1989; Mittermeier e Konstant, 1990; Auricchio, 1997) e
para o pr6prio muriqui-do-norte (B. hypoxanthus, em Co-
senza e Melo, 1998).

Apesar de ter ampla distribuigao no leste brasileiro, sao
conhecidas populagoes de muriqui-do-norte apenas para
Minas Gerais e Espirito Santo (Strier e Fonseca, 1996-
1997). No estado da Bahia, os 6ltimos registros feitos por
Aguirre (1971) remontam a d&cada de 60. Desde essa data,
nenhuma populagao de muriquis foi confirmada no estado
baiano, ate o ano de 2004.

Desde 1999, o Instituto Estadual de Florestas de Minas
Gerais e a Universidade Federal de Minas Gerais vem de-
senvolvendo trabalhos de reconhecimento da fauna de ma-
mfferos, considerando o grupo de primatas como indica-
dor de novas areas, nos vales do rio Jequitinhonha e Doce,
visando preservar essa rica biodiversidade (Hirsch, 2003;
Melo et al., 2002; Melo, 2004). Este estudo concentrado
trouxe detalhes sobre os principals fragments florestais
existentes nas referidas bacias e permit, hoje, direcionar
recursos e esforgos conservacionistas nas areas apontadas
como de importancia para a conservagao dessa fauna de
primatas diagnosticada e, conseqiientemente, dos demais
mamfferos e das demais especies tipicamente florestais.

Concomitante a essas iniciativas, o Ministtrio do Meio
Ambiente abriu um edital, em 2002, convocando institui-
9oes de pesquisa e ensino na tentative de melhorar nosso
conhecimento biol6gico acerca dos principals biomas brasi-
leiros, baseado nos diversos workshops nacionais que foram
realizados na 6ltima decada e no infcio dessa (Brasil, MMA,
2002). A Conservacao Internacional do Brasil associou-se as
principals instituigoes de ensino de nfvel superior do Estado

Neotropical Primates 12(3), December 2004

(Universidades Federais de Minas Gerais e de Vigosa, Uni-
versidade do Estado de Minas Gerais, campus de Caran-
gola e a Pontificia Universidade Cat61lica de Minas Gerais
- PUC Minas) e teve aprovada sua proposta de inventario
da biodiversidade nos vales do Mucuri e Jequitinhonha,
Minas Gerais e Bahia, nos dominios da Mata Atlantica e em
Areas consideradas prioritarias ou que necessitam de maiores
investigao6es cientificas apontadas por Conservation Inter-
national et al. (2000). Este inventario gerou novos dados de
ocorrencia para o muriqui-do-norte (Brachyteles hypoxan-
thus), que sao aqui apresentados e discutidos.

No present estudo e em trabalho previo (Melo, 2004),
foram enfatizadas amostragens nos fragments florestais lo-
calizados nas areas consideradas prioritarias de ndmero 213
(Vit6ria da Conquista/Jordania, BA), 217 (Salto da Divisa,
MG) e 221 (remanescentes da regiao de Te6filo Otoni,
MG) do mapa "Avaliaqao e Ak6es Prioritarias para a Con-
servacao da Biodiversidade da Mata Atlantica e Campos Su-
linos" (Conservation International et al., 2000). 0 ndmero
de fragments amostrados depended de uma analise de via-
bilidade feita com toda a equipe do projeto, envolvendo os
demais grupos temAticos, ap6s analise de imagens de satdlite
(escala 1:250.000). Aqui, sao relatados apenas os resulta-
dos dos locals onde a presenga do muriqui pode ser con-
firmada, quais sejam, a Reserva Biol6gica da Mata Escura,

municipios de Jequitinhonha e Almenara (16o20'29.0" Sul
e 41o00'42.0" Oeste) e a regiao da Fazenda Duas Barras,
em Santa Maria do Salto (16o24'23.5" Sul e 4003'15.4"
Oeste; Figura 1) (mais detalhes abaixo).

Andrade (2004) caracterizou a regiao da Mata Escura com
clima que corresponde ao tipo Cwa de Kbppen, mesot&rmi-
co com verses quentes e estacao seca no inverno. Nos pontos
mais altos (1.100 m de altitude), o clima e mais ameno,
tipo Cwb de Kbppen, com temperatures mais brandas (An-
drade, 2000a). JA a Fazenda Alto Cariri, nos municipios de
Salto da Divisa e Santa Maria do Salto, e caracterizada pelo
clima tropical quente 6mido com tries meses secos. A Area
encontra-se pr6xima a isoeta com 1.250 mm de precipita-
o6es e as chuvas distribuem-se entire a primavera e o verao,
sendo os meses de precipitacao maxima entire dezembro e
fevereiro (Nimer, 1989; Andrade, 2000b).

0 levantamento de primatas que ocorrem nas localidades
de estudo foi feito atraves de tries m&todos principals: 1)
amostragens em transectos lineares ("censos") conduzidas
durante o perfodo diurno e noturno para o registro de ma-
mfferos de medio e grande porte; 2) uso de playback para a
confirmaqao de presenga de primatas; e 3) entrevistas com
moradores locals, antigos cacadores e pesquisadores que jA
atuaram na Area.

Figura 1. A localizacao de ambas as dreas de estudo, vale Jequitinhonha, Minas Gerais e Bahia.


4Q U FW 40 OD-W

49 T-W


Neotropical Primates 12(3), December 2004

Os "censos" (line-transect sampling, Buckland et al. 2001)
foram feitos em trilhas ja existentes, bem como em bordas
de mata e estradas de terra pelo interior dos fragments em
dois horarios principals, no infcio damanha (06:00-10:00h)
e no infcio da noite (18:00-22:00 h). Um minimo de tries
censos diurnos e tries noturnos foram realizados/fragmento/
estacao. 0 registro de mamfferos em geral foi feito atraves
de observao6es visuais diretas ou atraves de vocalizao6es,
pegadas, fezes, carcacas, pelos, espinhos e demais vestigios
encontrados. 0 ritmo da caminhada foi mantido o mais
pr6ximo possivel de 0,5-1,0 km/h. 0 infcio e o t&rmino dos
censos foram anotados, assim como a distancia percorrida.
As distIncias foram medidas atraves de aparelho de GPS.
Nos censos noturnos utilizamos uma lanterna Maglite re-
carregavel de 12 watts e Petzl modelo Duo. Todos os regis-
tros foram anotados em caderneta de campo previamente
preparada, com espagos para hora, local, tipo de floresta,
tipo de registro, e ndmero de individuos. Adicionalmente,
Melo (2004), alim de entrevistas com moradores locals, uti-
lizou insistentemente da t&cnica de playback, na tentative de
otimizar os encontros (Melo e Mendes, 2000).

O present estudo foi realizado em quatro Areas dos 36 frag-
mentos florestais visitados por Melo (2004). Dos 167 km de
censo e mais de 180 km de caminhadas percorridos, obtive-
mos apenas dois registros de muriqui-do-norte (Brachyteles
hypoxanthus) num total de 74 avistamentos de primatas.

Melo et al. (2002) confirmaram a ocorrencia de muriqui
para a regiao da Mata Escura, hoje a Reserva Biol6gica Fe-
deral da Mata Escura, em Jequitinhonha, totalizando 14
individuos distribufdos em dois grupos sociais distintos.
Mais recentemente, Melo et al. (in prep.) puderam confir-
mar o ndmero total de individuos de um dos grupos vistos
em 1999, que chegou a 25 animals, considerando adults,
jovens e infants.

Apesar de terem sido raros os relates references ? ocorrencia
de muriquis no vale, esse registro no municipio de Jequiti-
nhonha (REBIO Mata Escura) e de extrema importIncia,
pois as populao6es consideradas relictuais de distribuiiao
mais ao norte no estado de Minas Gerais eram as popula-
o6es do Parque Estadual do Rio Doce, em Marlikria e da
Fazenda C6rrego de Areia, no municipio de Peganha (Mit-
termeier et al., 1987; 1989; Strier e Fonseca, 1996-1997;
Hirsch et al., 2002).

Em Agosto de 2004, durante censo realizado na Fazenda
Duas Barras, municipios mineiro de Santa Maria do Salto e
baiano de Guaratinga, pudemos confirmar mais uma popu-
lacao de muriquis para o vale. A confirmaqao dessa popula-
gao traz um duplo significado para a conservacao dessa es-
p$cie tao ameagada, cuja populacao mundial nao ultrapassa
a cifra de 1.000 individuos de vida livre (Mittermeier et al.
2005), ja que reforga a existencia de mais uma localidade.
Somada as demais, aumenta as esperancas de consolidadao
da luta pela preservacao de seu habitat e de sua sobreviven-
cia em long prazo. No caso especifico dessa regiao, ambos
os governor do estado da Bahia e de Minas Gerais podem

somar esforgos na tentative de implantar uma Unidade de
Conservacao (UC) de protegao integral na Area.

A presenga do muriqui nessa regiao, alem de ser uma con-
firmaqao obtida ap6s cinco anos de busca por Melo (2004),
merece destaque, pois pode ser considerada a primeira po-
pulacao de muriquis constatada para o estado da Bahia
ap6s as investiga6oes pioneiras feitas por Aguirre (1971).
Alkm desse avistamento feito, computamos diversos relatos
fieis da ocorrencia da especie para toda a regiao chamada
de Alto Cariri. 0 grupo identificado apresenta pelo menos
sete individuos e se encontrava em um grande fragmento
de mata que abrange ambos os estados. A fazenda Duas
Barras possui aproximadamente 1.200 ha de floresta pri-
mAria e e continue com os demais fragments florestais si-
tuados em todo o maciso da serra do Alto Cariri, que pode
somar mais de 18 mil ha de mata em bom estado de con-
servacao. Como metade dessa Area se encontra em territ6-
rio baiano, temos uma situaqao singular onde encontramos
esp&cies com suas distribui6oes alocadas em ambos os es-
tados. Essa condigao interessante sugere que podem existir
outros grupos de muriquis ao long deste maciso florestal,
que inclui extensos trechos totalmente inseridos em terri-
t6rio baiano, pois tivemos muitos relatos da ocorrencia de
grupos com mais de 30 individuos, tanto ao norte quanto
nessa regiao central onde observamos o primeiro grupo
de muriquis.

Em outro projeto de pesquisa, Melo et al. (in prep.) realiza-
ram uma visit a Reserva Particular do Patrim6nio Natural
(RPPN) Estacao Veracruz, em Porto Seguro (BA), de acordo
com informa6oes de moradores locals que teriam vistos mu-
riquis na Area. Nao obtivemos, porem, nenhum indfcio que
pudesse confirmar sua ocorrencia durante a visit feita, o
que vem ratificar as informa6oes de seu desaparecimento da
Estacao Veracruz e vizinhancas hA pelo menos 40 anos, mo-
tivado aparentemente pela forte pressao de caca na regiao.
Aguirre (1971) aponta quatro localidades distintas, mas re-
lativamente pr6ximas a atual RPPN Veracruz, como Areas
cujas popula6oes de muriquis foram extintas no passado,
incluindo o Parque Nacional (PARNA) Monte Pascoal e a
Fazenda Pontal, em Itamaraju.

Um 6nico relato obtido durante o trabalho levanta suspei-
tas recentes de um muriqui abatido por indios no Parque
Nacional Monte Pascoal, hA cerca de cinco anos. Embora
remota, nao se pode descartar totalmente a possibilidade de
que uma populacao remanescente de muriquis ainda ocorra
nas Areas deste PARNA.

0 muriqui (Brachyteles spp.) nao teve seu status de conserva-
dao, em Minas Gerais, avaliado como criticamente ameaga-
do, pois durante a anAlise da lista mineira da fauna ameagada
de extingao, esse tAxon era considerado um genero mono-
tfpico. A sua taxonomia foi revisada alguns anos depois por
Rylands et al. (2000), onde houve separaqao em duas esp&-
cies (muriqui-do-sul, B. arachnoides e muriqui-do-norte, B.
hypoxanthus). Tanto a lista national quanto a lista da IUCN
trazem essa 6ltima especie como criticamente ameacada de

Neotropical Primates 12(3), December 2004

extingao no Brasil e no mundo, respectivamente (Brasil,
IBAMA, 1992; IUCN, 2004).

Comparando cornm o trabalho feito por Rylands etal. (1988)
na regiao do medio Jequitinhonha, nenhuma das duas po-
pulagoes citadas nesta regiao por estes autores pode ser con-
firmada. Na Fazenda Ramaiana, em Joafma (MG) (ponto
32 em Rylands et al., 1988), os moradores mais antigos in-
formaram ter ocorrido muriqui na drea, mas em tempos re-
motos. Muitas especies da avifauna identificadas no vale do
Jequitinhonha foram coletadas nessa localidade, sendo que
registros hist6ricos importantes, como do macuco (Tinamus
solitarius), demonstram que a regiao sofreu intense pressao
de caca ou mesmo teve sua area de floresta tao reduzida
que a especie nao subsiste mais (G. T. Mattos, com. pess.).
Da mesma forma, B. hypoxanthus pode ter chegado a um
nimero reduzido e o seu exterminio por cacadores pode ter
selado seu destino, bem como na maioria dos fragments de
mata primAria relativamente grandes que foram detectados,
mas que nao deixam quaisquer indfcios da presenca recent
da especie. A segunda localidade, conhecida como Fazenda
Nossa Senhora das Gragas (ponto 30; Rylands et al., 1988),
tem possibilidades remotas ou sequer alguma possibilidade
de abrigar populagoes de muriqui.

Relatos confiiveis da presenca de muriquis foram obtidos na
Fazenda Alto Cariri (16o18'53.8" Sul e 39o59'42.4" Oeste),
situada mais ao norte da Fazenda Duas Barras, pordm no
municipio de Salto da Divisa (MG). A presenca da especie
nesta localidade deve ser muito provavel, jA que essa Area e
continue com a mata da Faz. Duas Barras onde os muriquis
foram vistos.

HA outro fragmento florestal situado na Fazenda Avenida
(15o47'44.0" Sul e 40o32'55.0" Oeste) divisa de municd-
pios de Bandeira e Jordania, em Minas Gerais, com Ma-
carani e Maiquinique, na Bahia, uma Area que tambem
possui extens6es consideriveis de mata e que se encontra
em excelente estado de conservacao. Todas as visits feitas
por Melo (2004), este estudo e por Melo etal. (in prep.) nao
puderam confirmar a presenca de muriqui nessa Area, apesar
de terms obtido entrevistas relatando a sua ocorrencia. En-
tretanto, consideramos que a localizacao de novos grupos
de B. hypoxanthus nao deve ser descartada e que estudos
futures sejam priorizados na Area.

Uma terceira Area visitada e que nos levou a conhece-la
foi exatamente a indicaqao, pelo seu proprietArio, de que a
Fazenda Limoeiro (16o02'57.0" Sul e 40o51'02.0" Oeste),
em Almenara (MG), abrigava grupos de muriquis. Isso se
baseou em relato feito por um morador local que disse ter
abatido um indivfduo adulto de muriqui. Pordm, em ne-
nhuma das duas visits realizadas na Area foi possfvel confir-
mar a ocorrencia desse taxon (14 a 18 de novembro de 2003
e 20 a 24 de abril de 2004). A Area de mata e grande (mais
de 9.000 ha), existem grotas extensas bastante fngremes e de
dificil acesso. Mesmo assim, ha uma constant retirada de
madeira, o que nos leva a crer que a especie, se de fato ainda
subiste no local, estA no limiar da extincao.

A redescoberta de grupos de animals de especies de mami-
feros de grande porte, como o muriqui-do-norte, reforga a
necessidade urgente de se conhecer melhor as Areas visitadas
com relacao aos demais elements da biota regional e exige
uma agao imediata dos governor estadual e federal na con-
servacao dessa biodiversidade, jA que a caca predat6ria, o
fogo constant e a retirada de madeira sao ao6es de deterio-
raqao permanent nessas Areas.

Agradecimentos: Especiais agradecimentos aos financiadores
desse trabalho, como o Instituto Estadual de Florestas, a
Margot Marsh Biodiversity Foundation, a SEMAD-MG,
& Wildlife Service, o CNPq atraves das bolsas de estudos
concedidas e a Conservagao Internacional do Brasil. Aos
demais parceiros de equipe do PROBIO (Romulo Ribon,
Renato Feio, Luciana Nascimento, Alexandre Salino) e do
IEF (Denize E Nogueira e Priscila M. Andrade). Ao prof.
Anthony Rylands pela credibilidade e incentive durante
o doutorado de E R. Melo. Agradecemos aos alunos da
UEMG/FAFILE de Carangola por ter nos ajudado durante
a campanha da Faz. Duas Barras, Erica dos Reis Rodes, Mar-
cello Silva Nery e Silvia Lucilia Fonseca de Souza, atraves do
projeto financiado pela Fundacao Biodiversitas e CEPAN
(028M/012004). A Maria Elisa Castellanos-SolA pela idea-
lizaqao do trabalho original e ao tecnico do IEF local, Gio-
vani Alves de Moura, pelo auxflio de todas as horas e apoio
is misses empreendidas. Ambos tiveram papeis cruciais no
desenvolvimento das pesquisas cientificas que vem ocorren-
do no vale Jequitinhonha desde 1999.

Fabiano R. Melo1'2, Adriano G. Chiarello3, Michel B.
Faria1, Pedro A. Oliveira3, Rafael L. A. Freitas3, Fernan-
do S. Lima'14 e Daniel S. Ferraz', 'Ciencias Biol6gicas,
FAFILE/UEMG, Campus de Carangola, Praga dos Es-
tudantes 23, Santa Emflia, Carangola 36800-000, Minas
Gerais, e-mail: ; 2Centro de Estu-
dos Ecol6gicos e Educaq~o Ambiental, Rua Capara6 122,
Centro, Carangola 36800-000, Minas Gerais; 3Programa de
P6s-graduacao em Zoologia dos Vertebrados, PUC-Minas,
Av. Dom Jose Gaspar 500, Coraqao Eucarfstico, Belo Ho-
rizonte 30535-610, Minas Gerais; 4Instituto de Pesquisas
Ecol6gicas IPL, Caixa Postal 47, Nazard Paulista 12960-
000, Sao Paulo, Brasil.


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Primates 10(3): 115-119.


Jerimiah Luckett, Elizabeth Danforth
Kim Linsenbardt, Jill Pruetz


We conducted a study at the privately owned El Zota Bio-
logical Field Station in Costa Rica to assess the effects of
forest management techniques on primate ecology and be-
havior. While many conservation-oriented studies note the

Neotropical Primates 12(3), December 2004

need for "corridors" to promote dispersal between isolated
habitat fragments, few studies provide quantitative infor-
mation on their use by primates. From July to August 2002,
we studied the three primate species that occur at the El
Zota Biological Field Station in Costa Rica- Cebus capuci-
nus, Ateles ..-rr .. Alouatta palliata-to compare their use
of planted versus naturally forested areas. We collected ap-
proximately 25 hours of data to quantify the general activi-
ties exhibited by primates in these types of habitat.


Field site
The El Zota Biological Field Station (1057.6'N,
8375.9'W) in northeastern Costa Rica (see Pruetz and
LaDuke, 2001) is affiliated with the non-profit organiza-
tion DANTA: Association for Conservation of the Tropics,
which, in addition to organizing university-level field cours-
es, seeks to preserve biodiversity in this region of the coun-
try. The station is owned by the Hiner Ramirez family of
Guapiles and is also affiliated with the Fundaci6n Neotropi-
ca in Costa Rica. El Zota has offered field courses in topics
such as Primate Behavior and Conservation, and Tropical
Herpetology since 2001 and provides an opportunity for
students to compare intact forest ecosystems with disturbed
and fragmentary habitats. Covering some 1000 ha, much of
the site is primary humid and swamp forest (700 ha), but
it also includes areas of planted native and non-native trees
(-270 ha), as well as pastures formerly used for cattle (30
ha) but now planted with food crops or used for grazing by
horses. Emergent trees typical of primary lowland forest in
Costa Rica, such as Dipteryxpanamensis, are widely logged
but common at El Zota. Some of the notable fauna there
include jaguar (Panthera onca), tapir (Tapirus bairdii), great
green macaw (Ara ambigua), king vulture (Sarcoramphus
papa), brown caiman (Caiman crocodylus), the fer-de-lance
or terciopelo (Bothrops asper), and strawberry poison-dart
frog (Dendrobates pumilio).

The large-scale plantings at El Zota are part of a reforesta-
tion plan in areas previously deforested and used as cattle
pasture. Both native and exotic species were originally
planted for sustainable tree harvesting, especially Gmelina
arborea (Meliaceae), Carapa guianensis (Meliaceae), and
Hyeronima alchorneoides (Euphorbiaceae). Gmelina arbo-
rea is a fast-growing tree indigenous to Asia (Arbonnier,
2000) and is the primary species planted at El Zota. It
is planted at a density of approximately 82 trees per ha
(Linsenbardt, unpubl. data) and produces fruits edible to
humans, which are possibly also eaten by the non-human
primates. Other tree plantings appear to be similarly
spaced and probably occur at similar densities. Hyeronima
is a large forest tree native to Costa Rica, characterized
by small berry-like drupaceous fruits that are edible for
humans (Martin et al., 1987). Carapa guianensis, also
native, is a medium-sized to emergent tree especially prev-
alent in rich soils and swamps (Gentry, 1993). The age of
the plots at this site for all three species is approximately
8 to 10 years.

Study subjects
The three species of primates at El Zota overlap in range as
well as diet. The mantled howling monkey (Alouatta pal-
liata) is folivorous and frugivorous, preferring young leaves
to mature foliage (Sussman, 2000). Flowers and a small
amount of insect material also constitute a significant por-
tion of its diet on a seasonal basis (Burton, 1995). Fruit
accounts for approximately 80% of the diet of the black-
handed spider monkey (Ateles .., r .., i. while leaves make
up the remaining 20% (Burton, 1995). It has large day and
home ranges due to its widely and patchily distributed food
resources (Sussman, 2000). This means that this species in
particular needs large contiguous forest remnants or smaller
protected islands connected by large travel corridors. The
white-faced capuchin (Cebus capucinus) has the most varied
diet. It is frugivorous and insectivorous. Fruit comprises
65% of its diet, while insects (20%) and leaves (15%) also
figure prominently (Burton, 1995). Capuchins have also
been known to eat eggs, small vertebrates, buds, and berries
(Burton, 1995).

At El Zota, these three primate species use their mutual
forest habitat in structurally different ways due in part to
their variable dietary requirements. The plantation trees
could potentially be used alternately or in combination as
food resources, travel corridors, or resting areas between
separated parts of the secondary forest. To date, while use
of the planted plots has been described anecdotally, no
quantification has been made as to the nature of that use
or possible species-specific patterns that may exist. This
study is one of the first to describe patterns of forest usage
exhibited by these three primate species in relation to the
plantation stands.

Capuchin groups at El Zota are typical of those reported
elsewhere (Sussman, 2000), having a multi-male, multi-
female social system. Two groups of capuchins, each of
more than 10 individuals, were identified in the main 50-ha
study area. The spider monkeys showed the typical fission-
fusion social system characteristic of this species (Sussman,
2000), and the largest party was observed to contain 18 in-
dividuals. Mantled howling monkeys are the most common
species at El Zota. Five multi-male, multi-female groups,
each including between four to seven adults in addition
to lone individuals, were the focus of behavioral observa-
tions at El Zota. Pruetz and LaDuke (2001) estimated that
primates at El Zota occur at densities of approximately 35
howling monkeys/km2, 30 capuchins/km2, and 28 spider

Observers (KL, JL, JP) followed a systematic observation
schedule in which focal group follows were attempted
throughout the day to control for temporal bias in primate
activity. Over 100 hours in the field yielded approximately
25 hours of systematic data, although contact with primate
groups accounted for over 50 hours. A system of trails (3
km) that followed natural ecological contours, such as
streambeds, was used in addition to more randomly-ori-
ented transects (1 km) and roads passing through planta-

Neotropical Primates 12(3), December 2004

tions (2 km). Scan-sampling of all group members was used
to record general activity patterns, such as rest, travel and
feeding, as well as habitat (gallery forest, secondary forest,
plantation) and precise location using trail markers placed
every 10 meters.


The three species used areas of planted trees for movement/
travel and resting, with the proportion of time devoted to
moving within plantations greater than for other activi-
ties observed in this habitat (see Fig. 1). Howling monkeys
spent relatively more time resting in plantation areas (26%
of time) than spider monkeys and capuchins (17% and
16%, respectively). Primates were never observed to feed on
the non-native Gmelina. Although spider monkeys were not
seen to feed in Hyeronima trees in the plantations during sys-
tematic observations, they were observed doing so at other
times, and all three species fed in Hyeronima trees wherever
they occurred in natural forest at El Zota. Like Ateles, Cebus
rested and moved within planted areas, but were never seen
to forage there. Alouatta fed on Hyeronima in planted areas,
and moved through these areas, but most of the feeding
that we observed there was during a single feeding bout on
a lone Ficus left standing in a Gmelina grove. One group of
approximately 10 howlers frequented this particular tree.


Various studies have demonstrated that primates are able to
adapt to corridor habitats. Lion-tailed macaques (Macacac
silenus), for example, readily adapt to tea gardens in the
Western Ghats of India, where native rainforest fragments
are separated by tea and coffee plantings as well as eucalyp-
tus trees, and where non-native plants outnumber native
plants (Singh et al., 2001). The macaques fed more often
on non-native plants than native ones, and changes ob-
served with the introduction of non-native species to the
macaques' habitat included their tendency to forage on the
ground for insects and fallen fruits. Birth and survivorship
rates at this site increased, possibly in part due to increased
food sources in the corridors (Singh etal., 2001). This study
demonstrated that corridors can be implemented in degrad-
ed natural areas to help the animals adapt to a changing

Other studies have demonstrated that plantation corri-
dors provide new food sources for primates. Brown lemurs
(Lemurfulvus) near Perinet, Madagascar, have been record-
ed using old mixed eucalyptus and pine plantations as a new
food source (flowers of both pine and eucalyptus) as well as
for travel, resting and sleeping, although population densi-
ties in these areas were significantly lower than in natural
forest, and new eucalyptus plantations were used very little
(Ganzhorn, 1985, 1987). Nonetheless, Ganzhorn (1987)
concluded that non-native plantation corridors were effec-
tive in connecting natural forest areas and extending the
natural habitat of primates, as well as in creating a buffer
zone between the natural forest and outside disturbances.

Similarly, Zanne et al. (2001) concluded that introduced
pine plantations may attract more indigenous mammals,
including primates, than agricultural or fallow land in
Uganda. They likened these abandoned plantations to wild-
life refuges in an area where the landscape is fragmented. In
this case, such habitat could be interpreted as augmenting
the associated refuges. Finally, Bicca-Marques and Calega-
ro-Marques (1994) demonstrated that wild black howlers
(Alouatta caraya) in southern Brazil relied on introduced
plant species during periods of native fruit scarcity. Clearly,
although introduced species are viewed as problematic by
most wildlife conservationists, non-native or non-indige-
nous species may prove to be beneficial short-term solutions

CaMs cawnus



NrFAted 1842 67.4
ue 0 e58 3oa





Feld IF.4 b.1

S157 ,33

Figure 1. The use of planted versus non-planted (natural) areas
by three species of primates at El Zota Biological Station. The
graphs show the percentage of time dedicated to feeding, resting
and moving in these areas. The numbers in the tables show the
proportion of time each species spent feeding, resting and moving
in the two different habitats.

Neotropical Primates 12(3), December 2004

for the long-term conservation of primate species in some
contexts, especially in isolated forest fragments.

The propagation of non-native trees might be considered in
certain cases because fast-growing species could provide travel
corridors for primates or access to native food trees, as was
observed with primates at El Zota. A narrow corridor of non-
native, rapidly growing tree species could be supplemented
with slower-growing native species. These plantations could
also act as buffers and wildlife refuges in fragmented habitats
and would appear to be preferable to fallow or agricultural
land (Zanne et al., 2001). At El Zota, trees were originally
planted to provide timber for making paper. Research into
the ecology of introduced species before implementing such
a project is crucial. Although Gmelina trees at El Zota pro-
vide travel pathways for all three primates, the extensive
root network of this introduced species prohibits significant
undergrowth where it is thickly planted, stifling succession
by native species. For this reason, future management plans
at El Zota include intensive clearing of areas after Gmelina
harvest and the propagation of native tree species, such as
Hyeronima (J. Ramirez, pers. comm.). Using native trees and
allowing succession to occur would provide areas conducive
to supporting wildlife, rather than merely providing corri-
dors for wildlife movement (Lindenmayer and Nix, 1993).
Although all the plantations were originally part of a sus-
tainable wood-harvesting scheme, the primates' use of these
areas for travel has caused the management to change its
harvesting plans. Tentative plans are to harvest certain trees
while sparing stands that serve as pathways for primates. By
cooperating with biologists, the owner and managers of this
private reserve allow for a land-use regime that is compatible
with the conservation of biodiversity.

Acknowledgments: Thanks to the Hiner Ramirez family and
DANTA for contributing to the conservation of Costa Rican
biodiversity; the village of El Zota, the Primate Behavior
and Ecology class of 2002, Andy Brei, Heather Davis, Tom
LaDuke, Maribel Mora, Melissa Remis, Pedro Robles, Jes-
sica Westin, Teresa and William and family, and the Costa
Rican government for striving to sustain such diverse flora
and fauna.

Jerimiah Luckett, Department of Biology, Franklin Col-
lege, Franklin, Indiana 46131, USA, Elizabeth Danforth,
Department of Anthropology, Iowa State University, Ames,
Iowa 50011, USA, Kim Linsenbardt, Department of Sociol-
ogy and Anthropology, Purdue University, West Lafayette,
Indiana 47907, USA, and Jill Pruetz, Department of An-
thropology, Iowa State University, Ames, Iowa 50011, USA.


Arbonnier, M. 2000. Arbes, Arbustes et Lianes des Zones
Seches d'Afrique de l'Ouest. Centre de Coop&ration
International en Recherche Agronomique pour le D&-
veloppement (CIRAD) Museum National d'Histoire
Naturelle (MNHN) The World Conservation Union
(IUCN), Paris.

Bicca-Marques, J. C. and Calegaro-Marques, C. 1994.
Exotic plant species can serve as staple food sources for
wild howler populations. Folia Primatol. 63: 209-211.
Burton, Francis. 1995. Multimedia Guide to Non-Human
Primates. Prentice Hall.
Ganzhorn, J. U. 1985. Utilization of eucalyptus and pine
plantations by brown lemurs in the eastern rainforest of
Madagascar. Primate Conserv. (6): 34-35.
Ganzhorn, J. U. 1987. A possible role of plantations for
primate conservation in Madagascar. Am. J. Primatol. 12:
Gentry, A. H. 1993. Woody Plants of Northwest South
America. The University of Chicago Press, Chicago.
Lindenmayer, D. B. and Nix, H. A. 1993. Ecological
principles for the design of wildlife corridors. Conserv.
Biol. 7: 627-630.
Martin, E W., Campbell, C. W. and Ruberte, R. M.
1987. Perennial Edible Fruits of the Tropics: An Inventory.
Agriculture Handbook No. 642, U. S. Department of
Agriculture, Washington, DC.
Pruetz, J. D. and LaDuke, T. C. 2001. New field site:
Preliminary census of primates at El Zota Biological Field
Station, Costa Rica. Neotrop. Primates 9: 22-23.
Singh, M., Kumara, H. N., Kumar, M. A. and Sharma, A.
K. 2001. Behavioural responses of lion-tailed macaques
(Macaca silenus) to a changing habitat in a tropical
rain forest fragment in the Western Ghats, India. Folia
Primatol. 72: 278-291.
Sussman, R. W. 2000. Primate Ecology and Social Structure.
Volume2: New WorldMonkeys. Pearson Custom Publishing,
Needham Heights.
Zanne, A. E., Keith, B., Chapman, C. A. and Chapman, L. J.
2001. Protecting terrestrial mammal communities: Potential
role of pine plantations. Afr J. Ecol. 39: 399-401.


Bernardo Urbani

Previously, I reviewed chronicles that reported on or illus-
trated Neotropical primates in the XVt and XVI' centuries
(Urbani, 1999). Recently, I found two new documents that
are important for understanding how New World monkeys
were initially represented in Europe and Asia.

The first is an Ottoman map of 1513 made by the Turkish
Admiral Piri Re'is (1470-1554), a navigator and polyglot
who spoke Greek, Arabic, Italian, Spanish, and Portuguese.
This work, known as the Piri Re'is Carte de L'Atlantique
(90 x 65 cm), is housed at the Topkapi Sarayi Museum in
Istanbul, Turkey (La Ronciere et al., 1984: plate 28). The
polychrome map was lost until 1929 and was part of a larger
planisphere. Monkeys were illustrated but not mentioned
in the text (Afetinan, 1954; McIntosh, 2000) (Fig. 1). In
addition to Portuguese and Arab sources, Piri Re'is may
have drawn from a chart by Christopher Columbus, appar-
ently found in a Spanish ship captured by the Turks in the

Neotropical Primates 12(3), December 2004

Figure 1. Two monkeys of the New World in the Piri Re'is' Carte
de L'Atlantique 1513. One is to the right of a cynocephalus (on the
left of the map) and the other to the right of an acephalus (on the
right of the map) (La Ronciere etal., 1984: plate 28).

Mediterranean Sea around 1501. In fact, Piri Re'is' map
may reflect the earlier Columbus map of 1498 (La Ronciere
et al., 1984: 218), which coincidentally is the year that Co-
lumbus, in his travels, first reported on monkeys in America
(Urbani, 1999). In the highly detailed map of Piri Re'is,
baboon-like monkeys in the New World were drawn for the
first time (Fig. 1). It is possible to infer that these illustra-
tions were made with African primate referents, as were the
reports by other travelers in the New World such as Ameri-
go Vespucci (who referred to Neotropical primates as ba-
boons and macaques; Urbani, 1999) and Arabic chroniclers
(Kruk, 1995). On the other hand, Piri Re'is might have
obtained another original source on New World monkeys
directly from the Europeans. Two primates are represented
and associated with mythical animals, one "dancing" with a
cynocephalus (dog-head) and another with a fruit in its hand
together with an acephalus (headless) (Fig. 1). These mon-
keys were illustrated as inhabiting the area that is currently
Colombia, Brazil, and Venezuela.

In February 1595, the English pirate captain Sir Robert
Dudley (1574-1649), voyaging in the West Indies, en-
tered the Gulf of Paria (Venezuela) from the southwest
at Serpent's Mouth, leaving it by the Dragon's Mouth in
order to arrive at the Isle of Trinidad. Of this island, he
said, "the country is fertile, and ful of fruits, strange beasts
and foules, where of munkeis(3), babions and parts were
in great abundance [sic]" (Dudley, 1899: 71). He also in-
dicated that the local name for primates in Trinidad was
"howa" (Dudley, 1899: 78). Of interest is that the editor,
G. F Warner, wrote a footnote citing Charles Kingsley
(1819-1875): "(3)His 'munkeys' were, of course, the little
Sapajous; his 'babions' no true Baboons, for America dis-
dains that degraded and dog-like form, but the great red
Howlers (Kingsley, At last, p.69)." In principle, it is the
first reference that we know of for monkeys from a Ca-
ribbean island, and specifically Trinidad. Considering the
two primates of this island (Phillips, 1998), the "munkeis"
are most likely Cebus j .. trinitatis, whereas the "ba-
bions" refer to Alouatta seniculus insulanus, both endemic

Acknowledgments: To the personnel of the UIUC libraries
for their cooperation, and Paul Garber and Anthony B.
Rylands for their suggestions. As always to Tania Urquiza-
Haas. B. U. is supported by a Fulbright-OAS Scholarship.
The author would appreciate comments and references for
future updates.

Bernardo Urbani, Department of Anthropology, Uni-
versity of Illinois, 109 Davenport Hall, 607 S. Mathews
Ave., Urbana, Illinois 61801, USA, e-mail: uiuc.edu>.


Afetinan, A. 1954. Life and Works of the Turkish Admiral:
Piri Reis. The Oldest Map of America Drawn by Piri Reis.
Tuirk Tarih Kurumo Basimevi, Ankara. Translated to
English by Leman Yolay.
Dudley, R. 1899. Robert Dudley's voyage to the West
Indies, 1594-1595, narrated by Himself. In: The voyage
of Robert Dudley, afterwards styled Earl of Warwick and
Leicester and Duke j. '.. ,. to the West Indies,
1594-1595, narrated by Capt. Wyatt, by Hi r and by
Abram Kendall, master, G. F Warner (ed.), pp.67-79. The
Hakluyt Society, London.
McIntosh, G. C. 2000. ThePiriReisMap of1513. University
of Georgia Press, Athens.
Kingsley, Ch. 1896. At last: A Christmas in the West Indies.
Macmillan and Co., Ltd., London.
Kruk, R. 1995. Traditional Islamic views of apes and
monkeys. In: Ape, Man, Apeman: Changing Views Since
1600, R. Corbey and B. Theunissen (eds.), pp.29-42.
Leiden University, Leiden.
Phillips, K. A. 1998. Tool use in wild capuchin monkeys
(Cebus j.. trinitatis). Am. J. Primatol. 46(3): 259-
La Ronciere, M., Mollat du Jourdin, M., Azard, M.-M.,
Raynaud-Nguyen, I. and Vannereau, M.-A. 1984. Les
Portulans, Cartes marines du XII' au XVII siecle. Office
du Libre S. A., Friburg (Switzerland).
Urbani, B. 1999. Nuevo mundo, nuevos monos: Sobre
primates neotropicales en los siglos XV y XVI. Neotrop.
Primates7(4): 121-125.


Adrian A. Barnett


The majority of primate genus names are derived from Latin
or Greek roots, typically referring to some aspect of their
biology. Among the pitheciines, for example, Chiropotes is
derived from the Greek "khefr" (hand) and Latin "potare"
(to drink). This is a reference to the bearded saki's habit,
originally reported by Humboldt (1811: see Hershkovitz,

Neotropical Primates 12(3), December 2004

Table 1. Meanings of the generic names of non-pitheciine
Neotropical primates. Fr. = French, Gr. = Greek, L. = Latin.
Name Derivation
Alouatta alouette, Fr. 'lark'
(i.e., harbinger of dawn)
a, Gr. 'not',
Aotus otus, Gr. 'of the ear'
(i.e., 'hidden-eared')
a, Gr. 'not'
Ateles teleios, Gr. 'complete'
(referring to thumbless hand)
brachy, Gr. 'short'
Brachyteles teleios, Gr. 'complete'
(referring to nearly-thumbless hand, i.e., short
Callicebus kalos, Gr. 'beautiful'
kdbos, Gr. 'a monkey'
Calithrix kalos, Gr. 'beautiful'
thrix, Gr. 'hair'
Cebuella kdbos, Gr. 'a monkey'
-ellus, L. diminutive suffix
Cebus kebos, Gr. 'a monkey'
lagos, Gr. 'a hare'
Lagothrix trikos, Gr. 'hair'
(refers to pelage woolliness)
leon, L. 'lion'
Leontopithecus to, L. diminutive
pithekos, Gr. 'ape'
Saimiri "a Brazilian Portuguese name for a small monkey"*
Sagunus sagouin, Fr. 'a squirrel monkey'*
a -inus, L. 'like'
*from Gotch (1979)

1985), of drinking by dipping a hand into a bromeliad or
water-filled tree hole and then licking the wet fur. The genus
Pithecia comes from the Greek names for "ape" ("pithekos":
see Table 1 for further examples). However, this direct deri-
vation is not the source for the third pitheciine genus, Ca-
cajao, a name with no classical roots.

Like the classical derivations of most generic names, common
English names for Neotropical primates generally note some
obvious feature of the animal that-as is common in folk
taxonomies-provides a simple description of the animal
(Brown, 1985; Morren, 1989; Cormier, 2000; Mourao et
al., 2002). This is seen with "howler," "spider," and "squir-
rel" monkeys, the common names of Alouatta, Ateles and
Saimiri, respectively. Uacari does not fit this pattern, for
its origins are independent of any European language. This
paper, then, seeks to answer the following questions: How
did the name Cacajao come into use when it has no classical
roots, what is the origin of" uacari," and what are the actual
meanings of these names? Likewise I discuss what this may
tell us about the inclusion of local names into a taxonomic
system based on the terminology of classical languages.

Uacaris are medium-sized Amazonian primates (3-5 kg)
with short tails and a dentition adapted for a diet of hard
fruits (Barnett and Brandon-Jones, 1997). Endemic to the
Amazon basin, there are seven recognized forms (Hershkov-
itz, 1987) in two species: the bald uacari, Cacajao calvus (five

subspecies), and the black-headed uacari, C. melanocephalus
(two subspecies). Sousa e Silva Jinior and Martins (1999)
recorded the existence of a sixth bald uacari, which might
or might not be a new subspecies. Unusual in appearance,
uacaris have been described as "one of the most grotesque
of all primates" (C. A. Hill, 1965, p.140), and a monkey of
"melancholy aspect... emaciated... bedraggled" (W. C. 0.
Hill, 1960, pp.236-237). Humboldt (1811, p.316; 1812,
p.359) provided the first description of a uacari and named
it Simia melanocephala (in keeping with the time's highly in-
clusive sense of genus [see Defler and Hernandez-Camacho,
2002]), recording the common name of "Le Cacajao." By
1823 the all-embracing category Simia was no longer em-
ployed, and Johann Baptist von Spix (1823, p.12) named
the animal he collected Brachyurus ouakary. This genus
stood until 1840, when Lesson recognized its preoccupa-
tion by Brachyurus Fisher 1813 (a genus of rodent, itself
later synonymized with Lemmus). Deprived of this quite
appropriate term (brachyurus means "short-tailed"), Lesson
proposed-though without explaining why-that the
genus be renamed Cacajao. Isidore Geoffroy Saint-Hilaire
(1847), apparently unaware of Lesson's change, continued
the use of Brachyurus when describing (as Brachyurus calvus)
what is now C. calvus calvus, and did so again when describ-
ing what is now C. c. rubicundus (I. Geoffroy Saint-Hilaire
and Deville, 1848).

The names cacajao and uacari are evidently derived from
native Amazonian languages: both Humboldt and Spix
specifically noted that the names they used for their speci-
mens were those given by the local people at each collec-
tion locality. These terms, then, originated from native lan-
guages that were once spoken within the geographic range
of Cacajao melanocephalus. This range covers a large area
of northwestern Amazonia (see Hershkovitz, 1987; Bar-
nett and Brandon-Jones, 1997) and overlaps with an area
of considerable linguistic diversity (see maps in Dixon and
Aikhenvald, 1999). Uacaris occur in large groups, spend
much of the year being highly visible in riverside forests,
are hunted (Barnett and Brandon-Jones, 1997), and fre-
quently appear in folk taxonomies (e.g., Defler, 2003). The
Yanomami name for C. m. melanocephalus, for instance, is
hisho-hoshimi (Boubli, 1999). Given that "hashimi' means
"bad, unpleasant, worthless" and "hisho" refers to the area
between the nose and upper lip (Gail Goodwin Gomez,
pers. comm.), a loose translation could be "ugly snout", a
phrase that would certainly be in-line with the slightly pe-
jorative nature of many other local names for members of
the genus. However, Gail Goodwin Gomez (pers. comm.)
has cautioned that while this is a grammatically possible
phrase, it is unknown whether it would be acceptable to
a native speaker. Indeed, in his dictionary of the Venezu-
elan dialect of Yanomami, Lizot (2004, p.10) says "ho6s6mi
Zool., mono chucuto; Cacajao melanocephalus (Cebidae).
Es poco frecuente en la region habitada por los Yanommi
centrales." Gomez points out that the s/sh alternation is
found elsewhere in the Yanomami languages, and that it is
"linguistically quite normal to find a reduplicatedd' form,
[such as] hosomi hosomi," or the variant transcribed by

Neotropical Primates 12(3), December 2004

Boubli as honsho-honshome (where "on" refers to the nasal-
ized "o" vowel). So, Boubli's term is a reduplicated variant
of the term identified by Lizot (2004) in his dictionary of
Yanomami. Thus, the name may not be pejorative after all,
but simply monomorphemic, which cautions against hasty
interpretations of felicitous word combinations under such

Hershkovitz (1987) established the type locality for Hum-
boldt's specimen as a Salesian mission on the Rio Casiqui-
are, and the indigenous inhabitants of the mission were
said to use cauiri for C. m. melanocephalus (Humboldt,
1811). The Spanish missionaries called it chacuto, mono feo
or mono rabon; the second literally means "ugly monkey"
and so echoes the rather pejorative Yanomami name. The
third term refers to its short tail, and parallels rabicd, used in
Brazilian Amazonia (da Cunha and Barnett, 1989) as does
macaco mal-acabado ("unfinished monkey") reported by
Hershkovitz (1987). "Short tail" is also the direct meaning
of several indigenous names for C. melanocephalus, includ-
ing tschitschi in the language of the Tariana, who occupy
the upper Rio Vaupes in Colombia (Alexandra Aikhen-
vald, pers. comm; Koch-Grunberg, 1911), and tchitchi of
Baniwa, a language spoken mainly on the Rio Igana and
its tributaries on the Brazilian/Colombian frontier and on
the upper Rio Guainfa, Venezuela (Robin Wright, pers.
comm.). Piconturo or pitiontouro is a regional name for the
golden-backed uacari, Cacajao melanocephalus ouakary and
is often heard among settler (caboclo) communities on the
upper Rio Negro and its tributaries, including the Uapes/
Vaupes and the Curicuriarf; it is also used in the town of
Sao Gabriel do Cachoeira (da Cunha and Barnett, 1989).
This name appears to be a Europeanized (Spanish or Portu-
guese colonizers) rendition of ..-.- the name for the
animal in Tucano (Ramirez, 1997; Alexandra Aikhenvald,
pers. comm.). Ramirez (1997) gives p(nasalized i)ko as aroot
for "tail" (p.145), and turu (p.198) as "short." These varied
names, however, are not often used outside Amazonia and
shed no light on the provenance of cacajao and uacari.

The Origins of Cacajao and Uacari

According to Humboldt, cacajao or cacahao is a "Marabi-
tanas" Amerindian name for this monkey. "Marabitanas,"
however, is not recognized as a linguistic entity today, nor
did it exist at the time of Humboldt's visit to northwest-
ern Amazonia (Loukotka, 1968; Tovar and Tovar, 1984;

Victor Golla, pers. comm.). More likely, this was the name
of a village that was mistaken for an ethnic identity (but
see below). In Humboldt's time the Rio Casiquare region
was probably peopled by speakers of Bard, once the most
widespread of Maipurean (or Arawak) languages, originally
spoken from the Rio Branco to the upper Orinoco (Alex-
andra Aikhenvald, pers. comm; Victor Golla, pers. comm.)
but now nearly extinct (Aikhenvald, 1995). In Bard, the
term kakdhau (stressed on the second syllable) has been re-
corded to stand for the uacari (Alexandra Aikhenvald, pers.
comm.). This name does not appear to "mean" anything
in the descriptive sense, following the general pattern of
North Amazonian languages, in which descriptive names
for animals are generally rare (Alexandra Aikhenvald, pers.
comm.). Auricchio and Grantsau (1995) believe cacajao is
onomatopoeic for the uacaris' high-pitched "kah-kah" con-
tact calls. This might have been the origin of the name in
Bard, especially since elsewhere in the range of Cacajao mela-
nocephalus the common name for the uacari is bicd, which
almost certainly derives from their plosive "bee-koh!" alarm
call (A. Barnett, pers. obs.). The native names of many pri-
mate species are often close mimics of their various calls (see
Table 2 for Southeast Asian examples).

By the time of von Humboldt's visit, the Marabitanas did
not exist as a people, apparently having been exterminat-
ed by intertribal warfare in the late 1760s (Robin Wright,
unpubl. ms.). The word "Marabitanas" as recorded by
Humboldt may have been a place name derived from the
people's name or ethnic group (ethnonym) (Alexandra
Aikhenvald, pers. comm.), or it may have come from the
name of a Bard leader, as a number of prominent individu-
als seem to have used it. Little is known about the Marabita-
nas (Robin Wright, unpublished ms.), although one docu-
ment (Miss6es Salesianas do Amazonas, 1933, p.25) reports
that they were "aliados dos Arihini" or "allies of the Ari-
hini," a subgroup of the Bard. (Contra Nimuendaji [1932],
they were a cultural rather than a linguistic subgroup: see
Aikhenvald [1995]). This reputed alliance implies that the
two groups, Bard and Marabitanas, were linked by trade or
by language (Wright, 1991; Ramirez, 1997).

While traveling in the region, Karl Martius (1863) record-
ed kakayau as the name used for C. melanocephalus in the
area of the Braso Casiqiuare/upper Rio Negro. However,
the word kakdhau does not fit the pronunciation patterns
of Bard. Alexandra Aikhenvald (pers. comm.) notes: "I am
quite confident that kakdhau in Bard is a loan. One reason is

Table 2. Examples of onomatopoeic local names for Asian primates. (Taxonomy follows Groves, 2001).
Local name and language Latin and English names Source
Wow-wow (Malay) Hylobates lar Pocock (1939)
Pio (Bhotia) Macaca assamensis pelops Pocock (1939), Prater (1965)
Kra (Malay) Macacafascicularis Wood (1885), Finn (1929), Payne etal. (1985)
Sahu (Lepcha) Semnopithecus schistaceus Pocock (1939)1, Brandon-Jones (1999)
Wanga (Marachi) Semnopithecus dussumieri Pocock (1939)2, Brandon-Jones (1999)
1As Semnopithecus entellus achilles. 2As Semnopithecus entellus achates.

Neotropical Primates 12(3), December 2004

that such long roots (three syllables) are atypical for the lan-
guage. The other reason is that the sound "h" in Bard is very
restricted. It is never found in the middle of a morpheme
(for example, a root)." The shape and sound of the word
also stand out as highly unusual in the language, especially
the glottal fricative h, which is rarely found in that place
in a word and in that juxtaposition to other sounds (see
Aikhenvald, 1995).

There are two alternatives for the origin of this word in Bard.
First, it may be a very recent loan; the source person for Ai-
khenvald's dictionary of Bard, the last fluent speaker of the
language, was old and used a number of Spanish loan words,
such as playa for "beach." So kakdhau may have entered his
vocabulary via regional Spanish speakers. Alternatively, it
may be a loan from much longer ago, reflecting the status of
Bar&-speaking people of the upper Rio Negro as compara-
tively recent arrivals in the Casiquiare/upper Rio Negro area
(Derbyshire and Pullman, 1998). When they first entered
the region, the Bard may have borrowed names from other
tribal groups for the fauna that were new to them, as is
often the case (see Pike, 1959; Hunn, 1997; Atran, 1990;
Brown, 1984; Berlin, 1992; Cotton, 1996; Minnis, 2000
for other examples). One source of loan words may well
have been the Marabitanas, and one of those loaned words
may well have referred to a short-tailed primate with a sin-
gular vocalization. Before European contact, the upper Rio
Negro probably had over a hundred distinct languages, an
estimated 70% of which are now extinct (Ramirez, 1997;
Aikhenvald and Dixon, 1999; Aikhenvald, pers. comm.).
Given this ongoing cultural attrition, what we present there
cannot be firmly proven. What appears clear, however, is
that the word is not descriptive; it is merely reflective-an
onomatopoeic derivative.

This word (pronounced wah-KAR-ee) is now the accepted
English common name for all monkeys in the genus Ca-
cajao. It seems we owe this word to Spix, who wrote of the
"ouakary" monkey in his Simiarum et Vespertilionum Brasil-
iensium species novae of 1823, noting it to be the local name
where he collected his type specimen. Latinized to Ouakaria,
this name was briefly used for the genus proper by Gray in
1849, after Lesson (1840) replaced it with Cacajao.

While Humboldt's collection locality is quite precise (San
Francisco Solano Mission, Rio Casiquiare, Venezuela), that
of Spix is not. "Habitat in sylvis fluminibus Solimbens et
Iga interjectis" (Spix, 1823, p.13), the only geographical
reference in the original description of the species, does not
provide a collection point. Therefore, although Spix ac-
knowledges that "uacary" is a local name ("'Tespece de singe,
A quelle le nome Ouakary est applied par les habitans" [Spix,
1823, p.13]), the linguistic group from which this name
originated cannot be determined. Spix's reference to the Rio
Igi is a mystery in that the black-headed uacari he illus-
trates is not known to occur there (restricted to left bank of
the Rio Japura). It may be merely a reference to show the
habitat type occupied (riparian forest), rather than an actual

locality. The forests of the Rio Iia (the Brazilian stretch of
the Rio Putumayo) are occupied, at least on the right bank,
by Cacajao calvus rubicundus (see Hershkovitz, 1987).

Acariis used for C. m. ouakaryin Lingua Geral, a trans-Am-
azonian trading language (Stradelli, 1929). Lingua Geral is
based on a creole version of Tupinamba, from the Tupf-
Guarani branch of the Tupf language family, from what is
now Maranhao and Pard (Jensen, 1999; Alexandra Aikhen-
vald, pers. comm.). However, despite the widespread use of
acariin Lingua Geral to refer to uacaris (e.g., Tatevin, 1910;
Stradelli, 1929), what the word actually means is unknown
(Victor Golla, pers. comm.). It may be monomorphemic
(i.e., like "cat," but unlike "green woodpecker," it does not
mean anything per se [Denny Moore, pers. comm.]).

Thus, it seems that members of the pitheciine genus Ca-
cajao owe both their common and scientific names to words
deeply rooted in unrelated Amazonian languages, attached
to specimens independently collected and named by two
different 199t-century explorers working in widely separated
areas of the Rio Negro basin.


So, we have an explanation for the provenance of the names
and some understanding of their meanings in the origi-
nal languages. But why were these strange, non-European
names retained? Despite the uacari's obvious and unusual
physical characters-such as their odd facial appearance
and a tail one-third their body length (unique among
Neotropical primates)-it would seem that no European
common name for uacaris has ever been widely used. Given
that the common name for Chiropotes, the bearded saki,
helps distinguish it from the genus Pithecia, then "brush-
tailed saki" or "bob-tailed saki" might be sensible alterna-
tives to uacari; yet old wildlife encyclopedias (e.g., Broderip,
1857; Wood, 1885; Vogt and Specht, 1888; Miles, 1897;
Boulenger, 1936) used no common name other than vari-
ants of the word uacari.

Common names will often describe a new taxon by com-
bining two familiar animals, often unrelated, which seem to
encompass elements of the new form-for example, shrew
opossums (Caenolestidae), otter shrews (Potamogalidae),
and kangaroo rats (Dipodomysspp.). But some animals resist
all efforts to be described by amalgamation, and so we have
common names such as aye-aye (Daubentonia madagas-
cariensis), binturong (Arctictis binturong), cacomistle (Bas-
saricusspp.), goral (Naemorhedusspp.), kangaroo (Macropus
spp.), kinkajou (Potus flavus), llama (Lama glama), okapi
(Okapia johnstoni), peccary (Tayassu pecarn), serow (Capri-
cornis spp.), and tamaraw (Bubalus mindorensis). Likewise,
a local name for Cacajao was adopted as the common name
for want of any suitable European term. Such borrowing of
words from existing native folk taxonomies in circumstanc-
es of zoological uncertainty must have been very common
in the 18th and 19th centuries when new mammal species
were being described in numbers never seen before or since.

Neotropical Primates 12(3), December 2004

(Baratay and Hardouin-Fugier [2002] note that only 10%
of the mammal species known in 1993 were recognized in
1800; by 1890 that figure had risen to 50%.)

In effect, the formal adoption of a native name acknowledg-
es that what has been named is so far outside the standard
frame of reference that the entity defines itself; the local name
emphasizes the exotic nature of the animal and becomes its
own definition. This process is nicely demonstrated by the
uncertainty over what to call the recently discovered Asian
bovine Pseudoryx nghetinhensis. After several unsatisfactory
(and less than euphonious) attempts-"Loatian Ox-Ante-
lope," "Vu Quang Ox"-it was a regional name, "Sao La,"
that was finally adopted (see Nowak, 1999; Macdonald,
2001). For the third genus of pitheciines we must conclude
that Europeans, unable to elaborate on a previous common
name, defaulted to the local version, implicitly accepting
the incomparability of these highly specialized primates.
Uacari and cacajao, above all, seem to be a subliminal codex
that conveys the meaning "strange".

Acknowledgments: I thank the following people for kindly
sharing their knowledge and time: Alexandra Aikhenvald,
La Trobe University, Australia (who was particularly help-
ful and generous with her time); Dan Everett, Manchester
University; Victor Golla, Humboldt State University, USA;
Gale Goodwin Gomez, Rhode Island College, USA; Al
and Cheryl Jensen, Summer School of Linguistics; Denny
Moore, University of Manchester, England; and Robin
Wright, State University of Campinas, Sao Paulo, Brazil. I
am also grateful to Marie Monaghan, Michael Palmer and
Ann Sylph of the Zoological Society of London Library,
Larry Currey and Laura Burkhart at the California Acad-
emy of Sciences General Library, and staff of the University
of California Berkeley Life Sciences Library. Ann MacLar-
non (University of Surrey Roehampton), Rebecca Shapley
(Akodon Ecological Consulting), and Alexandra Aikhen-
vald commented on early drafts of this paper. The editorial
comments of John M. Aguiar and Anthony Rylands consid-
erably improved the original manuscript.

Adrian A. Barnett, Centre for Research in Evolutionary An-
thropology, School of Life and Sport Sciences, Roehampton
University, West Hill, London SW15 3SN, UK and Dept.
of Anthropology, California Academy of Sciences, Golden
Gate Park, San Francisco, CA 94118, USA (Research As-
sociate). E-mail: .


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Neotropical Primates 12(3), December 2004

Professor Dr. Harald Sioli, founder of
Amazonian Limnology and a major in-
fluential campaigner for the conservation
of the region, died on Thursday, 14 Oc-
tober 2004, aged 94. He was the former
Director of the Department of Tropical
Ecology of the Max-Planck-Institute for
Limnology, Plan. Most of his career was
dedicated to research-medical, physi-
ological, and, primarily, limnological-in Brazil. In 1954 he
became head of the Department of Limnology of what was
then the newly founded Instituto Nacional de Pesquisas da
Amaz6nia (INPA) in Manaus. He returned to Germany in
1956 as Head of the Department and Director of the Hydro-
biological Institute in Plan, later the Max-Planck Institute for
Limnology (MPIL), which resulted in the long-term collabo-
ration with INPA that continues today.

We owe to Harald Sioli the classification of the Amazonian
rivers (black-, white-, and clearwater) and their inundation
systems (igapd and vdrzea), and the entire basis of our under-
standing of much of Amazonian ecology and limnology. With
his explanations (his warnings) that the deforestation of the
Amazon would lead to a major increase in atmospheric carbon,
it was Harald Sioli who, unwittingly, gave rise to the widely
cited myth of the Amazon as "the lungs of the planet."

Two issues of Amazoniana were dedicated to Prof. Sioli on
the occasion of his 90th birthday-Volume 16(3/4) and
Volume 17(1/2). In the first of these, Dr Wolfgang Junk
(2001) wrote an appraisal of Sioli's scientific work and pub-
lished his curriculum vitae, including a full listing of his
scientific publications. Pages 169-172 of Volume 18(1/2) of
Amazoniana provided a series of short memoirs and com-
ments from numerous world-renowned scientists, friends,
and colleagues, including such as Sir Ghillean Prance, Dr
Jtirgen Haffer, Dr Eneas Salati, Dr Herbert Schubart, Prof.
Ernesto Medina, Prof. Nigel Smith, Prof. Loki Schmidt, Dr
William Rodrigues, and Dr Jorge Arias. Wolfgang Junk,
Director of the Working Group for Tropical Ecology of the
MPIL, wrote "During his active working period and also
thereafter, Professor Sioli exerted a strong influence on the
development of tropical ecology at an international level and
created an everlasting monument-his pioneering scientific
work about Amazonia" (in litt., 18 October 2004).


Junk, W. J. 2001. Appraisal of the scientific work of
Harald Sioli. Amazoniana 16(3/4); 285-297. [Includes
a curriculum vitae (pp.287-290) and a list of scientific
publications (pp.290-297)].
Schaller, E 2004. Nachruf/Obituary. Prof. Dr. Harald Felix
Ludwig Sioli (1910-2004). Amazoniana 18(1/2): 163-


The Tijuca forest in the municipality of Rio de Janeiro has
a long history of disturbance and modification from human
activities. It harbored a large indigenous population until the
16th century when, following European colonization, forests
throughout the Atlantic coast of Brazil were intensively
logged for brazilwood (Caesalpinia echinata). Highly valued
for its red/purplish dye and its wood, brazilwood became
commercially extinct in the Atlantic forest, including Rio de
Janeiro. Changes in the landscape of Tijuca were, however,
more drastic still in the 18th century, with the destruction of
most of its forest for coffee plantations.

In the middle of the 19' century the rivers feeding the city
of Rio de Janeiro were drying up, and the Emperor Dom
Pedro II, recognizing the link with deforestation in the
upper reaches and headwaters of the rivers, ordered that
the forest be restored. Both exotic and native trees were
used, along with a very large number of slaves to plant
them (Dean, 1996). In the first decades of the 20th century,
numerous non-native animals were introduced, including
primates. The restored forest of Tijuca was made into a
national park of 3,466 ha in 1961, and today is one of the
largest urban national parks in the world. The forest is now
well-established, although it lacks many of the species that
originally occurred there (see Coimbra-Filho and Aldrighi,
1971; Coimbra-Filho, 2000).

The original primate community of these forests would
predictably have included the southern muriqui (Brachyteles
arachnoides), the howler monkey (Alouatta guariba), the
black-horned capuchin monkey (Cebus nigritus), the
buffy-tufted-ear marmoset (C. .-.' aurita) and, in
lower elevations, the golden lion tamarin (Leontopithecus
rosalia) (Aguirre, 1971; Coimbra-Filho and Aldrighi, 1971;
Coimbra-Filho et al., 1973; Rylands et al., 1993). Neither
the black-fronted titi ((. .... nigrifrons) nor the northern
masked titi ((C .... personatus) have been recorded, and
it is not known whether they were ever present there (see
Cunha, 2003).

None of these species can be found in the Tijuca forest today.
Introduced common marmosets ((. .-'. jacchus) from
northeast Brazil and what we believe are hybrid capuchin
monkeys (Cebus) are the most abundant primates (Cunha,
2005). Marmosets and capuchin monkeys are both highly
favored as pets, and over the centuries many of them have
undoubtedly been let free or escaped. The capuchins we
have seen are very variable in pelage coloration and in the
form of their tufts and are certainly not pure-breeding Cebus
nigritus, but a mixture of different species, likely C. nigritus,
C. libidinosus, and C. robustus (Jose de Sousa e Silva Jr., pers.
comm.). The Amazonian squirrel monkey (Saimiri), and

Neotropical Primates 12(3), December 2004

the black-tufted-ear marmoset ((. .-'. penicillata) from
Central Brazil are also reported to occur in Tijuca, but are
evidently scarce as they were not seen during my fieldwork
in 2004. A single capuchin monkey was observed together
with a group of ( .-'. jacchus.

The abundance of common marmosets and the hybrid
capuchins is, we believe, related to the abundance of cer-
tain food resources, some of them non-native such as the
jackfruit (Artocarpus lhern,:op/9'//,h), particularly appreciated
by the capuchin monkeys (Cunha, 2005). Both marmosets
and capuchin monkeys adapt well to urban conditions,
entering backyards, gardens, and houses around the park
to find food. Since 1970, the zoologist and conservation-
ist Adelmar E Coimbra-Filho has warned of the potential
negative effects of these overabundant, non-native monkeys
on other species, specifically on bird populations, nestlings,
and eggs preyed upon by both Cebus and C .-( jacchus
(Coimbra-Filho and Aldrighi, 1971). C. jacchus also present
a potential threat to human health, harboring a variant of
the rabies virus (Morais et al., 2000) that has killed at least
eight people in the northeastern state of Ceara (Favoretto
et al., 2001). It is also a host for the Chagas' disease para-
site (Trypanosoma cruzi) and with marmosets now spread-
ing throughout the state, a potential source of infection
for Leontopithecus rosalia, surviving now only in the north
(Morais Jr., 2005).

Acknowledgments: This study was part of the MSc. thesis of
the first author. Carlos E. Grelle, Rui Cerqueira, and Antho-
ny B. Rylands made helpful comments on an earlier version
of this manuscript. The Brazilian Higher Education Au-
thority (CAPES) and Inter-American Development Bank/
Program de Despoluigao da Bafa de Guanabara/Fundacao
Estadual de Engenharia do Meio Ambiente/Instituto Terra
de Preservagao Ambiental provided financial support.

Andr6 Almeida Cunha and Marcus Vinicius Vieira, Pro-
grama de P6s-Graduacao em Ecologia and Laborat6rio de
Vertebrados, Departamento de Ecologia, Universidade Fe-
deral do Rio de Janeiro, Caixa Postal 68020, Rio de Janeiro
21941-590, RJ, Brazil, e-mail: ,


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Parque Nacional da Tijuca, GB, Brasil. Brasil Florestal 4:
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Mata Atldntica Brasileira. Cia. das Letras, Sao Paulo.
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F. A. A. and Mattos, C. A. de. 2001. Rabies in marmosets
((. .-.' jacchus), Ceara, Brazil. Emerging Infectious
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de Primatologia, p.57. Pontificia Universidade Catolica do
Brasil, Porto Alegre, RS, 13-18 February, 2005.
Morais, N. B., Rodin, B. N., Chaves, H. H. M., Neto,
J. B. de and Silva, L. M. da. 2000. Rabies in tamarins
((. .-'. jacchus) in the state of Ceara, Brazil, a distinct
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A. 1993. Systematics, geographic distribution, and some
notes on the conservation status of the Callitrichidae.
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Ecology, A. B. Rylands (ed.), pp.95-120. Oxford Science
Publications, Oxford.


The Peruvian government published the Supreme Decree
No. 34-2004-AG in the official journal El Peruano on 22
September 2004. This decree approves the categorization of
threatened Peruvian wildlife and at the same time prohib-
its hunting, capturing, owning, transporting, and exporta-
tion for commercial purposes. An appendix to the decree
lists 301 vertebrate species in different IUCN categories.
The decree will hopefully represent an important tool to
implement conservation action for, and stimulate research
on, threatened Peruvian wildlife. Table 1 lists the primate
species included in this appendix.

El 22 setiembre 2004, el gobierno peruano public en el
diario official "El Peruano" el Decreto Supremo no. 34-
2004-AG, que aprueba la categorizaci6n de species amen-
azadas de fauna silvestre y al mismo moment prohibe su
caza, capture, tenencia, transport y exportaci6n con fines
comerciales. El decreto lleva un anexo, en la cual estan lis-
tadas 301 species de vertebrados en diferentes categories
de la UICN. Este decreto represent un instrument im-
portante para implementar acciones de conservaci6n y al
mismo moment estimular investigaciones cientificas en las

Neotropical Primates 12(3), December 2004 155
Table 1. Categorization of Peruvian primate species.
Cuadro 1. Categorizaci6n de species de primates del Peri.
CR Critically endangered/En peligro critic
No primates listed no incluye species de primates
EN Endangered/En peligro
Alouatta palliata Mantled howler monkey Mono coto de Tumbes
Aotus miconax Andean night monkey Mono nocturno, musmuqui
Ateles belzebuth White-bellied spider monkey Maquisapa vientre blanco, mono arana
Oreonaxflavicauda Yellow-tailed woolly monkey Mono choro de cola amarilla
Saguinus labiatus Red-bellied tamarin Pichico de barriga anaranjada
"_____________________________ _____Pichico pecho anaranjado
VU Vulnerable/Vulnerable
Ateles chamek Black spider monkey Maquisapa
Cacajao calvus Red uakari Huapo colorado, huapo rojo
Callicebus oenanthe Andean titi monkey Toc6n, tit
Caicebus torquatus llhanded titi mo nkey Toc6n de collar, toc6n negro
Ca _____eus toruatus _Yellow-handed titi monkey
Callimico goeldli Goeldi's monkey Pichico de Goeldi, supay pichico
Lagothrix cana Geoffroy's woolly monkey Mono choro
Lagothrix lagotricha Humboldt's woolly monkey Mono choro
NT Near Threatened/Casi amenazado
Alouatta seniculus Red howler monkey Mono coto
Lagothrixpoeppigii Poeppig's woolly monkey Mono choro

species amenazadas. Cuadro 1 present las species de pri-
mates incluidas en este anexo segdn las categorfas.

Eckhard W Heymann, Abteilung V, -ilr, r ..1...1. .~i, &
Soziobiologie, Deutsches Primatenzentrum, Kellnerweg 4, D-
37077 Gbttingen, Germany, e-mail: .

Reference / Referencia

El Peruano, Lima, 22 de setiembre de 2004, pp.276853-


In September 2004, Marcela Santamaria G6mez of the Wild-
life Research Group, Department of Anatomy, University of
Cambridge, UK, defended her doctoral thesis, supervised
by David J. Chivers and titled, "The effect of home range
reduction on the ecology of red howler monkeys in Central
Amazonia." It was supported by the Biological Dynamics
of Forest Fragments Project (Smithsonian Institution and
Institute Nacional de Pesquisas da Amazonia), the Funda-
gao 0 Boticario para a Protegao da Natureza, Apiros, Brit-
ish Federation for University Women (BFUW), Instituto de
Pesquisa Ambiental da Amazonia (IPAM), the University of
Cambridge, Selwyn College, the Department of Anatomy,
the American Society of Primatologists, and Idea Wild. The
following is a summary of the thesis.

The loss of effective seed dispersers in forest remnants has
been stated to disrupt dispersal services that alter the dynam-

ics of tropical forests. Howler monkeys are efficient seed
dispersers that prove to be exceptionally tolerant to habitat
fragmentation by surviving in very small forest fragments.
In this context, the effect of home range reduction caused
by habitat fragmentation on the feeding ecology of red
howler monkeys (Alouatta seniculus) and on their subse-
quent role as primary seed dispersers in Central Amazonia
is examined.

The study was carried out at the Biological Dynamics of
Forest Fragments Project (BDFFP) near Manaus, Brazil.
Two howler groups living in fragments of 2.5 ha and 12
ha (Groups 1 and 2, respectively) were habituated during
a four-month period. Systematic data on the diet, activity
budgets, and use of space were collected for each group on
a monthly basis between January 2002 and January 2003.
Comparisons on seed dispersal were also made from faecal
analysis between the two fragments (Co 2.5 ha and Co 12
ha) and continuous forest (Km41).

At the three study sites, a strong seasonality in plant-part
production was recorded within the howlers' home ranges,
dividing the year into three seasons: fruiting (January-
May), leafing (June-September) and flowering (October-
December), but low fruit availability was found within the
small fragment. Both groups were frugi-folivorous, with
drastic seasonal variations based on plant-part availability.
Group 1 responded to low fruit supply by eating more
flowers and by repeatedly crossing a secondary-vegetation
gap (50 m) to obtain fruit from an adjacent continuous
forest. Forest fragmentation drastically reduced the howlers'
home range from about 20 ha in Km41 to 7.2 ha and 2.5
ha in the fragments (63% and 88% reduction, respectively).

Neotropical Primates 12(3), December 2004

Home range was composed of 60% edge habitat for Group
1 and only 26% for Group 2. A more drastic reduction
in the home range of Group 1 resulted in a net decline in
food plants, which affected decisions of time allocation in
this group: howlers living in the small fragment rested more
and fed less throughout the study. Group 1 seemed to live
in a more demanding environment and was presumably at
its limits of flexibility, whereas Group 2 was well adjusted
to a less disturbed habitat.

Although the reduction of the howlers' home range de-
creased the number of seeds and plant species dispersed,
howlers were effective seed dispersers for most of the spe-
cies they consumed at the three study sites. In fragments,
up to 77% of small to large seeds of fruits exploited by
Groups 1 and 2 were dispersed by endozoochory; passing
through the howlers' digestive systems-lasting about 20
hours-had a positive or neutral effect on germination
success. Despite the fact that Group 1 deposited seeds at
shorter distances from fruit sources than Group 2, up to
93% of seeds were moved away from the parental crown
by both groups.

Although howlers are generally regarded as less effective
dispersers because they produce large faecal clumps with
seed aggregation, this is not always the case. Five main
aspects of deposition patterns emerged from this study,
which indicated that the view of categorizing a species as a
good or bad disperser is misguided.

1) Howlers defaecated mainly in synchrony (c. 80%), but
also separately (20%).
2) In both defaecation types, howlers dispersed seeds in
latrines and random sites in the forest, but more often in
the former (up to 66%).
3) Seed deposition was spatially clumped; this patchiness
was associated not only with sleeping areas, but also with
latrines located outside them.
4) In latrines, howlers deposited more seeds of more species
than in random sites, but at both fragments seed densities
were similar between defaecation sites.
5) The fate of seeds delivered in howlers' multispecies seed
depositions varied greatly according to seed species at all
sites and to seed density at Km41. Nonetheless, after nearly
one year 51% of the seedlings had survived in Km41, 50%
in Co 12 ha, and 23% in Co 2.5 ha.

In conclusion, howlers living in small and medium frag-
ments are contributing to the maintenance of regeneration
processes through seed dispersal; consequently, the species
is a key element to consider in management and restoration
programmes of fragmented landscapes.

Marcela Santamaria G6mez, Wildlife Research Group,
Department of Anatomy, Downing Street, Cambridge
CB2 3DY, UK, e-mail: .


G6mez, M. S. 2004. The effect of home range reduction on
the ecology of red howler monkeys in Central Amazonia.
PhD thesis, Wildlife Research Group, Department of
Anatomy, University of Cambridge, UK.


On 17 November 2004, at the opening of the third IUCN
World Conservation Congress, the 2004 IUCN Red List of
Threatened Species was launched alongside the Global Spe-
cies Assessment: a detailed analysis of the Red List data and
the first major assessment of the status of global biodiversity.
The 2004 RedList is now available on the Red List website,
, as well as on the mirror sites
and .

This year's version includes a new search feature. It is now
possible to search for species by their most recent assess-
ment date. If you are looking for a complete list of assess-
ments on the Red List (including a complete list within a
particular country or region), you should select "All" from
the "Red List Assessment Years" list box on the search page.
If you are looking for a list of species reassessed or added to
the list in a particular year, you should select that year from
the list box (e.g., for a list of species reassessed or added to
the list this year, select "2004" from the list box).

We have also added an Acknowledgements page this year to
recognize and thank all the contributors to the list. Without
the enthusiasm, generosity, patience, and hard work of all of
our volunteer contributors, the production of the Red List
would not be possible.

Caroline Pollock, Research Assistant, IUCN/SSC Red List
Programme, 219c Huntingdon Road, Cambridge CB3
ODL, UK, e-mail .


Laury Cullen Jr., Research Coordinator at Instituto de Pes-
quisas Ecol6gicas (IPL) based in Nazard Paulista, Sao Paulo,
is a recipient of The Rolex Awards for Enterprise, sponsored
by Rolex S.A. The award was announced in Paris on 29
September 2004. A key aspect of Cullen's receiving this
award was his project "Transforming Farmers into Con-
servationists to Preserve the Atlantic Forest and its Fauna."
Over the last nine years, Cullen has focused on protecting
the remaining forest fragments in the west of the state of
Sao Paulo, working with small farmers and landowners and
demonstrating techniques and systems in agroforestry that
promote the recovery of degraded soils as well as the preser-
vation and recovery of forest fragments and their fauna. He
is currently planning to increase the number and extent of
forest corridors in the region, while simultaneously helping

Neotropical Primates 12(3), December 2004

to promote the economic wellbeing of at least 400 farmers.
IPL was founded in 1992 specifically to protect the black
lion tamarin, Leontopithecus chrysopygus, one of the many
species that will benefit directly from the forest restoration
resulting from Cullen's project. Cullen is currently conduct-
ing research for his doctoral thesis at the Durrell Institute for
Conservation and Ecology (DICE) of the University of Kent,
UK. The deadline to register for "The Rolex Awards for En-
terprise 2006" is 31 May 2005. Websites: awards.com>, .


No dia 05 de outubro de 2004, foi langada oficialmente a
rede 'speciesLink' criada pelo Centro de Referencia em Infor-
maqao Ambiental (Cria), Estado do Sao Paulo, Diretor Pre-
sidente Vanderlei Perez Canhos. Integrada ao Sistema de In-
formaqao Ambiental do Programa Biota/FAPESP (SinBiota),
a rede, que devera permitir a integraqao dinamica de dados
sobre a biodiversidade paulista, comeqa alum das fronteiras
do Estado: a coleqao do Jardim Botanico do Rio de Janeiro j d
esta integrada ao sistema.

O sistema permit a integraqao de diferentes grupos taxono-
micos por meio de bancos de dados distribufdos e protocolos
de comunicaqco. Corn isso, sera possivel ligar, no future, as
coleq6es biol6gicas a outras redes de informaqao do pats e do
exterior, por meio de "software" livres. A nova estrutura envol-
ve registros de microrganismos, Acaros, insetos, repteis, mamf-
feros, peixes e tipos de madeira. A rede compartilharA infor-
mao6es de coleo6es das tres universidades paulistas e de nove
institutes de pesquisa, alum do Jardim Botanico fluminense. 0
speciesLink devera ser utilizado como embriao para o desenvol-
vimento de uma rede brasileira de coleo6es cientificas.

"Corn o objetivo de acomodar a biodiversidade tanto sob o
ponto de vista geogrAfico como taxon6mico, a expectativa e
que o sistema tenha 750 mil registros ate 2006", preve Canhos.
A iddia e que esses aplicativos possam ajudar na resolucao de
problems como proteqao de esp&cies ameagadas, mudancas
climAticas e planejamento de Areas de conservacao.

"Corn o avanco das ferramentas de analise, sfntese e visualiza-
cao dos dados, as coleo6es que ficarem de fora de uma plata-
forma como o speciesLink tenderao a ficar menos competitivas
e menos visiveis para a comunidade cientifica", disse Canhos.

O mecanismo fisico que viabiliza o novo sistema foi estru-
turado a partir de servidores que permitem a integraqao de
informao6es por meio da Rede ANSP (Academic Network
at Sao Paulo), a conexao de internet avangada do Estado de
Sao Paulo e tambem um program da FAPESP Mais infor-
mao6es: .

Fonte: Thiago Romero, Agencia de Noticias da Fundacao de
Amparo a Pesquisa do Estado de Sao Paulo (FAPESP), 06
de outubro de 2004. Sftio de internet: fapesp.br>.


Ten students and professors from Bolivia and Peru partici-
pated in a field course on primates held in the Department
of Pando, Bolivia, during August 2004. Leila Porter (Uni-
versity of Washington) gave lectures on primate behavior,
ecology, and conservation at the Universidad Amazonica
de Pando (UAP) in the city of Cobija, and students con-
ducted field projects at the UAP's newly established Ta-
huamanu Biological Field Station. Claudia Coca Mendez,
of the Center for the Investigation and Preservation of the
Amazon (CIPA), UAP, served as the course coordinator.
The goal of the course was to introduce students to con-
cepts concerning primate research and conservation, and to
provide participants with the opportunity to conduct re-
search on wild primate groups.

The participants were presented with one week of intensive
lectures in Cobija. These courses provided an overview of
primate taxonomy, behavioral ecology, the role of primates
in their ecosystems, concepts of primate conservation bi-
ology, and conservation management strategies. Following
these lectures, participants developed team research propos-
als that they presented to each other for peer review. After
due revision, students and professors traveled to the field
station to start their research projects. At the field station,
local guides helped the teams to complete four projects:
1) a census of primate species' densities at the site; 2) an
estimation of niche overlap between Saguinusfuscicollis and
S. labiatus, 3) an examination of the diet of S. labiatus; and
4) a study of social behavior in juvenile S. fuscicollis. In ad-
dition to their research projects, participants were given les-
sons on how to capture callitrichines using blinds and traps
and how to collect fruit phenology data. Following comple-
tion of field research, students returned to Cobija to analyze
their data, prepare a final report, and present their results to
other professors and students from the UAP.

The Field Station is home to 11 species of primates, and
a number of primate groups (of four species) have been
habituated for field studies. The station therefore serves as
an excellent location for teaching about primate behavior
and ecology. In addition to learning about concepts related
to primate behavior, ecology, and conservation, students
became familiar with methods for primate research and de-
veloped important contacts across Bolivia and Peru. Due to
the success of this first Pandino primate field course, more
courses are anticipated for the future.

Special thanks to the Primate Action Fund (Conservation
International Margot Marsh Biodiversity Foundation) for
financial support for the course.

Leila Porter, Department of Anthropology, University
of Washington, Seattle, WA 98195-3100, USA, e-mail:

Neotropical Primates 12(3), December 2004


The first studbook for the European zoo collection of Cal-
licebus cupreus was produced in 2003, with data current up
to 31 December 2002 (Webster, 2004). Darren Webster,
of Blackpool Zoo, UK, has produced a second studbook
covering another year of births, deaths, and movements
current through 31 December 2003. These studbooks were
produced due to the large number of individuals taken in
by European zoos from the California Regional Primate Re-
search Center, Davis, during the years 2000-2002.

The red titi has a wide range on the upper Amazon, report-
ed to be restricted to the east of the Rio Ucayali in Peru to
the south of the Rio Amazonas-Solim6es in Peru and Brazil,
extending east as far the Rio Purus in Brazil (Hershkovitz,
1990; Van Roosmalen etal., 2002). Although quite variable
in pelage patterns, its key distinguishing features include
the lack of a frontal white blaze across the forehead, red-
dish-brown cheeks and sideburns, reddish-brown forearms
and forelegs and an overall buff-brown agouti crown, back,
upper arms, thighs, and tail. The tail can be overall similar
to the back (dark brownish) or very much paler to varying
extents from the tip to its proximal portion. Andrew Kitch-
ener, Curator of Mammals and Birds of the National Muse-
ums of Scotland in Edinburgh, discusses this variability in
a short essay in the studbook, "Understanding morphologi-
cal variation in the red titi, ( .... cupreus" (pp.24-25)
and provides a protocol and recommendations for cosmetic
postmortems (pp.26-27), requesting that carcasses be sub-
mitted to him for registration and preservation.

In 2003, the programme for the red titi monkey was up-
graded from a European Studbook (ESB) to a full-blown
European Endangered Species Programme (EEP) by the
European Association of Zoos and Aquaria (EAZA). The
living population registered in the studbook on 31 Decem-
ber 2003 was 36 (18.17.1) in eight institutions. Six indi-
viduals (2.4.0) in two institutions were not imported from
Davis, are of unknown origin, and are not included in the
breeding population. Two institutions (the Edinburgh Zoo
and the Skansan Akvarium, Sweden) joined the breeding
programme. All collections in a breeding situation produced
young in 2002 and 2003. There were seven births (3.2.2)
in 2003 with five (3.1.1) surviving. Two adults died and a
third individual was transferred to a private breeder and is
no longer part of the breeding program. Overall growth for
the breeding population was, therefore, two.

The studbook also includes information on the inbreeding
coefficient, age pyramid and births, deaths and transfers,
and full historical listing. There are contact details for dip-
loid chromosome sampling and a list of ( .... karyo-
types (adapted from Hershkovitz, 1990). The eight institu-
tions collaborating in the EEP are Apenheul Nature Park,
The Netherlands (0.2.0); Basel Zoo, Switzerland (2.0.0);
Berlin, Germany (2.3.0); Blackpool Zoo, Great Britain

(5.4.0); Bristol Zoo, Great Britain (1.1.1); Edinburgh Zoo,
Great Britain (2.1.0); LaVallke des Singes, Romagne, France
(3.5.0); and Skansan Akvarium, Sweden (3.1.0).

Darren Webster, Blackpool Zoo, East Park Drive, Black-
pool FY3 8PP, England, UK, e-mail: @blackpoolzoo.org.uk>, and Andrew Kitchener, Curator
of Mammals and Birds, National Museums of Scotland,
Chambers Street, Edinburgh EH1 1JF, Scotland, UK.


Hershkovitz, P. 1990. Titis, New World monkeys of the genus
( .. iCebidae, Platyrrhini): A preliminary taxonomic
review. Fieldiana Zoology, New Series, (55): 1-109.
Van Roosmalen, M. G. M., Van Roosmalen, T. and
Mittermeier, R. A. 2002. A taxonomic description of the
titi monkeys, genus ( .... Thomas, 1903, with the
description of two new species, ( .... bernhardi and
( .... stephennashi, from Brazilian Amazonia. Neotrop.
Primates 10(Suppl.): 1-52.
Webster, D. (compiler). 2003. European t'udlbook, for Red
Titi Monkeys (Callicebus cupreus). No 1. Blackpool Zoo
Park, Blackpool, UK. 32pp. Data current to 31/12/2002.
Webster, D. 2004a. A European studbook for the red titi
((C .... cupreus). Neotrop. Primates 12(1): 38.
Webster, D. (compiler). 2004b. European Studlbook for Red
Titi Monkeys (Callicebus cupreus). No 2. Blackpool Zoo,
Blackpool, UK. 27pp. Data current to 31/12/2003.


On 4 November 2004, Sally Walker received the Heini Hedi-
ger Award from Ed McAlister, President of the World Associ-
ation of Zoos and Aquariums (WAZA). Sally Walker is Coor-
dinator for South Asia for the IUCN/SSC Primate Specialist
Group. The following paragraphs are from McAlister's com-
ments describing her life and conservation achievements.

Sally Walker was born in the United States of America.
She first travelled to India in the early 1970s as a scholar of
Sanskrit, the ancient and sacred language of the Hindus. A
heightened awareness of Asian cultures encouraged Sally to
become a student of Yoga. This deep cultural immersion also
awakened a profound interest in the concept of a "wholeness
of nature," increasingly expressed in a concern for the welfare
and conservation of the threatened wildlife in India. Sally's
first practical commitment to this concern was as a volunteer
at Mysore Zoo, where in 1981 she established the Friends of
Mysore Zoo Society-an educational, public relations, and
scientific support group, the first of its kind in Asia. So began
a life-long dedication to Indian, and subsequently, all Asian
Zoos and a firm commitment to develop, through education
and science, their role and mission in conservation.

Walker's early success resulted in an appointment to the Na-
tional Zoo Advisory Board of the Department of Environ-

Neotropical Primates 12(3), December 2004

ment, Government of India. Government officials encour-
aged Sally to set up the Zoo Outreach Organisation (ZOO)
in 1985. Although based in India, the organisation now ex-
tends throughout South Asia, including the nations of Ban-
gladesh, Bhutan, Nepal, Maldive Islands, Pakistan, Sri Lanka,
and Afghanistan. This is a region that contains a tremendous
amount of biodiversity but is bedevilled by problems such as
language differences, political differences, economic, cultural,
and social differences, not to mention ongoing conflict. We
are all aware of the recent devastation wrought in Afghani-
stan. The Zoo Outreach Organisation, with Sally's guidance
and support, has successfully tackled these problems.

Working both for zoo and field conservation, the zoo organi-
zation has become a model for successfully building networks
to facilitate or catalyse conservation action. It brings together
stakeholders, e.g., scientists, other specialists, and laypeople,
to form a consensus on a variety of conservation issues. Two
monthly publications, ZOOS' PRINT and ZOO ZEN,
founded and edited by Walker, carry the work of the orga-
nization far beyond South Asia. A number of other publica-
tions have emanated from this organization, one of the most
recent being the excellent "Mammals of Afghanistan."

Walker's other work has included lobbying initiatives, pro-
vision of educational material, and the development of
scientific training programs. In particular, as Convenor of
the IUCN/SSC Conservation Breeding Specialist Group
- India, she has been instrumental in establishing the now-
celebrated CBSG Regional Networks. Walker's early work
with zoos led to a six-year membership on the Central Zoo
Authority, an autonomous body of the Ministry of Environ-
ment, Government of India, and membership on Ministe-
rial committees rare and unusual honours indeed for those
born outside India.

An expert in seeking support for her many initiatives; Walker
has become an institution within the Zoo profession, and has
demonstrated an extraordinary capacity for achieving suc-
cessful outcomes, often from extremely limited resources.

Walker is a dedicated conservationist and has achieved a
great deal, not only in the region in which she had decided
to live and work but all around the world. She is an inspira-
tion to all of us, and I have great pleasure, on behalf of the
WAZA Council and you, its members, on handing over the
Heini Hediger Award to her today.

Ed McAlister, President, World Association of Zoos and
Aquariums, P.O. Box 23, CH-3097, Liebefeld-Bern,


We have initiated a project in a contact zone between two
species of Alouatta in Brazil. Contact zones in this genus

have been highlighted in the literature since the early
1970s. We would like to establish a database of research
projects focusing on scientific questions related to con-
tact zones, broadly defined including sympatric zones.
Although we are particularly interested in contact zones
in the Neotropics, other geographical regions are also of
interest (for example, the zones of contact among baboon
species and subspecies). These studies are of particular
importance to primatologists studying behavioral mecha-
nisms of reproductive isolation (vocalizations and others
such as reinforcement) as well as the preservation of pri-
mate biodiversity due to increasing fragmentation of habi-
tats and other anthropogenic perturbations.

We conducted a search of speciationn" on PrimateLit,
which yielded only 123 citations. Most of these refer-
ences pertained to Paleotropical species and only one
paper mentioned bioacoustic character displacement
(without any data). Studies such as the one we propose
have been conducted with insects, anurans, fish, and birds
(see, for example, Loftus-Hills and Littlejohn, 1992;
also see Johnstone, 1997), which could provide helpful
templates for primate research on these topics. The en-
vironmental (abiotic and biotic) changes experienced by
numerous taxa provide natural experiments for changes
in signaling systems, species integrity, and other impor-
tant topics for long-term projects involving multidisci-
plinary input (for example, identification of hybrids with

We solicit your interest on these and related topics, including
registry of any contact zones of which you may be aware
or where you are currently focusing your studies. Please
contact one or both of us at the addresses below (preferably
via e-mail).

Clara B. Jones, Department of Psychology, Fayetteville State
University, 1200 Murchison Road, Fayetteville, NC 28301,
USA, Tel: 910-672-1575, e-mail: ;
. Webpage: clara.jones.socialpsychology.org>. Theoretical Primatology
Project (TPP) Website: .

Jdlio C6sar Bicca-Marques, Faculdade de Biociencias/
PUCRS, Av. Ipiranga, 6681 Pd. 12A, Porto Alegre 90619-
900, Rio Grande do Sul, Brazil, Tel: +55 (51) 3320-3545,
ext. 4742, e-mail:.


Johnstone, R. A. 1997. The evolution of animal signals. In:
Behavioral Ecology: An Evolutionary Approach, J. R. Krebs
and N. B. Davies (eds.), pp.155-178. Blackwell Science,
Oxford, UK.
Loftus-Hills, J. J. and Littlejohn, M. J. 1992. Reinforcement
and reproductive character displacement in Gastrophryne
carolinensis and G. olivacea (Anura: Microphylidae): A
reexamination. Evolution 46: 896-906.

Neotropical Primates 12(3), December 2004


In honor of Dr. Charles Southwick's long-
standing commitment to conservation
education, we have developed the Charles
Southwick Conservation Education Com-
mitment Award. This award is dedicated
to recognizing individuals living in primate habitat coun-
tries who have made a significant contribution to formal
and informal conservation education in their countries. The
amount of the award is $750: $500 is given to the recipient,
and $250 will be given in the recipient's name to a project of
their choosing in their community.

The IPS Education Committee is pleased to announce the
following recipients of the 2004 Charles Southwick Conser-
vation Education Award:

Dr. Surendra Mal Mohnot, Emeritus Professor and Chair-
man, Primate Research Center, Jodhpur, India, has a long
and distinguished career in teaching, research, and public
conservation education in India. He is the founding direc-
tor of the School of Desert Sciences, a teaching and research
center devoted to the conservation of wildlife, environments,
and people in the Great Thar deserts of western India. He has
trained and mentored more than 40 pre- and post-doctoral
students and young investigators, many of whom are now
working for the conservation of India's natural resources.

Mr. Wilberforce Okeka founded the Kakamega Environ-
mental Education Program (KEEP), a program focused on
environmental education for school children living near
the Kakamega Forest in Kenya. The educational goal of the
KEEP project is to teach local people about the wonder and
beauty of the forest and the importance of conserving it.
By partnering with local Kenyan organizations, KEEP has
also become involved in facilitating lifestyle changes that re-
lieve pressure on the forest, including tree planting, income
generation from on-farm cultivation of forest products, and
fuel-efficient cooking technology.

The Lawrence Jacobsen Education Development Award sup-
ports the initiation and long-term support of primate conser-
vation education programs. This education award supports
field conservation programs, work with local communities
and/or schools, and programs that provide training in con-
servation education techniques. The Education Committee
is pleased to announce the following recipients of the 2004
Lawrence Jacobsen Education Development Award:

Randy Kyes was awarded $500 to support a conservation
education program in the elementary schools around Tang-
koko-Batuangus-Duasudara Nature Reserve, North Sulawe-
si, Indonesia.

Sandra Correa was awarded $500 to support an educational
exhibit for Alouatta seniculus as part of Fundaci6n Ecolom-
bia's program to reintroduce animals rescued from the pet
trade to a restored forested area in Colombia.

Please join us in congratulating these deserving individuals
as they develop conservation education programs aimed at
preserving primates in their region.

Anne Savage, IPS VP for Education, Disney's Animal King-
dom, P.O. Box 10,000, Lake Buena Vista, FL 32830, USA,
e-mail: .


The CD of the XVIII' Congress of the International Pri-
matological Society (IPS) held in Adelaide, South Austra-
lia in 2001 is now available. The price is A$35 plus $A3
postage and handling within Australia. Overseas costs will
be US$25 plus US$4.00 for postage and handling. If you
would like a copy, please contact Graeme Crook, e-mail:
, with the subject heading "IPS
Congress CD."

Graeme Crook, President, Australasian Primate Society,
PO Box 500, One Tree Hill, SA 5114, Australia.


SThe ASP Conservation Committee
would like to solicit grant proposals
for the ASP Conservation Small Grants
competition of 2006. These grants, up to $1500, are
specifically designed to help fund conservation research
or related projects, including conservation education.
ASP members working in habitat countries are especially
urged to apply or to help someone from a habitat country
submit a meaningful project that can be a portion of a
larger effort. Guidelines for the grant applications are
available on the ASP website, ,
and may also be obtained from Dr. Janette Wallis, Chair
of the ASP Conservation Committee, ABTI-American
University of Nigeria, e-mail net> or .

Please note that the Conservation Small Grants will be
awarded early again this year, in order to expedite the
delivery of grant money to the winners in time for the
summer months when many of these projects begin. The
deadline for grant proposals is 16 January 2006. Materials
may be submitted online at the ASP website beginning
in early December or sent as an e-mail attachment to the
Committee Chair at the address above. Successful grants
will be announced in late March, 2006.

Neotropical Primates 12(3), December 2004


A Primatologia no Brasil- 8, edited by Sergio L. Mendes
and Adriano G. Chiarello. 2004. IPEMA Instituto de
Pesquisas da MataAtlantica and SBPr- Sociedade Brasileira
de Primatologia, Vit6ria, Espirito Santo, Brazil. 340pp.
ISBN 8599058010. This volume presents 26 chapters
based on presentations at the 9' Brazilian Congress of
Primatology held in Santa Teresa, Espirito Santo, Brazil
in July 1999. Contents: Os prim6rdios da primatologia
no Brasil A. F. Coimbra-Filho, p.11; Feeding biology of
Neotropical primates-D. J. Chivers & M. Santamarfa, p.37;
Reproductive strategies of New World primates: Interbirth
intervals and reproductive rates K. B. Strier, p.53;
Evolutionary trends of neotropical primates according to
the AP68 and AP40 microsatellites -M. Ruiz-Garcia, M. I.
Castillo and D. Alvarez, p.65; Biogeography of Amazonian
primates S. F. Ferrari, p.101; Interao6es ecol6gicas entire
mico-ledo-dourado (Leontopithecus rosalia Linnaeus, 1766)
reintroduzido e mico-estrela (( .-' jacchus Linnaeus,
1758) introduzido em fragments de MataAtlantica, RJ -
A. G. Affonso, C. R. Ruiz-Miranda and B. Beck, p.123; 0
mico-ledo-dourado (Leontopithecus rosalia Linnaeus, 1766)
minimize os customs do comportamento de brincadeira?
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behavior and selectivity in mantled howler monkeys,
Alouatta palliata. Diss. Abst. Int. B64(11): 5396.
Xu, X. M. 2004. Functional organization of visual cortex in
bush babies and owl monkeys revealed by optical imaging
of intrinsic signals. Diss. Abst. Int. B65(3): 1185.


XXIIIrd Annual Conference of the Australasian Primate
Society, 12-13 March, 2005, South Australian Museum,
Adelaide, South Australia. Twenty-minute sessions will be
reserved for each paper. Abstracts should be received before
1 February 2005. Information: Graeme Crook, President,
Australasian Primate Society, PO Box 500, One Tree Hill,
SA 5114, Australia, e-mail: . For
further details visit htm>.

Primate Society of Great Britain 2005 Spring Meeting,
22-23 March, 2005, Chester College. For more information
contact: Paul Honess, PSGB Meeting Officer, Department
of Veterinary Services, University of Oxford, Parks Road,
Oxford OX1 3PT, UK, e-mail: or
visit the website at .

IX Simposio de Antropologia Fisica, 4-8 April, 2005,
Habana, Cuba. El Museo Antropol6gico "Montand" y la
Catedra de Antropologfa "Lufs Montand" de la Facultad
de Biologfa de la Universidad de La Habana, la Sociedad
Cubana de Antropologfa Biol6gica, la Sociedad de
Estudios Primatol6gicos Eopithecus de Mexico, convocan
al IX Simposio de Antropologfa Ffsica "Luis Montand",

el V Congreso Primates como Patrimonio Nacional, el II
Coloquio Primates a traves del Caribe y el II Coloquio de
Antropologfa "Manuel Rivero de la Calle", del 4 al 8 de abril
del 2005. Correspondencia: Dr. Armando Rangel Rivero,
Secretario, Museo Antropol6gico Montand, Calle 25 #455,
entire J e I. El Vedado, Facultad de Biologfa, Universidad
de La Habana, Ciudad de La Habana, Cuba, e-mail:
, website: wisc.edu/pin/IX_SIMPOSIO DE ANTROPOLOGMA

2005 Meeting of the Mexican Society of Primatologists,
4-7 May, 2005, Instituto de Ecologfa, Xalapa, Veracruz,
Mexico. For information: Juan Carlos Serio Silva, Presidente,
Asociaci6n Mexicana de Primatologfa AC, Departamento de
Biodiversidad y Ecologfa Animal, Instituto de Ecologfa AC,
km 2.5 antigua carretera a Coatepec, No. 351 congregaci6n
El Haya, CP 91070, Apartado Postal 63, Xalapa, Veracruz,
Mexico, Tel: +52 (228) 8 42 18 00 ext 4109 /4110 (Fax: ext
4111), e-mail: .

Fourth Annual Callitrichid Behavioral Husbandry and
Management Workshop, 21-22 May, 2005, Washington,
DC, USA. The Callitrichid Behavioral Husbandry
and Management Workshop will be presented by the
Cotton-top Tamarin SSP and hosted by the US National
Zoo in Washington, DC. For more information,
see the Workshop's website at: si.edu/ConservationAndScience/EndangeredSpecies/

19th Annual Meeting of the Society for Conservation
Biology, 15-19 July, 2005, Universidade de Brasflia, Brasflia,
Brazil. Theme: "Conservation Biology: Capacitation and
Practice in a Globalized World." The chair is Miguel Marini,
Zoology Department, Universidade de Brasflia. Contact:
SCB 2005 Local Organizing Committee, Departamento de
Zoologia, IB, Universidade de Brasflia, 70910-900 Brasflia,
DF, Brasil, telefax: +55 61 307-3366, e-mail: <2005@conbio.
org>, website: .

Association of Tropical Biology and Conservation 2005
Annual Meeting, 23-29 July, 2005, Uberlandia, Brazil. The
venue will be the Uberlandia Convention Center. For more
information write to the Chair of the Organizing Committee,
Kleber del-Claro, Laborat6rio de Ecologia Comportamental
e Interagoes, Universidade Federal de Uberlandia, Caixa
Postal 593, Uberlandia 38400-902, Minas Gerais, Brazil, e-
mail or .

IX International Mammalogical Congress, 31 July-5
August, 2005, Sapporo, Japan. Organizing Committee:
MAMMAL2005, c/o Field Science Center, Hokkaido
University, N11 W10, Sapporo 060-0811, Japan, e-mail:
, website: www.imc9.jp>.

1t Congress of the European Federation of Primatology, 9-
12 August, 2005, Gbttingen, Germany. The Congress will be

Neotropical Primates 12(3), December 2004

hosted by the German Society for Primatology (GfP) at the
German Primate Centre (DPZ), University of Gbttingen. It
will coincide with the 9th Congress of the German Society.
European students and researchers working on all aspects
of primatology are invited to attend. Registration from 1
November 2004 to 30 March 2005. For more information
contact Peter M. Kappeler, President EFP, German
Primate Center (DPZ), Abteiling Verhaltensforschung &
Okologie, Kellnerweg 4, D-37077 Gbttingen, Germany,
e-mail: , website: primatologie.de/EFP2005/index.htm>.

28'h Annual Meeting of the American Society of Primatolo-
gists, 17-20 August, 2005, Portland, Oregon. The meeting
will be held at the Benson Hotel and hosted by the Oregon
National Primate Research Center. Call for abstracts and the
meeting announcement will be sent electronically to all ASP
members in mid-December 2004. Deadline for proposals
for symposia, roundtables or workshops is 17 January, 2005.
Deadline for abstracts for contributed papers, symposia
speakers, workshops and roundtable discussions is 14 Febru-
ary, 2005. If a paper version of the meeting announcement
is preferred, please contact: Larry Williams, Program Co-
Chair, Tel: +1 251-460-6293, Fax: +1 251-460-6286, e-mail:
. For more information, please
contact: Dr. Kristine Coleman, chair of the local organizing
committee of the ONPRC at .

29th International Ethological Conference, 20-27 August,
2005, Budapest, Hungary. For more information, write to:
IEC2005, Department of Ethology, Ebtvbs University, 1117
Budapest, Hungary, or subscribe to the e-mail newsletter at

COHAB 2005: First International Conference on Health
and Biodiversity, 23-25 August, 2005, Galway, Ireland.
This important global event will provide an international
forum for scientists, professionals, policymakers, and
stakeholders to address the issues linking environmental
health, human health, biological diversity, and international
development. Themes to be discussed at the conference
include the Millennium Ecosystem Assessment and the
U.N. Millennium Development Goals; biodiversity,
genetic resources, natural products and drug discovery;
nature, culture, sociology and mental health; biodiversity
in nutrition, agriculture and food production; pathogen
pollution and the ecology of infectious disease; the use of
wild species as sentinels of environmental health; functional
biodiversity and ecosystem services; and ethnomedicine
and traditional knowledge. A special session of COHAB
2005 will discuss policy options for addressing the health
and development problems posed by biodiversity loss,
with relevance to public health and development planning
policies worldwide. COHAB 2005 is being organised by
an international consortium led by the United Nations
Development Programme, the United Nations Environment
Programme, the Global Environment Facility, the World
Conservation Union, the Centre for Health and the Global
Environment at Harvard Medical School, and other

international organizations. Full details of the conference
may be found at http://www.cohab2005.com. Enquiries
should be directed to Conor Kretsch, COHAB Director,
e-mail: .

Measuring Behavior 2005 5th International Conference
on Methods and Techniques in Behavioral Research, 30
August 2 September, 2005, Wageningen, The Nether-
lands. Measuring Behavior will offer an attractive mix of
presentations, demonstrations, discussions, meetings, and
much more (see gram/index.html> for details). Proceedings of the 2002
meeting are available at mb2002/index.html>. Deadline for proposals of Symposia
and SIGs: 1 December 2004. For more information, con-
tact: Prof. Dr. Louise E. M. Vet, Program Chair, Measuring
Behavior 2005, Conference Secretariat, P.O. Box 268, 6700
AG Wageningen, The Netherlands, Tel: +31-317-497677,
Fax: +31-317-424496, e-mail: ,
website: .

Sixth Meeting of the Asociaci6n Primatol6gica Espaniola,
27-30 September, 2005, Facultad de Psicologfa, Universidad
Complutense de Madrid, Madrid, Spain. Sponsored by the
Asociaci6n Primatol6gica Espafiola (A.P.E.), this Meeting
will focus on the themes of Child Ethology, Conservation,
Great Apes and Humans: Similarities and Differences, and
Tool Use. For more information please see the website at
or contact Dr. Fernando
Colmenares () or Dra. Maria
Victoria Hernandez-Lloreda ().

2005 Annual Meeting of the Conservation Breeding
Specialist Group, 29 September- 1 October, 2005, Syracuse,
New York, USA. Beginning with a late-afternoon ice-breaker
on Wednesday, the meeting will run through Saturday,
ending with an afternoon and dinner at the Rosamond
Gifford Zoo. Regional network meetings will take place on
Tuesday, 27 September, and a Steering Committee meeting
on Wednesday, 28 September. Accommodations are at the
Genesee Grande Hotel (http://www.geneseegrande.com),
which offers a variety of rooms and rates. The deadline for
registration is 1 August 2005; for more information, email
a request to <2005cbsg@cbsg.org> or visit their website at

New World Primate Workshop (A Focus on Cebids), 30
September 1 October, 2005, Cleveland, Ohio, USA. The
Cleveland Metroparks Zoo announces a workshop on New
World Primates that will focus on the captive care of Cebids
in U.S. institutions. Informal roundtable discussions will in-
clude the following topics: diet and health, social groups and
mixed species, enrichment and training behaviors, and pop-
ulation management. The workshop will begin at 10 a.m. on
Friday, 30 September, and end at 4 p.m. on Saturday, 1 Oc-
tober. Attendance is limited to 50 people and registrants will
be asked to complete a pre-meeting survey regarding their
experiences with Cebids. The workshop will be held on the
zoo grounds. Some meals will be provided and local lodging

Neotropical Primates 12(3), December 2004

suggestions can be provided. Registration fee = $25. For more
information and a registration form, contact Tad Schoffher at
216-635-3332 or .

8th World Wilderness Congress, 30 September 6 October,
2005, Anchorage, Alaska, USA. Over a thousand delegates
from dozens of nations will attend the Eighth WWC, with
additional events in Kamchatka and the Russian Far East.
Convening every three to four years, the theme of this year's
Congress is "Wilderness, Wildlands and People-A Partner-
ship for the Planet." This Congress will generate accurate, up-
to-date information on the benefits of wilderness and wild-
lands to both contemporary and traditional societies, and will
examine the best models for balancing wilderness and wild-
lands conservation with human needs. For more information,
see the Congress website at .

60th World Association of Zoos and Aquariums Annual
Conference, 2-6 October, 2005, New York, New York,
USA. The 60th WAZA Annual Conference will be hosted
by the Wildlife Conservation Society and held at the
Marriott Marquis hotel. The theme of the meeting will be
"Wildlife Conservation: A Global Imperative for Zoos and
Aquariums." Additional information will be made available
on the conference website at .

III Congresso Brasileiro de Mastozoologia, 12 a 16 de
outubro de 2005, realizado pela Sociedade Brasileira de
Mastozoologia (SBMz) e a Universidade Federal do Espfrito
Santo (UFES), no SESC Praia Formosa em Aracruz, Espfrito
Santo. 0 event reunira pesquisadores, profissionais e
estudantes com o objetivo de apresentar, analisar e discutir
trabalhos cientfficos, descobertas e tendencias no estudo
dos mamfferos. 0 tema dessa edicao e "Diversidade e
Conservacao de Mamfferos," que sera abordado sob diversos
aspects durante o event, que contara com a participated
de especialistas ligados a instituio6es de ensino e pesquisa
nacionais e estrangeiras, bem como outros profissionais que
atuam em 6rgaos governamentais, na iniciativa privada e
em organizao6es nao-governamentais. Somente serao aceitas
inscrio6es pela internet. Podera ser realizada a inscrihao
online do congress ate o dia 31 de maio, e o envio dos
resumes podem ser feitos ate o dia 30 de junho de 2005.
Mais informao6es: .

Counting Critters: Estimating Animal Abundance
and Distance Sampling, 17-21 October 2005, Disney's
Animal Kingdom, Orlando, Florida, USA. This five-day
workshop will introduce participants to the most impor-
tant methods of estimating animal abundance in a rigor-
ous but accessible way. In the first half of the workshop,
we cover plot sampling, distance sampling, mark-recapture
and removal methods. We explain the common key statis-
tical concepts underlying the methods, use custom-written
simulation software to understand how the methods work,
and discuss which method to use when. In the second
half, we focus on distance sampling in more detail. We
discuss practical issues such as use of the Distance soft-
ware, field methods, and survey design. The workshop is

aimed at anyone who needs to estimate wildlife density
or abundance and is taught by leading researchers from
the Centre for Research into Ecological and Environmen-
tal Modelling at the University of St Andrews, Scotland.
Registration for this workshop is now open. Since all of
our previous workshops in the USA have been oversub-
scribed, we encourage everyone interested to register as
soon as possible. For more details, please see ruwpa.st-and.ac.uk/counting.critters/> or contact Rhona
Rodger, Workshop Organizer, CREEM, University of St
Andrews, The Observatory, St Andrews, Scotland KY16
9LZ, tel: +44 1334 461842, fax: +44 1334 461800, e-

Primer Congreso Colombiano de Primatologia,
Asociaci6n Colombiana de Primatologfa, del 2 al 4
noviembre de 2005, Bogota, Colombia. El Primer
Congress Colombiano de Primatologfa tendra tres Areas
Tematicas para la presentaci6n de los trabajos: Biologia
y Ecologia studios en ciencias bAsicas que incluyen
morfologfa, taxonomfa, sistemAtica, gen&tica, biologfa
molecular, evoluci6n, biodiversidad, comportamiento
y ecologfa; Medicina studios en anatomfa, fisiologfa,
medicine, clinic, patologfa, epidemiologfa, nutrici6n, y
restricci6n de primates; y Conservacidn y Manejo (in situ /
ex situ) investigaci6n aplicada y gesti6n multidisciplinaria,
herramientas conceptuales y tecnicas dirigidas a la
conservaci6n, uso y aprovechamiento, trabajo comunitario,
comercio, mantenimiento en cautiverio, reproducci6n,
tecnicas de capture, manipulaci6n, registro y marcaje,
enriquecimiento ambiental, rehabilitaci6n, disposici6n
de primates decomisados, normatividad y legislaci6n. La
ponencia debe incluir informaci6n nueva, se pueden enviar
resdmenes de temas presentados en reuniones anteriores
pero su aporte al Congreso debe ser clave, general discusi6n
constructive o representar temas emergentes. Para mayor
informaci6n del Congreso, puede visitar la siguiente pAgina
web: , o en
el correo electr6nico .

Primate Society of Great Britain (PSGB), Winter
Meeting 2005, 9 December, 2005. Flett Theatre, The
Natural History Museum, London. The theme is "Primate
Evolution 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: . See website: psgb.org/Meetings/Winter2005.html>.

V Gdttinger Freilandtage "Primate Diversity Past,
Present and Future," 13-16 December, 2005. University
of Gbttingen and German Primate Center, Gbttingen,

Neotropical Primates 12(3), December 2004

Germany. OrganizedbyPeter M. Kappeler. Confirmedinvited
speakers: Diversity in the Past: Extinct primate communities
- John F1, I, (State University of New York, Stony Brook).
Diversity Today: Diversity of Malagasy primates -Anne Yoder
(Yale University); Diversity of American primates -Anthony
B. Rylands (Conservation International); Diversity of Asian
primates Jatna Supriatna (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 Diversityfor Tomorrow: Diversity and conservation
hotspots Russell A. Mittermeier (Conservation
International); Extinction biology- Carlos Peres (University
of East Anglia); Conservation genetics George Amato
(Wildlife Conservation Society); Conservation genetics
- Michael Bruford (Cardiff University); Reintroductions
- Carel P. van Schaik (University of Zirich). Comparative
Perspectives: Speciation in birds Trevor Price (University of
Chicago); Bird taxonomy and conservation Robert Zink
(University of Minnesota). Contact: Prof. Dr. Peter M.
Kappeler, Deutsches Primatenzentrum (DPZ), Kellnerweg
4, D-37077 Gbttingen, Tel/Fax: +49-551-3851-284/291, e-
mail: , website: de/sociobiology/GFT2005/index.htm>.


Ecology in an Era of Globalization: Challenges and
Opportunities for Environmental Scientists in the Americas,
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. Abstracts
should address one of the meeting's three subthemes:
invasive species, human migration, and production. The
invasive species subtheme includes such topics as dispersal
of invasive plant and animal species, emerging diseases, and
resistance of local ecosystems to invasive species and disease.
The human migration subtheme includes the environmental
effects of international and local emigration and immigration
on recipient and source areas. Potential topics include
infrastructure development needs and impacts, effects 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 consider the need
to communicate with broad audiences. For more information,
or to submit an abstract, visit .
Deadline for abstract submissions: September 16, 2005.

75th Annual Meeting of the American Association for Phys-
ical Anthropology, 5-12 March 2006, Anchorage, Alaska,
USA. For program information, please contact the Program

Chair, Lyle W. Konigsberg, Department of Anthropol-
ogy, 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 Anthropology, Univer-
sity of Alaska Anchorage, Anchorage, AK 99508, USA, tel:
907-786-6839, fax: 907-786-6850, e-mail alaska.edu>. Website at annmeet>.

Primate Society of Great Britain (PSGB), Spring Meeting
2006, 27-28 March, 2006, University of Stirling, Stirling,
Scotland. The theme is "Primate Mentality and Wellbe-
ing." 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
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
Primatologists (ASP), 16-19 August, 2006, San Antonio,
Texas. Sponsored by Southwest National Primate Research
Center. Tentative deadline dates are: Notify program chair
of intent to offer a symposium or workshop by 5 December,
2005; Symposia and Workshop abstracts with confirmed
list of participants due to program chair by 9 January, 2006;
all final abstracts are due for symposia, oral, and poster
presenters by 6 February, 2006. See the ASP website for
updates and further information: meetings/index.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
available 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 Baldi, Chair of the Local Organising
Committee, at .


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, 1919 M St. NW, Suite 600, Washington,
DC 20036, Tel: 202 912-1000, Fax: 202 912-0772, 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, 1919 M St. NW, Suite 600,
Washington, DC 20036, USA.

Printed on New Leaf Reincarnation Matte 80# cover (100% recycled/50% post-
consumer waste, processed chlorine free) and New Leaf Reincarnation Matte 70#
text (50% recycled/30% post-consumer waste, elemental chlorine free). By using this
environmentally friendly paper, Conservation International saved the following resources:
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For more information about New Leaf Paper, go to ww newleafpaper. corn.

Neotropical Primates
A Journal and Newsletter of the IUCN/SSC Primate Specialist Group
Vol. 12(3), December 2004


Short Articles

The Mottled-face Tamarin, Saguinus inustus, in the Amana Sustainable Development Reserve, Amazonas, Brazil
Luciane L. de Souza, Helder L. Queiroz andJose Mdrcio Ayrest........................................................................................... 121
Preliminary Observations on the Mottled-face Tamarin (Saguinus inustus) on the Lower Rio CaquetA,
Colombian Amazonia
Erwin Palacios, Adriana Rodriguez and Claudia Castillo ................................................................ 123
Movements of Alouatta palliata Among Forest Fragments in Los Tuxtlas, Mexico
Salvador Mandujano, Luis Arturo Escobedo-Morales and Rodolfo Palacios-Silva............................................................................ 126
Aspects of the Behavioral and Endocrine Ontogeny of Six Moustached Tamarins, Saguinus mystax (Callitrichinae)
Maren Huck, Petra Lottker, Eckhard W Heymann and Michael Heistermann.............................................................................131
Social Structure of Alouatta guariba clamitans: A Group with a Dominant Female
Jodo M. D. Miranda, Itiber P. Bernardi, Rodrigo E Moro-Rios, Lucas M. Aguiar,
Gabriela Ludw ig and Fernando C. Passos ..................................... ............................................................................................... 135
Substrate Manipulation by Alouatta guariba clamitans in Solving a Locomotor Problem
Fldvia Koch and Julio Cesar Bicca-M arques.................... .......... ....................... ...................... ........ .....................138
Novos Registros de Muriqui-do-Norte (Brachyteles !\ ... ..- ,l no Vale do Rio Jequitinhonha, Minas Gerais e Bahia
Fabiano R. Melo, Adriano G. Chiarello, Michel B. Faria, Pedro A. Oliveira,
Rafael L. A. Freitas, Fernando S. Lima and Daniel S. Ferraz................................................................................................. 139
Planted Trees as Corridors for Primates at El Zota Biological Field Station, Costa Rica
Jerimiah Luckett, Elizabeth Danforth, Kim Linsenbardt and fill Pruetz .......................................143
Further Information on Neotropical Monkeys Reported in the XVI Century
B ernardo U rbani ................................................... ................................................................................................................ 146
The Meanings of Cacajao and Uacari: Folk Etymology in Neotropical Primate Taxonomy
A drian A Barnett......................................... .............................. ........................................... 147

In M em oriam : H arald Sioli ......................................... ................................................................................................. 153

N ew s .................................................... ............ ........................................ ..................... ..... 153

P rim ate Societies ........................................ .............................. ........................................... 160

R recent Publications....................................... ............................. ......................................... 161

M eetin g s .................................................................... ............................................................................................... ..... 165

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