Front Cover
 Back Matter
 Back Cover

Title: Neotropical primates
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Permanent Link: http://ufdc.ufl.edu/UF00098814/00053
 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: August 2007
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: VID00053
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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Table of Contents
    Front Cover
        Front Cover
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    Back Matter
        Back Matter 1
        Back Matter 2
    Back Cover
        Back Cover
Full Text

ISSN 1413-4703



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

Volume 14



Erwin Palacios
Liliana Cort6s-Ortiz
Jl1io C6sar Bicca-Marques
Eckhard Heymann
Jessica Lynch Alfaro
Liza Veiga
News and Book Reviews
Brenda Sol6rzano
Ernesto Rodriguez-Luna
PSG Chairman
Russell A. Mittermeier
PSG Deputy Chairman
Anthony B. Rylands



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

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

ISSN 1413-4703 Abbreviation: Neotrop. Primates

Erwin Palacios, Conservaci6n Internacional -Colombia
Liliana Cortes Ortiz, Museum of Zoology, University of Michigan, Ann Arbor, MI, USA
Jdlio Cisar Bicca-Marques, Pontificia Universidad Cardlica do Rio Grande do Sul, Porto Alegre, Brasil
Eckhard Heymann, Deutsches Primatenzentrum, Gattingen, Germany
Jessica Lynch Alfaro, Washington State University, Pullman, WA, USA
Liza Veiga, Museu Paraense Emilio Goeldi, Belem, Brazil

News and Books Reviews
Brenda Sol6rzano, Instituto de Neuroetologia, UniversidadVeracruzana, Xalapa, M6xico
Ernesto Rodriguez-Luna, Instituto de Neuroetologia, Universidad Veracruzana, Xalapa, Mexico

Founding Editors
Anthony B. Rylands, Center for Applied Biodiversity Science Conservation International, Arlington VA, USA
Ernesto Rodriguez-Luna, Instituto de Neuroetologia, 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
Carolyn M. Crockett, Regional Primate Research Center, University of Washington, Seattle, WA, USA
Stephen E Ferrari, Universidade Federal do Para, Belem, Brazil
Russell A. Mittermeier, Conservation International, Arlington, VA, USA
Marta D. Mudry, Universidad de Buenos Aires, Argentina
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
Coordinator, Special Section on Great Apes, Liz Williamson
Regional Coordinators Neotropics
Mesoamerica, Ernesto Rodriguez Luna
Andean Countries, Erwin Palacios and Eckhard W. Heymann
Brazil and the Guianas, M. Cecilia M. Kierulff
Regional Coordinators -Africa
West Africa, John E Oates
East Africa, David Mbora
Regional Coordinator- Madagascar
Jirg U. Ganzhorn
Regional Coordinators -Asia
China, Long Yongcheng
Southeast Asia, Jatna Supriatna and Christian Roos
South Asia, Sally Walker

PSG General Coordinator/Website Manager, John M. Aguiar

Layout: Kim Meek and Glenda Fabregas, Center for Applied Biodiversity Science, Conservation International, Arlington, VA, USA
Editorial Assistance: John M. Aguiar, Center for Applied Biodiversity Science, Conservation International, Arlington, VA, USA

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

Front cover: A bald uakari, Cacajao calvus calvus, from the Brazilian Amazon. Photo by Russell A. Mittermeier.

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

Neotropical Primates 14(2), August 2007 47


Jesus Martinez1 and Robert B. Wallace1' 2

1 ji-.-'. Conservation Society, Greater Madidi Landscape Conservation Program, Casilla 3-35181, San Miguel, La Paz,
2 i .. j.' Conservation Society, 185th Street and Southern Boulevard, Bronx, New York, 10460, USA, e-mail: <


We present information on the distribution of the endemic Bolivian primates Callicebus olallae and C. modestus based
on a series of site visits in the Beni Department. Callicebus olallae was registered at four localities and C. modestus at
11 localities, and in combination with negative data and localities for other Callicebus species in the region we defined
the distributional boundaries for both endemics. Mean group size was 2.7 individuals for C. olallae and 3 individuals
for C. modestus. The results highlight the restricted distribution of both species, with C. olallae having one of the highest
levels of primate endemism in the world and an "occurrence area" of 400 km2, and considering forest fragmentation
in the region, more realistically an "occupancy area" of 50 km2. Although C. modestus is found in slightly larger areas
(1,800 km2 and 450 km2 respectively) the distribution area is still extremely restricted. The restricted distributions together
with ongoing forest fragmentation due to cattle ranching, unregulated ecotourism activities, and principally the scheduled
improvement of the "Northern Corridor" road that passes through this region, means that both species should be consi-
dered highly endangered.

Key Words: Geographical range, group size, physical characteristics, Southwestern Beni Department, Bolivia


Presentamos informaci6n acerca de las areas de distribuci6n de los dos primates end6micos de Bolivia, Callicebus olallae y
C. modestus, mediante la visit de localidades en el departamento del Beni. Callicebus olallae fue registrado en 4 localida-
des y C. modestus en 11 localidades, y en combinaci6n con datos negatives y localidades para otras species de Callicebus
en la region definimos los limits de distribuci6n para las dos species end6micas. El promedio del tamafio de grupo para
C. olallae fue 2.7 individuos y para C. modestus 3 individuos. Los resultados resaltan la distribuci6n altamente restringida
que ambas species presentan, siendo C. olallae una de las species con mayor grado de endemismo de Sur America al tener
un "area de ocurrencia" de 400 km2 y, considerando el alto grado de fragmentaci6n del bosque, de forma m~s realista un
"irea de ocupaci6n" de 50 km2. Aunque C. modestus ocupa areas mayores (1,800 km2 & 450 km2 respectivamente) muestra
tambikn un elevado grado de endemismo. Esta situaci6n, junto con la fragmentaci6n del bosque debido a las actividades
de ganaderia, ademis de actividades no reguladas de ecoturismo y, principalmente el proyectado mejoramiento del tramo
vial del "Corredor Norte" que atraviesa la zona habitada por estos primates, conlleva a considerar a ambas species como
altamente amenazadas.

Palabras Clave: Rango geogrifico, tamafio de grupo, caracteristicas fisicas, suroeste del departamento Beni, Bolivia

Introduction of their continental distribution is scant, most dramati-
cally demonstrated by the recent discoveries of new species
The titi monkeys (genus Callicebus) are considered the (Hershkovitz, 1988, 1990; Van Roosmalen et al., 2002;
most diverse primate genus in the Neotropics, with Wallace et al., 2006). A good example of the poor distri-
29 recognized species (Van Roosmalen et al., 2002; Wal- butional knowledge regarding Callicebus is the case of the
lace et al., 2006). Clearly defined distributional patterns, two Bolivian endemics, Callicebus modestus and Callicebus
habitat preferences, food habits, and behavior of many of olallae (Kinzey, 1981; Hershkovitz, 1988, 1990; Ander-
these species are still unknown. Indeed current knowledge son, 1997; Rylands et al., 2000; Van Roosmalen et al.,

Neotropical Primates 14(2), August 2007

2002). Collected by the Olalla brothers and described by
Linnberg (1939) based on the examination of just three
collected individuals, the specimens have subsequently
been considered as full species in taxonomic revisions of
Callicebus (Hershkovitz, 1990; Kobayashi, 1995). Never-
theless, for almost seventy years no field data were obtained
about these apparently range-restricted Bolivian endemic
primate species.

In light of this situation, in 2002 the Wildlife Conserva-
tion Society began investigating the distribution of both
species (Felton et al., 2006). These preliminary studies
confirmed the presence of C. modestus and C. olallae, or
at least two distinct phenotypes (Felton et al., 2006), in
a zone near the suggested collection sites of the Olalla
brothers (Linnberg, 1939; Anderson, 1997), which com-
prises non-continuous forest patches interfacing with a
natural savannah system (Felton et al., 2006). However,
their overall distributional pattern and extent remained

A critical issue in the study of these species is their rather
similar morphological features (Linnberg, 1939; Hershko-
vitz, 1990) that together with variable lighting conditions
during field observations make accurate identification a
challenge. This situation calls into question the original
taxonomic distinctions made by Linnberg (1939) and
subsequently supported by others (Hershkovitz, 1990;
Kobayashi, 1995), especially when considering the small
sample size and the restricted and apparently overlapping
distribution area determined for both species. The original
specimens for both species were collected within 65 km of
each other (Felton et al., 2006), although it is important
to note that some sources have questioned the collection
locations for other primate species reported by the Olalla
brothers in other areas of South America (Aquino and En-
carnaci6n, 1996).

Establishing the taxonomic and conservation status of
these endemic, range-restricted species is of urgent con-
cern, particularly considering imminent plans to improve
the road system in the immediate region. This improve-
ment includes the paving of a major road and is likely to
have a deleterious effect on forest cover in the immediate
vicinity due to the colonization effect (Reid and Landivar,
1997; Forman and Deblinger, 2000; Trombulack and Fris-
sell, 2000). In this way, conservation research efforts for
C modestus and C olallae need to prioritize establishing
the range of these Bolivian endemics, estimating popula-
tion size, as well studying titi monkey genetics in order
to confirm their taxonomy in a phylogenetic framework.
Here we present the results of research conducted between
March 2004 and December 2006 regarding the distribu-
tional limits of C olallae and C modestus in the Beni De-
partment of Bolivia. Additionally we report on preliminary
data regarding their biology.


Study area
The area evaluated (Fig. 1) stretched from the Beni River to
the east of the Mamord River (eastern and western limits of
survey), and from the Rurrenabaque Yucumo road to Rib-
eralta (southern and northern limits of survey). We visited
43 sites across the region, placing more effort in the area
of the Yacuma River and in drier forest patches between
the Beni and Mamord rivers. Site selection was facilitated
using satellite images, and corresponded to the locations of
the original collection sites, the initial field observations of
Felton et al. (2006), new observations obtained during this
study, and reports from other researchers and local people.
However, it is important to remark that in general local
people only recognize titi monkeys at the genus level.

Determining the distributional limits
At each locality, we conducted preliminary non-structured
interviews with local people using photographic material
and vocalization recordings to determine the presence of
Callicebus and to identify specific potential locations for fur-
ther observation. We then actively searched for Callicebus
groups at a series of locations using vocalization 'playbacks'
at regular intervals in an attempt to elicit vocal responses.
Searches lasted between 06:00-10:00 h because these peri-
ods correspond to their highest vocal activity; complemen-
tary searches were made between 16:00-18:00 h. Whenev-
er we located a group, we recorded the time and method of
detection (response to playback, direct visual observation,
or spontaneous vocalizations). We observed groups with
Zeiss 10 x 40 binoculars and filmed them with a Sony Digi-
tal 8 Video Camera TRV 361. Vocalizations were recorded
with a Sony TCM-5000-EV tape recorder equipped with a
unidirectional Sennheiser ME 67 microphone. Upon first
sighting, we noted group size, composition, position, and
height in the forest canopy, as well as the geographic lo-
cation obtained with a GPS (Garmin 12 XL III Plus and
V). We also noted general habitat characteristics for each
observation locality.

Data analysis
Considering our initial lack of knowledge regarding the
morphological features of these monkeys in the field, spe-
cies identifications were made using available visual infor-
mation and comparing these data with descriptions made
by Lbnnberg (1939), Hershkovitz (1990), and also with
Felton et al. (2006), who had obtained video footage of
reported groups. To improve our identifications we also
compared images with photographs of specimens at the
Royal Natural History Museum of Stockholm, Sweden
(A612105 C modestus and A632187 C olidllwc). In order
to assist in the interpretation of distributional patterns, we
developed a map of all known localities, including the orig-
inal collection sites (Linnberg, 1939; Anderson, 1997),
the observations by Felton et al. (2006) and our ongoing
observations. The map also included sites where we did not
register Callicebus, although we differentiated between sites

Neotropical Primates 14(2), August 2007

where local people reported Callicebus and those where titi
monkeys were completely unknown.


Species identification
During this study, we encountered 66 groups of Callicebus
at 20 localities. Of these, 14 groups were Callicebus olallae,

31 were Callicebus modestus, 16 were Callicebus donacophi-
lus, and for five groups we were unable to assign a species
identification. These new records have dramatically in-
creased the number of observations for the endemic species
(Table 1).

For each species, we have identified diagnostic morpho-
logical characteristics helpful to distinguish the species

I Observation Sites of Callicebus

* -- a "--
in "- A ^


Figure 1. Geographic location of sightings of (:.... ,,, during the field survey. For current knowledge status of (:.... ,,, in Bolivia, we
are including in this figure the locations reported for C. aureipalatii and C. donacophilus from Anderson (1997), Wallace et al. (2006) and
Felton et al. (2006) and unpublished WCS observations.

Table 1. Number of documented C:.... ,,'n modestus and (: .... ,,', olallae groups.
Species L6nnberg (1939) Felton et al. (2005) This study Total groups
C. olallae 1 3 14 18
C. modestus 1 2 31 34
C cf. modestus* 0 1 0 1
* Supposed hybrid groups between endemic species (Felton et al. 2006).

Neotropical Primates 14(2), August 2007

in field. These features relate mainly to pelage coloration
patterns. Callicebus olallae has rather long, brown reddish
body pelage. Under good light conditions, a wide orange
band along each hair is visible, with the tip of the hair ap-
pearing brown. The tail is more grayish but the color does
not contrast strongly with the coloration of the body fur;
the base of the tail is lighter both dorsally and ventrally.
White ear tufts are very conspicuous. Narrow rims of black
hair that reach the ears are distinguishable around the faces
of some individuals. Hands and feet are black with some
white hairs visible. Another important feature is the verti-
cally elongated form of the head with a clearly prominent
and oval-shaped mouth especially noticeable in adult males
(Fig. 2). Body fur color appears more intensely red when
individuals are in direct sunlight.

Callicebus modestus has a non-uniform orange-brownish fur
color caused by the presence of alternate bands of orange
and dark brown hair, known as agouti pelage. The tail does
not have the conspicuous lighter basal zone, and the gray-
ish color of the tail is very noticeable and highly contrasting
with the rest of the body fur. Conspicuous white ear tufts are
present as in C. olallae. Hands and feet are also black with
light hairs visible, though not as light as in C olallae. How-
ever, this last trait is not easily distinguishable. Contrasting
with C. olallae, the head of adult male C. modestus is wider
laterally with a not too prominent pentagon shaped mouth
area apparently due to a more prominent nasal area not
observed in C. olallae (Fig. 3). The tail appears black if ex-
posed directly to sunlight, and appears lighter if backlit.

Determining the distributional limits
According to our observations, C olallae is largely restrict-
ed to riverine habitats on the Yacuma River, although one
group occurred 5 km east of the Maniqui River but in simi-
lar habitat. A partial preference for drier forest patches is
noticeable for C modestus, which occurred on the eastern
and western sides of the Yacuma River. One C modestus
group occurred just 100 meters east of the Maniqui River,
but this area was a higher and drier forest area. Overall,
these data give preliminary support to the hypothesis that
C olallae is found in relatively humid and riverine forest

Figure 2. Adult male C:,,....n olallae at Estancia La Asunta
(Yacuma River). Photo by M. Spanowicz.

Figure 3. Adult male C.. /,, modestus from Aguaizal (Northeast of Santa Rosa). Photo by M. Spanowicz.

Neotropical Primates 14(2), August 2007

in this patchily forested landscape; whereas C. modestus is
found in relatively drier forest patches (Fig. 1).

We determined the Beni River as the distributional western
barrier for both Bolivian endemics. During 2005, WCS
researchers recovered a titi monkey skin from a hunted
individual from forest immediately adjacent to the Beni
River on the western side. This specimen represents Callice-
bus aureipalatii, the new species recently described from the
Madidi protected area and registered only on the western
side of the Beni River (Wallace et al., 2006). In June 2005,
we observed C. modestus groups in forest immediately adja-
cent and on the eastern side of the Beni River. However, it
is important to note that the C modestus location around
the San Marcos community was in a noticeably drier belt
of forest than the majority of the relatively humid forest
found immediately adjacent to the Beni River. Indeed, on
several previous visits to the community we failed to regis-
ter Callicebus in the more humid sectors of this forest.

The southern and southwestern limits for the two endem-
ics occur in two broad 10 km swaths on either side of the
Yucumo Rurrenabaque road, an area characterized by
lack of primary forest and low densities of wildlife related
to several human settlements. Colonists who settled in
these areas almost ten years ago do not report titi monkeys.
In addition, no confirmed records exist for Callicebus in the
Pil6n Lajas Biosphere Reserve and Indigenous Territory,
and indigenous people indicate that titi monkeys are not
present within the reserve. These negative reports together
with areas of humid forest located to the south (as observed
in satellite images) suggest that these larger blocks of more
humid forest represent a southern limit for the distribution
of both endemic species.

We extended the known distributional limits (Hershkovitz,
1990; Anderson, 1997) for both C olallae and C modes-
tus. We found groups of C modestus between Yacuma and
Maniqui Rivers, and groups of both species in the riverine
forests of the Maniqui River. Subsequently we investigated
reports of the presence of C modestus to the east of the
Mamord River (M. Herrera, pers. comm. to R. Wallace),
but instead were able to verify the presence of Callicebus
donacophilus at this location. Additional observations of

C donacophilus east of the Maniqui and Mamord Rivers
suggest that neither C modestus nor C olallae occur further
east of the Maniqui River. Indeed, the San Borja region is the
current known eastern distributional limit for both species.
To determine the northern limit, we extended our surveys
to the relatively tall and humid Amazonian forests located
south of Riberalta (see Fig. 1). Here we observed Callicebus
groups that resembled species within the C. cupreus species
group (van Roosmalen et al., 2002). Unfortunately, due to
observation conditions, we were unable to assign species-
level identification to these groups. Nevertheless, we are
certain that these observations do not represent either of
the endemic species. We suggest that the southern limit
of this more humid forest probably represents a northern
boundary for C. modestus and perhaps C. olallae although
further field confirmation is recommended.

In the highly fragmented forests between Santa Rosa del
Yacuma and San Borja and northeast of Trinidad on the
eastern side of the Mamord River we failed to register Cal-
licebus, and people here report not to have ever seen or heard
these monkeys. Forest vegetation in these areas was very
scarce and highly patchy, with relatively small and rather
scattered stands of forest. In addition, forests did not have
abundant canopy vines-a feature that characterized Cal-
licebus localities in the broader southwestern Beni region.

Biological data
Group sizes ranged from 1 to 5 members in C. olallae and
1 to 6 in C. modestus. Solitary individuals were observed
on one occasion for each species. Average group sizes were
similar with 2.7 and 3.0 individuals for C. olallae and
C. modestus, respectively (t= -0,936; df= 43; n = 31; n 2= 14;
p= 0.354; see Table 2). Group composition data showed
similar proportions of adult and juvenile members for both
species although observations for each species occurred in
different seasons, thereby limiting comparisons.

Despite the riverine and non-riverine division of the
observations for C. olallae and C. modestus, respectively,
both species occurred in a similar habitat known locally as
"chaparrales" and characterized by a relatively low forest
canopy with dense lianas and the notable presence of
motacd palms (Attaleaphalerata) and/or garabata terrestrial

Table 2. Group size and composition of C. olallae and C. modestus.
Species C. ollle % C. modestus %
Group size 14 31
Maximum 5 6
Minimum 1 1
Mean 2.7 3.0
Population structure (no. of individuals) 38 94
Adults 28 73.7 62 66.0
Subadults 1 2.6 3 3.2
Juveniles 7 18.4 16 17.0
Infants 2 5.3 13 13.8

Neotropical Primates 14(2), August 2007

bromeliads (Pseudoananas sagenarius). We observed a
slight tendency for more motacd at C. olallae sites, and
more garabati at C. modestus sites. Local people reported
that Callicebus feed on motacd palms, and we found seeds
similar to garabati in C. olallae feces. The average height
of Callicebus observations did not differ between the two
species (t=-1,175; df=43; n1=31; n2 = 14; p=0.094;
Table 3). For both Callicebus species, adult males typically
appeared alert in the presence of researchers and often
approached observers, while other group members usually
remained still and stayed in relatively high branches.
Usually, upon achieving direct visual contact, monkeys
changed from a territorial call to an alarm call directed at
the observers and the rest of the primate group. These calls
are short harmonics that cover a wide frequency range. On
other occasions, groups stayed in dense vegetation without

Other primate species observed in the forest patches where
C olallae and C. modestus occurred included Saimiri boli-
viensis, Cebus libidinosus, and Alouatta sara. At the Yacuma
River, we observed Alouatta caraya. We registered Ateles
chamek in the riverine forest along the Beni River.

Identified local threats to Callicebus
Local communities reported some hunting of both Callice-
bus species, but their small size does not make Callicebus a
preferred game animal compared with other primates and
ungulates. Hunting pressure appears to be greatest along
the Maniqui River. The administrative policies of the pri-
vate cattle farms, numerous in the region, usually prohibit
hunting on their properties. We obtained occasional reports
of Callicebus hunting near cattle farms to use the skin for
making ropes and fur to make thread, although these uses
are not unique to these species. Unfortunately, hunting is
more of a threat during logging activities where workers
obtain most of their food from hunted wildlife, including
Callicebus. This activity is only relevant in the Beni riverine
forest on the western side of the endemics' distribution.

In the Amazonian forests in the northern reaches of the
survey area, interviews and anecdotal observations suggested
high levels of primate hunting, including Callicebus, espe-
cially during an intensive but short annual period of Brazil
nut harvesting that represents the major income source for
a large proportion of the local population. Indeed, for some
people hunting remains an important subsistence and op-

portunistically commercial activity throughout the year as
many people wait for the next Brazil nut harvest. As sug-
gested in informal interviews with local people, intensive
hunting pressure is probably responsible for the dramatic
change toward more evasive behavior in Callicebus in this


This study represents a significant contribution towards the
biological knowledge of these poorly known endemic and
range-restricted species. Direct and prolonged observations
allowed us to determine critical diagnostic phenotypic
traits for both C. olallae and C. modestus under a variety
of lighting and weather conditions. Pelage coloration is the
clearest characteristic that allows species identification in
the field. For C. olallae for at least some animals we con-
firmed the black ring around the face originally described
by Lonnberg (1939), contrasting with observations of the
same species by Felton et al. (2006). The contrast between
the color of the tail and body pelage is another important
difference, with the basal zone conspicuously lighter in
C olallae. The longer and shaggier pelage of C. olallae was
another recognizable trait. Differences in the density of
white hair on the ear tufts were not considered diagnostic
because observations suggested variability across pheno-
types in the same groups, although this may be related to
observation conditions that did not allow for the distinc-
tion of subtle features. We also noticed some distinctions
in the head shape of adult males that concur with cranial
measurements reported by Kobayashi (1995).

Video recording was a useful methodology for identify-
ing these species, although we did find some inconsisten-
cies between video records and direct observations, with
more orange colorations registered by video instead of the
more grayish color observed directly. We noted that some
lighting conditions affected the digital sensors of the video
camera. This problem is solvable by recording individuals as
they move through distinct light conditions. These results
show the great importance of making detailed phenotypic
descriptions, as well as video recordings under different
light conditions. Even though the morphological features
cited above allow us to make some clear phenotypic dis-
tinctions between these two primates, the final taxonomic
status remains unclear considering the potential formation
of hybrid groups reported by Felton et al. (2006). Thus,

Table 3. Observation height and canopy height details for C,...../',,n modestus and (,:.... ,, oaillae.
Species Average height obs. (m) SD Max. values Min. values
C. olallae 6.8 3.5 15 3
C. modestus 8.8 3.8 20* 3
Average canopy height (m)
C. olallae 8.9 3.0 15 5
C. modestus 10.2 2.8 15 5
*Observed in emergent tree

Neotropical Primates 14(2), August 2007

genetic analyses are required to confirm their taxonomic
status, clearly fundamental from a conservation viewpoint.

Felton et al. (2006) reported a group of C. olallae (No-
vember 2002) with more than two adults and during this
study local people reported the same for C. donacophilus
(February-March 2005). However, all of our observations
consist of groups with up to four or five individuals with
only two adults, always a male-female pair. Callicebus spe-
cies are monogamous (Kinzey, 1981; Rylands et al., 2000;
Van Roosmalen et al., 2002), and therefore larger groups
containing several adults might be the result of temporary
or seasonal aggregations, offspring remaining in their natal
group after sexual maturation (Eckhard W. Heymann, pers.
comm. 2007), and/or temporal variations in group size due
to reduced dispersal opportunities in forest patches (Bicca-
Marques et al., 2002).

The key result for both C olallae and C modestus is the
extension of the eastern limit from the previously known
distributional area in the Beni and Yacuma Rivers (An-
derson, 1997; Felton et al, 2006). Our results also offer
a preliminary northern distributional limit for both spe-
cies although further work is required to establish how this
limit varies for each species. Our results show that C olallae
is restricted to riverine forests along the Yacuma and Ma-
nique Rivers, whereas C modestus seems to occupy patches
of forest in a larger area between the Beni and Maniqui
Rivers. We therefore suggest that C olallae has an extreme-
ly restricted distribution with groups almost entirely con-
centrated on the Yacuma River, and so far just one group
registered along the Maniqui River. Nevertheless, the habi-
tat preferences of both species need further study, particu-
larly given that both species are absent from smaller forest
patches in the region.

Distribution points for these endemic species and their
congeners in the broader region (Fig. 1), suggest IUCN
"extent of occurrence" distributional areas of ca. 1,800
km2 for C. modestus and ca. 400 km2 for C. olallae. Nev-
ertheless, because of the fragmented nature of the forest
landscape in the region, the IUCN "area of occupancy"
standard measure of distributional area seems more per-
tinent. Under this measure C modestus should be provi-
sionally considered Endangered (c.450 km2) and C. oallae
should be provisionally considered Critically Endangered
(c. 50 km2). To date no abundance estimates for these spe-
cies exist, however, given that they are not superabundant
in the forest patches we have visited, in terms of popula-
tions we recommend that both species be considered en-
dangered. The most important threat to both endemics is
the forthcoming improvement and paving of the "Corre-
dor del Norte" road. This road crosses the distributional
area of both species (Fig. 1) and although the use of heavy
machinery, noise, and pollution are concerns, there is a
high probability for increasing human settlements and
forest loss along the road. Given the suspected low popu-
lation numbers for both species, significant forest loss and

fragmentation would lead to a very fragile conservation
situation and a potential extinction risk, principally for
C. olallae. Information collected to date has been provid-
ed to the official environmental evaluation team for the
proposed Corredor del Norte road construction project;
this team was previously unaware of the presence of these
endemics and are now proposing to make specific consid-
erations within their recommendations. Subsistence and
commercial hunting is also a major threat in areas under-
going forestry and Brazil nut activities. Even though cur-
rent eco-tourism activities on the Yacuma River are not an
obvious threat to Callicebus, the presence of C olallae in
the tourism area indicates the need to monitor the impact
of this activity in the future.

The information gathered to date has been provided to
municipal authorities, and these meetings and presenta-
tions have drawn the attention of local decision-makers to
the presence of these endemic titi monkeys. Now the cre-
ation of municipal protected areas is being promoted in the
region. Municipal authorities are motivated by the status of
the Yacuma River region as a major tourism attraction; over
75% of the 20,000 tourists that visit Rurrenabaque every
year visit the Yacuma River region. Indeed, the two endem-
ic titi monkeys could well become important flagship spe-
cies for this initiative and we are working to ensure they are
included in the design and management of the proposed
municipal reserves. Finally, both C modestus and C. olal-
lae have restricted distributional ranges. The situation of
C. olallae is particularly alarming, and given the imminent
threat of major road improvements in the region, the race
is on to establish the conservation status of these endemics.
Knowledge of the precise northern distributional limits is
urgently required for both species, as well as population
estimates for both species in a series of strategically targeted
forest patches. Finally, determination of their taxonomic
status through genetic analyses is also an urgent priority.


We would like to thank the Wildlife Conservation Society,
the Gordon and Betty Moore Foundation, Primate Con-
servation Inc., the Margot Marsh Biodiversity Founda-
tion, and the Conservation International Primate Action
Fund for their financial support. We are also grateful to the
Caceres brothers (Jairo, Ernesto "Billy" and Jasmani) for
their support during fieldwork. We would like to thank the
National Directorate for the Protection of Biodiversity for
help in acquiring necessary research permits.


Anderson, S. 1997. Mammals of Bolivia: Taxonomy and
Distribution. Bulletin of the American Museum of Natu-
ral History. New York.
Aquino, R. and Encarnaci6n, F. 1996. Distribuci6n geogri-
fica de Saguinus tripartitus en la Amazonia del Perl.
Neotrop. Primates 4: 1-4.

Neotropical Primates 14(2), August 2007

Bicca-Marques, J. C., Garber, P. A. and Azevedo-Lopes,
M. A. 0. 2002. Evidence of three resident adult male
group members in a species of monogamous primate,
red titi monkey (Callicebus cupreus). Mammalia 66:
Felton, A., Felton, A. M., Wallace, R. B. and G6mez, H.
2006. Identification, distribution and behavioral obser-
vations of the titi monkeys Callicebus modestus Lonnberg
1939, and Callicebus olallae Lonnberg 1939. Primate
Conserv. (20): 40-46.
Forman, R. and Deblinger, R. 2000. The ecological road-
effect zone of a Massachusetts (U.S.A.) suburban high-
way. Cons. Biol. 14: 36-46.
Hershkovitz, P. 1988. Origin, speciation and distribution
of South American titi monkeys, genus Callicebus (Family
Cebidae, Platyrrhini). Proc. Acad. Nat. Sci. Philadelphia
140: 240-272.
Hershkovitz, P. 1990. Titis, New World monkeys of the
genus Callicebus (Cebidae, Platyrrhini): A preliminary
taxonomic review. Fieldiana Zoology. New Series 55:
Kinzey, W. G. 1981. The titi monkeys, genus Callice-
bus. In: Ecology and Behaviour of Neotropical Primates,
Vol. 1, A. E Coimbra-Filho and R. A. Mittermeier (eds.),
pp.241-276. Academia Brasileira de Ciencias, Rio de
Kobayashi, S. 1995. A phylogenetic study of titi monkeys,
genus Callicebus, based on cranial measurements: I. Phyl-
etic groups of Callicebus. Primates 36: 101-120.
Lonnberg, E. 1939. Notes on some members of the genus
Callicebus. Arkivfor Zoologi 31A (13): 1-18.
Reid, J. and Landivar, R. 1997. Consecuencias econ6micas
y biol6gicas de la construcci6n de caminos en las tierras
bajas de Bolivia: Un m6todo de evaluaci6n ripida. Docu-
mento Tcnico 53 /1997. BOLFOR, Santa Cruz.
Rowe, N. and Martinez, W. 2003. Callicebus sightings in
Bolivia, Peru and Ecuador. Neotrop. Primates 11: 32-35.
Rylands, A. B., Scheider, H., Langguth, A., Mittermeier,
R. A., Groves, C. P. and Rodriguez-Luna, E. 2000. An
assessment of the diversity of New World primates.
Neotrop. Primates 8: 61-93.
Trombulack, S. and Frissell, C. 2000. Review of ecological
effects of roads on terrestrial and aquatic communities.
Cons. Biol. 14: 18-30.
Van Roosmalen, M. G. M., Van Roosmalen, T. and Mit-
termeier, R. A. 2002. A taxonomic review of the titi mon-
keys, genus Callicebus Thomas, 1903, with the description
of two new species, Callicebus bernhardi and Callicebus
stephennashi from Brazilian Amazonia. Neotrop. Primates
10: 1-52.
Wallace, R. B., Gomez, H., Felton, A. and Felton, A.
2006. On a new species of titi monkey, genus Callicebus
Thomas, from western Bolivia (Primates, Cebidae) with
preliminary notes on distribution and abundance. Pri-
mate Conserv. (20): 29-39.

Neotropical Primates 14(2), August 2007 55


Paula Giraldo1' 2, Carolina G6mez-Posada1, Jesus Martinez1' 2 and Gustavo Kattan'

SFundacion EcoAndina/l' -j Conservation Society Colombia Program, Apartado Aereo 25527, Cali, Colombia, e-mail:
., .'............ >, , ,
2 Departamento de Biologia, Universidad del Valle, Cali, Colombia


As part of a study of responses by red howler monkeys (Alouatta seniculus) to forest fragmentation, we studied red howlers in
an extensive forest including a habitat mosaic of old-growth forest, secondary forest of different ages, and non-commercial
monospecific plantations of exotic Chinese ash (Fraxinus chinensis) and pines, to produce baseline information on resource
use and their role as seed dispersers. We followed three troops for six months, encompassing a dry and a wet season; in total,
they fed on 48 species of trees, including fruits, leaves and flowers, but each group relied on a limited set of species available
within its home range, particularly in the Moraceae and Cecropiaceae. This contrasts with lowland forest, where howlers
feed on a larger variety of species. We observed no season of generalized fruit scarcity, as is the case in the lowlands. However,
localized fruit scarcities in howler home ranges occurred. Howlers fed on relatively small quantities of leaves when fruit was
available, and became primarily folivorous when fruit was unavailable. Howlers transported a mean of 2.3 species of seeds
per fecal sample, to a mean distance of 116 m from the parent tree. More importantly, howlers are transporting seeds to ash
and pine plantations, helping to enrich these habitats.

Key words: Andes, Colombia, red howler monkey, frugivory, folivory, montane forest, cloud forest, seed dispersal, habitat


Como parte de un studio sobre la respuesta del mono aullador rojo a la fragmentaci6n de bosques, estudiamos una pobla-
ci6n en un bosque extenso que incluia un mosaico de habitats de bosques maduros, bosques de regeneraci6n de distintas
edades, y plantaciones monoespecificas de urapin o fresno de la China (Fraxinus chinensis) y pinos, para producer informa-
ci6n de base sobre patrons de uso de recursos y su papel como dispersores de semillas. Seguimos tres grupos de aulladores
durante seis meses, que incluyen una estaci6n seca y una h6meda. Los tres grupos se alimentaron de 48 species de irboles,
incluyendo frutos, hojas y en menor media flores, pero cada grupo dependi6 de unas pocas species de acuerdo a la dispo-
nibilidad de aquellas en su irea de actividad, particularmente en el Moraceae y Cecropiaceae. Esto contrast con los bosques
de tierras bajas, donde esta especie se alimenta de una variedad de species much mayor. No observamos una 6poca de
escasez generalizada de frutos, como si ocurre en tierras bajas, sin embargo, si observamos escasez localizada en las ireas de
actividad de los grupos estudiados. Los aulladores siempre consumieron pequefias cantidades de hojas cuando habia frutos
disponibles, y fueron principalmente folivoros cuando los frutos fueron escasos. Los aulladores dispersaron un promedio de
2.3 species de semillas en sus heces, a una distancia promedio de 116 m del arbol parental. Lo mris important, sin embargo,
es que estin transportando semillas a las plantaciones de urapin y pino, ayudando a enriquecer estos habitats.

Palabras clave: Andes, Colombia, mico aullador rojo, frugivoria folivoria, bosque montano, bosque nublado, dispersi6n de
semillas, restauraci6n de habitat

Introduction and cloud forest (Hernindez-Camacho and Cooper, 1976).
They can also use secondary forest and survive in isolated
The red howler monkey, Alouatta seniculus, is widely dis- patches (Rylands and Keuroghlian, 1988; Schwarzkopf
tribute in northern South America. It is found in the and Rylands, 1989). In Colombia the red howler is dis-
northern and western Amazon basin and east to the Guy- tribute from sea level to 2,400 m a.s.1. and occasionally
anas, and in the Andes from Colombia to Bolivia. Red up to 3,200 m a.s.1. (Hernindez-Camacho and Cooper,
howler monkeys occupy a variety of forest habitats, from 1976; Defler, 2003). Howler monkeys (Alouatta spp.) have
lowland rain forest in the Amazon, to gallery forest and a folivorous-frugivorous diet, although they do not possess
relatively open woodland in savannas, tropical dry forest, a highly specialized digestive tract to deal with leaves, nor

Neotropical Primates 14(2), August 2007

do they exhibit particularly low metabolic rates (Milton,
1998). As frugivores, howlers are recognized as important
seed dispersers in tropical forest, because they have long in-
testinal retention times and they move seeds away from the
parent trees (Julliot, 1996a; Yumoto et al., 1999). Second-
ary dispersers may then scatter these seeds and contribute
to their germination (Andresen, 1999, 2002).

The ecology of red howlers has been well studied in a vari-
ety of lowland forest types. However, there is only one de-
tailed study available for Andean populations (Gaulin and
Gaulin, 1982). This one-year study examined food habits
of one group of howlers at a cloud forest in the Central
range of the Colombian Andes, and reported consumption
of fruits and leaves of 34 plant species. There are reasons
to believe that aspects of the behavioral ecology of howl-
ers may differ between lowland and montane forest. First,
plant diversity decreases with elevation and species com-
position changes (Gentry, 1992; Cavelier et al., 2001), so
montane populations may have a different and more re-
stricted choice of food species. Second, primary productive
decreases with elevation in correlation with a decrease in
temperature. These factors may affect activity patterns and
space and resource requirements of howlers.

Montane populations of red howler monkeys in Colombia
are threatened by habitat destruction and fragmentation.
Recent estimates indicate that over 70 percent of Andean
forests have been transformed, and most of what remains is
in isolated patches, particularly within the howlers' range in
the inter-Andean valleys (Kattan and Alvarez-L6pez, 1996;
Cavelier et al., 2001). Across Latin America, howler mon-
keys have not escaped the negative consequences of human-
caused disturbances of natural habitats. Some populations
of mantled howlers (Alouatta palliata) have gone locally
extinct, and many populations of Alouatta spp. barely per-
sist in isolated fragments, under conditions that put them
at risk (Estrada and Coates-Estrada, 1988, 1996; G6mez-
Posada, unpublished data). However, some studies suggest
that howlers can persist in isolated forest patches, relying
on a limited subset of plant food species in the families
Moraceae and Cecropiaceae (Estrada and Coates-Estrada,
1986; Rylands and Keuroghlian, 1988; Schwartzkopf and
Rylands, 1989; Estrada et al., 1999).

Understanding the ecology of red howler monkeys and
their role as seed dispersers in montane forest fragments is
critical to the conservation of both the monkeys and their
habitat. Here we present results of a six-month study of
red howler monkey at Otin-Quimbaya Flora and Fauna
Sanctuary, a cloud forest site in the Central Cordillera of
the Colombian Andes. The study site is a restored forest,
formed by a mosaic of remnant patches of old-growth
forest, second-growth forest and non-commercial mono-
specific tree plantations that were established as part of a
revegetation program. Our study area (489 ha) is part of
a larger forested area (several thousand hectares) and is the
largest remnant in the howlers' range on the western slope

of the Central Cordillera of the Colombian Andes. This
study provides baseline data for a larger study, currently
in progress, documenting patterns of habitat and resource
use, and space requirements of red howler monkeys in
forest fragments in the Colombian Andes (G6mez-Posada,

Study Area and Methods

The study was conducted at the Santuario de Fauna y
Flora Otin-Quimbaya (0443'N, 7528'W), located on
the western slope of the Central Cordillera of the Andes
of Colombia, east of the city of Pereira. This reserve pro-
tects a mosaic of Andean forests of different regeneration
ages, including patches of old-growth forest, and patches of
different sizes of native Andean oak (Quercus humboldtii),
exotic Chinese ash (Fraxinus chinensis) and conifers. These
monospecific tree stands were planted in the 1960s as part
of a revegetation and soil stabilization program initiated by
the local utility authority to protect the Otin river drain-
age (Londofio, 1994). Tree plantations were abandoned to
natural regeneration and presently have a high plant di-
versity, particularly in the understory (Durin and Kattan,
2005), although the canopy is dominated by the planted
species. The Sanctuary encompasses 489 ha at elevations
between 1,800 and 2,100 m a.s.l., but is contiguous with
Ucumari Regional Park, which protects 3,980 ha of forest
between elevations of 1,750 and 2,600 m a.s.l. Precipita-
tion in the region is bimodal, with peaks of rain in April
and October and an annual mean of 2,712 mm (Estaci6n
Metereol6gica El Cedral, CENICAFE, 1995-2001), and
the mean annual temperature is 15C (Aguilar and Rangel,
1994). A mild dry season occurs in December-January and
a stronger one in July-August, when monthly precipita-
tion may be under 100 mm.

The study was carried out during July-December 2001,
including a dry and a wet season. We identified 11 monkey
troops (mean group size SD = 7.3 2.5 individuals) in an
area of 113 ha, and selected three groups (labeled C, D, and
E; Table 1) for intensive observations (Giraldo, 2003; Mar-
tinez, 2003). Each group used home range of 10.2 3.3 ha
during the six-month study period (G6mez-Posada et al.,
2004). The home range of troop C (8 ha) was completely
contained in mature forest. Troop D's home range (14 ha)
was next to the river, in an area of secondary forest (old
enough to have a developed canopy with interlocking tree
crowns), Andean oak, ash and pine plantations. Troop E
(8.7 ha) used mostly an ash plantation and a small area of
secondary forest. We tried to follow each group for three
to four consecutive days each month, making observations
between 06:30 and 17:00. We obtained 6-9 observation
hours per day, depending on weather; monkeys remained
inactive during cold days. The harsh topography also made
following troops difficult. We recorded diet every 15 min
using a slow scan method (Altmann, 1974, adapted by
Robinson, 1986); in each sample, we scanned the group for
ten minutes, and for each monkey, we noted the first item

Neotropical Primates 14(2), August 2007

consumed. In each scan we noted the type of food eaten
(leaves, mature or immature fruits, flowers), the troop's
location, and the habitat type. The proportions of use of
different food items were compared with a G contingency
test (Zar, 1996). Voucher samples of plants were collected
and identified at several national herbaria (see Rios et al.,

To estimate availability of resources used by howl-
ers, we measured fruit and new leaf production in three
400 x 4 m transects (total area sampled = 4,800 m2) distrib-
uted in an area of about 100 ha where the study troops oc-
curred. No previous studies have evaluated resource avail-
ability for montane populations of red howler monkeys so
there were no criteria for deciding transect size. We chose
to use fixed-width transects (Stevenson et al., 1998; Hem-
ingway and Overdoff, 1999) and the size of the transects
was decided by taking into account the lower diversity and
smaller spatial scale of montane forests, compared with
lowland forests (Gentry, 1992; Cavelier et al., 2001; Sil-
verstone-Sopkin, pers. comm.). We included all trees over
15 m tall. We monitored these transects once per month to
estimate new-leaf production (estimated as the proportion
of the tree's canopy with young leaves) and number of ripe
fruits in canopy trees. Fruits were directly counted in one
tree branch and multiplied by the number of branches to
extrapolate to the whole tree. We explored the correlation
of monthly fruit and leaf availability with intensity of use
(for each group) using Spearman's rank correlation coef-
ficient (Zar, 1996).

To complement feeding observations and document seed
dispersal, we collected feces from the forest floor and un-
derstory vegetation, as soon as monkeys defecated. Def-
ecation in howlers was a group activity; usually the alpha
male initiated a defecation bout and all members of the
group followed. Samples were washed, dried and filtered
to separate fiber and seeds, and we counted manually all
seeds >5 mm and estimated numbers of seeds <5 mm by
weighing samples of 100 seeds. To estimate dispersal dis-
tances from parent trees, we made ad libitum observations
(Altmann, 1974) while following a troop for at least two
continuous days; we recorded the time and place of con-
sumption of fruits at a particular tree, and the site where
seeds were defecated afterwards (making sure monkeys had
not fed on another tree of the same species). Points were
located on a map of the study area and dispersal distances

measured as a straight line. We included data only on def-
ecations that occurred after 20 h of beginning observations
of a troop feeding on a particular tree, to make sure that
seeds originated from this particular tree. Twenty hours is
the mean retention time estimated for red howlers by Jul-
liot (1996a), Andresen (1999) and Yumoto et al. (1999).


Availability of resources
In the three transects we recorded a total of 74 species of
trees and a mean (+ SD) of 83.3 30 individual trees per
transect, representing about one third of the species of
canopy trees in the study area (Rios et al., 2004). Avail-
ability of new leaves was low during the six months of the
study, but some species like Ficus spp., Cecropia -
and some Clusiaceae produced new leaves all the time. In
general, trees in this cloud forest were nondeciduous and at
any particular moment had a low proportion of the crown
(- 5 %) covered in new leaves. The only deciduous species
were Andean oak and Chinese ash (leaves not consumed
by howlers in this study). In July and August availability
was high because of synchronized leafing in Andean oak
(Fig. 1).

Both the total fruit availability and the number of spe-
cies in fruit peaked in August and November (Fig. 1).
The August peak was due to fruiting of Garcinia sp. and
Wettinia kalbreyeri. In November the peak was produced
by Heliocarpus americanus (which has a dry fruit not con-
sumed by howler monkeys) and the palm W kalbreyeri. In
general, fruit production in the forest showed little syn-
chronization, both intra- and interspecifically, and reflected
fructification events of individual trees. This was the case of
one individual tree of Ficus killipii, and one Garcinia sp.,
which produced fruit during two non-consecutive months;
this particular Ficus tree was an important food item for
troop E during this study (Appendix and see below).

Diet composition and food preferences
Although our aim was to follow each group for 3-4 days
per month, the rough topography of the study area made
it difficult to always locate and follow groups, in spite of
intensive search efforts. Groups C and D were followed
during five months and group E during four non-consecu-
tive months. We obtained a total of 1,643 feeding records
of fruits, leaves and flowers, in 388.3 hours of observation

Table 1. Size, composition and sampling effort (hours and days of observation) of three red howler monkey troops studied during six
months in the Central Cordillera of the Colombian Andes (M = male, F = female).
Group Adult Subadult Juvenile Infant Total No. hours /
M F M F M F No. days
C 1 2 1 1 5 119.4/ 18
D 1 3 1 1 1 1 2 10 136.4/ 19
E 1 2 1 1 1 2 8 132.0 / 15
Total 3 7 2 2 2 2 5 23 388.3 / 52

Neotropical Primates 14(2), August 2007

for the three howler groups in the six months of study. They
fed on 48 species of plants, in some cases including fruits
and leaves or fruits and flowers of the same species (Ap-
pendix). In addition, we once observed the adult male of
group C feeding on tree bark, and a juvenile twice sucked
and then dropped mouthfuls of moss obtained from a cy-
press tree.

The diet was mostly folivorous (49.2% of feeding records)
and frugivorous (45.1%); flowers had a low representation
in the diet (5.7%; Table 2). It was difficult to distinguish
young and mature leaves through binoculars, particularly
for species such as Paullinia sp., Ficus spp. and Macrolo-
bium colombianum, so we pooled all leaf-feeding records
in one category. The most important plant families, with
both fruits and leaves consumed, were the Moraceae
(17 species with 45.7 percent of the total of 1,643 feeding
records, including several species of Ficus representing 42.2
percent of feeding records) and Cecropiaceae (one species,
C. -. representing 17.2 percent of feeding records).
The leaves of two other species, Paullinia sp. (Sapindaceae)
and Macrolobium colombianum (Leguminosae) also made
up an important proportion of feeding records (Appendix).
During their daily travel routes (553.9 247.9 m; G6mez-
Posada et al., submitted), howlers always found at least one
fruiting tree within their home range each month. Particu-
larly important in this regard were C. -. and Ficus
spp., which usually had one or more trees in fruit within

Figure 1. Resource availability for howler monkeys in the Cordil-
lera Central of Colombia. Graph la shows the mean proportion
per transect of tree canopy with new leaves, and graph lb shows
the mean number of fruits per transect (N = 3 transects).

the study area. However, on any particular day howlers fed
on a combination of fruits, leaves, and occasionally flow-
ers, of several species. Monthly rates of consumption of
the different food items varied widely (Fig. 2). This reflects
intense and opportunistic feeding in particular trees when
resources were available, as was the case with individual
E .11//.p' and Garcinia sp. trees that produced fruit abun-
dantly in non-consecutive months. There were significant
differences among troops in diet composition (G = 328.7,
P < 0.01; Table 2).

m LVs Frt 1 Imm = FIw
100 1
: Troop C


so- Troop D

I eo43

TroopE 310



ms Troop E 102



Jul Aug Sep Oct Nov Dec

Figure 2. Diet composition of three groups of red howler monkeys
in the Central Cordillera of the Colombian Andes. Bars show the
proportion of feeding bouts of four items (Lvs = leaves, Frt = ripe
fruits, Imm = unripe fruits and Flw = flowers) in the diet, in ob-
servations made between July and December 2001. Numbers
above bars indicate number of observations per month.



Jud Aug Sep Oct

Nov Dec

Neotropical Primates 14(2), August 2007

Troop C, with a home range in mature forest, fed on a total
of 21 tree species and visited a mean of 5.4 (SD = 1.6) spe-
cies per day. The most important items in the diet of this
group were leaves of Paullinia sp. and M. colombianum, and
leaves and fruits of C. -. and two species of Ficus, in
combination accounting for 65 percent of feeding records
(Appendix). Another important species in this group's diet
was Otoba lehmannii, which produced fruit continuously
during this study, representing most of the immature fruit
category (Fig. 2); however, howlers only took a small bite
of the tissue of the area of insertion of the peduncle and
dropped the rest of the fruit. In July we were able to follow
this group for only one day, hence the small number of ob-
servations, all on 0. lehmannii (Fig. 2). During November,
this group fed heavily on flowers of Macrolobium colom-
bianum (Fig. 2), which flowered synchronously throughout
the forest.

Troop D, which had a mixture of ash and pine plantations
and secondary forest in its home range, had a more varied
diet. This troop fed on a total of 27 species, visiting a mean
of 4.7 (+ 2.1) species per day. This group fed to some extent
on immature fruits of Chinese ash (September and Novem-
ber; Fig. 2), but mostly relied on fruits and leaves of Cecro-
pia -. and several species of Ficus (Appendix). In July,
little fruit was available within this group's home range, and
monkeys relied heavily on leaves (mostly Cecropia; Fig. 2).

Finally, troop E had a home range mostly restricted to ash
plantation, with some secondary forest. The diet of this
group was limited to 12 species, with a mean of 4.4 ( 1.9)
species visited per day. However, three species represented
82.9 percent of feeding records (Appendix). This group
relied heavily on leaves of a small number of trees, except
during October and December, when it fed on fruits of
a particular Ficus killipii tree for 58.4 percent of feeding
records (Fig. 2). Other important items were C . .
(fruits and leaves) and Paullinia sp. (leaves). This troop
also fed to some extent on immature ash fruits in October
(Fig. 2 and Appendix).

Only 27 of the 48 species eaten by the howlers were rep-
resented in the three transects. Several species that were
important in the monkey's diet, such as Ficus 4/.1/.p., and
Paullinia sp. (Appendix), were not found in transects.
Some Macrolobium colombianum and Garcinia sp. trees

Table 2. Diet composition of three red howler monkey troops

were present in transects, but these species fruited asyn-
chronously throughout the study area. Howlers fed on
these species, but not at the time when they were fruiting
in transects; thus, we have no monthly estimate of fruit
availability for these species, and could not calculate any
preference index. There was no correlation between fruit
and new leaf availability (Fig. 1) and consumption of these
items by each group (Fig. 2; P > 0.3 in all cases). However,
leaf consumption in groups D and E decreased when fruit
availability increased.

Seed dispersal
Each troop defecated two to four times daily, usually under
a single tree. It was difficult to keep individual feces separate
because they were scattered around when hitting vegetation
under the tree, so we collected all feces from each group
defecation bout, combined them into a single sample, and
identified and counted numbers of seeds. Fecal samples
contained a mean of 2.3 (SD= 1.04) species of seeds. We
found a total of 9 species of intact seeds and 290,174 seeds
in 60 samples. Seed size was highly variable, but most seeds
(99.9%) were less than 5 mm long; this was the case with
C. -. (<3 mm), Ficus spp. (<0.8 mm) and Miconia
acuminifera (<0.5 mm), which were the most abundant
species in fecal samples (Table 3). The largest seeds found
in feces were Garcinia sp. (33.8 0.54 mm, N= 20) and
Allophylus mollis (8.2 0.05 mm, N= 20). Only five of
the 9 seed species found in feces were observed in feed-
ing records, while the other four species occurred in small
numbers and we could not identify them. All seeds of Ficus
spp. are very similar and we could not separate the differ-
ent species, however, we observed consumption of fruits of
at least five species of Ficus so the total number of species
in the feces could be 13 or more. Dispersed seeds showed
no evidence of parasitism, except for A. mollis, which had
some seeds parasitized by a coleopteran larva. We obtained
33 direct dispersal distances from the feeding point in the
parent tree to the defecation site (we could not obtain more
dispersal distances because howlers sometimes fed on dif-
ferent trees of the same species in a single day and because
we could not obtain 20 h of continuous following). In
14 cases, this distance was between 6 and 8 m (defecation
under same tree). In the other 17 cases, mean dispersal dis-
tance was 116.1 m (SD = 92.1). Because howlers frequently
visited the same feeding tree several times in a single day, it
was not possible to determine retention times.

in the Andes of Colombia, measured as percent of feeding bouts

Item Consumed Troop Three troops combined
Leaves 57.3 59.2 34.5 49.2
Ripe Fruits 17.0 31.4 60.6 39.2
Unripe Fruits 8.4 5.3 4.9 5.9
Flowers 17.3 4.0 0 5.7
N (feeding bouts) 405 601 637 1,643

Neotropical Primates 14(2), August 2007

Table 3. Number of seeds of nine species of plants found in 60 troop-composite fecal samples of three red howler monkey troops in the
Andes of Colombia.
Plant species Monkey troop
Cecropia telealba 54,545 56,587 58,276
Ficus spp. 5,461 4,870 100
Miconia acuminifera 43,076 38,986 28,085
Allophylus mollis 100 11 2
Garcinia sp. 7 0 0
Dendropanax macrophyllus 30 0 7
Simplocus sp. 2 11 0
Unidentified 1 8 0 0
Unidentified 2 2 8 0
Total 103,231 100,473 86,471


Resource availability and diet composition
We observed three important differences between mon-
tane and lowland howler populations in their feeding
ecology. First, during our six-month study, we observed
no times of generalized fruit scarcity at the level of the
entire forest, as is the case in lowland forest (Milton,
1980; Julliot, 1996a; Stevenson et al., 2000; Palacios and
Rodriguez, 2001). Instead, fruit was available in small but
relatively constant amounts, although localized scarcities
within the howlers' home ranges occurred (but our study
included only six months, so we have no information on
long-term patterns). Intra- and interspecific asynchrony in
the phenology of cloud forest trees has been reported for
other Andean sites (Ataroff, 2001; Cavelier et al., 2001).
Fruiting of many species, especially those important in the
monkeys' diet (Ficus, Cecropia), tended to occur asynchro-
nously. Howlers at our site usually combined fruit and
leaves in their diet, although leaf use tended to decrease
when fruits were available. They fed on the leaves of a wide
variety of species, but each species was consumed in small
amounts (as proportion of feeding bouts), although some
species (Paullinia sp., Ficus spp. and C. telealba) produced
new leaves all the time and were more continuously used
(Appendix). The same general feeding pattern was report-
ed by Gaulin and Gaulin (1982) for another site on the
same mountain range. We did not observe consumption
of oak leaves (Quercus humboldtii), as Gaulin and Gaulin
(1982) did, even though home ranges of two groups in-
cluded oak stands.

Second, the number of species of plants on which howl-
ers fed was small, probably reflecting lower plant diversity
compared to the lowlands. This is particularly true at our
study site, which is a restored, secondary forest. We re-
corded a total of 48 species in their diet (fruits and leaves
combined), but for one group this number was as low as
12 species in four months, and Ficus spp. and C . .
represented more than 60 percent of the diet of the three

groups. The three groups fed on a total of 14 fruit species but
these numbers may increase with longer study. The group
studied by Gaulin and Gaulin (1982) in a mature forest fed
on a total of 34 fruit species in one year; in contrast, studies
on lowland populations usually report higher numbers. For
example, a 2-year study of one troop of red howler mon-
keys in French Guiana reported a total of 195 species in
the diet, including 97 species of fruits (Julliot and Sabatier,
1993). Two other studies in eastern Colombia reported
17 species of fruits consumed by one troop in 9 days
(Yumoto et al., 1999), and 43 species consumed in
12 months by one troop (Stevenson et al., 2002). The third
difference is in the size and color of fruits. Howler monkeys
are reported to prefer large, yellow or orange-colored fruits
(Janson, 1983; Julliot, 1996b). At our site, most fruits con-
sumed by howlers were small (10-40 mm, except for C.
-.. I. and many were green or whitish (such as some
Ficus spp. and A. mollis) or purple (Miconia acuminifera),
and the only large and yellow fruit in their diet was Gar-
cinia sp. Our study did not encompass a full year cycle,
but few large and yellow or orange fruits in canopy trees
are available at this cloud forest site (Rios et al., 2004), and
most were not consumed by howlers.

Availability of resources varied among the three groups,
reflecting the heterogeneous nature of our study area. In
particular, the group living in ash plantation had a restrict-
ed diet, and resorted to leaves when little fruit was avail-
able within its home range. Most resources used by troops
in ash plantations are in neighboring secondary forest and
a few native trees dispersed within the plantations. Our
measures of monthly fruit and leaf availability did not cor-
relate with consumption, but this could be a result of our
transects not adequately representing resource availability
and our study not covering a full year cycle. The observed
intra- and interspecific asynchrony in fructification sug-
gests that a better estimate of fruit availability in mon-
tane forest could be obtained by placing more but shorter
transects dispersed throughout the study area and inside
howler home ranges.

Neotropical Primates 14(2), August 2007

Monkeys at our study site depended heavily on a few
species in the Moraceae, especially Ficus spp., and Cecro-
pia telealba. This capacity to rely on a limited set of fruit
species, in addition to folivory, allows howler monkeys
to survive in small and degraded forest fragments, and in
anthropogenic habitats (Rylands and Keuroghlian, 1988;
Schwartzkopf and Rylands, 1989; Coates-Estrada, 1996;
Estrada et al., 1999, 2002; G6mez-Posada et al., 2005).
Throughout tropical America, the Moraceae (particularly
Ficus, Morus, Brosimum, Poulsenia and Pseudolmedia) and
Cecropiaceae have emerged as critical resources for dif-
ferent species of howlers, in particular in second-growth
habitats (Gaulin and Gaulin, 1982; Milton, 1991; Julliot
and Sabatier, 1993; Silver et al., 1998; Estrada et al., 1999,
2002; G6mez-Posada et al., 2005).

Seed dispersal
Howler monkeys are important members of the
seed-disperser guild in tropical forests, because of their
abundance, biomass, flexibility of habitat use, and the sheer
number of seeds they move within and between habitats
(Estrada and Coates-Estrada, 1984; Julliot, 1996a; Silver et
al., 1998; Yumoto et al., 1999). At our study site, howlers
defecated intact seeds of at least nine species that were able
to germinate (Giraldo, 2003). The number of dispersed
seed species at our site, however, is small compared to low-
land forest sites. For example, one group of howlers dis-
persed nine species in 9 days in eastern Colombia (Yumoto
et al., 1999) and one group in French Guiana dispersed 86
species in 2 years (Julliot, 1996a).

On the other hand, howlers were seed predators for three
species at our site (Fraxinus chinensis, Wettinia kalbreyeri
and Otoba lehmannii) because they ate unripe fruits. They
also sometimes dropped the large seeds of Garcinia sp.
under the parent tree after partially eating the fruit. Howler
feces at our site always had seeds, usually of several species.
Most seeds were small, and in preliminary tests many were
removed by dung beetles (Giraldo and G6mez-Posada,
unpublished data), so were less likely to be predated by
rodents. Removal of seeds by secondary dispersers such
as dung beetles contributes to increased seed survival (Es-
trada and Coates-Estrada, 1986; Andresen, 1999, 2002).
Although many seeds were deposited under the parent tree,
monkeys also moved seeds to different areas of their home
ranges. Most importantly, they moved seeds between habi-
tats, in particular to the ash plantation, enriching the ash
plantation and catalyzing secondary succession.


We thank the staff of the Santuario de Flora y Fauna
Otin Quimbaya for logistical support at the park. The
study was funded by a grant from the John D. and Cath-
erine T. MacArthur Foundation. Estela Quintero helped
with seed identification and William Vargas and Phillip
Silverstone-Sopkin with plant identification. We thank
Pablo Stevenson for comments on the manuscript.


Aguilar, M. and Rangel, J. 0. 1994. Clima del Parque Re-
gional Natural Ucumari y sectors aledafos. In: Ucumari,
un Caso Tipico de la Diversidad Bidtica Andina, J. 0.
Rangel (ed.), pp.39-56. Corporaci6n Aut6noma Re-
gional de Risaralda, Pereira.
Altmann, J. 1974. Observational study of behavior: Sam-
pling methods. Behaviour 49: 227-277.
Andresen, E. 1999. Seed dispersal by monkeys and the fate
of dispersed seeds in a Peruvian rain forest. Biotropica 31:
Andresen, E. 2002. Primary seed dispersal by red howler
monkeys and the effect of defecation patterns on the fate
of dispersed seeds. Biotropica 34: 261-272.
Ataroff, M. 2001. Venezuela. In: Bosques Nublados del
Neotrdpico, M. Kapelle andA. Brown (eds.), pp.397-442.
Institute Nacional de Biodiversidad, Santo Domingo de
Cavelier, J., Lizcano, D. and Pulido, M. T. 2001. Colom-
bia. In: Bosques Nublados del Neotrdpico, M. Kapelle and
A. Brown (eds.), pp.443-496. Institute Nacional de Bio-
diversidad, Santo Domingo de Heredia.
Defler, T. 2003. Primates de Colombia. Serie de Guias Tropi-
cales de Campo 4, Conservaci6n Internacional, Bogoti.
Durin, S. M. and Kattan, G. H. 2005. A test of the util-
ity of exotic tree plantations for understory birds and
food resources in the Colombian Andes. Biotropica 37:
Estrada, A. and Coates-Estrada, R. 1984. Fruit eating and
seed dispersal by howling monkeys (Alouatta palliata) in
the tropical rain forest of Los Tuxtlas, Mexico. Am. J Pri-
matol. 6: 77-91.
Estrada, A. and Coates-Estrada, R. 1986. Frugivory by
howling monkeys (Alouatta palliata) at Los Tuxtlas,
Mexico: Dispersal and fate of seeds. In: Frugivores and
Seed Dispersal, A. Estrada and T. H. Fleming (eds.),
pp.93-104. Dr W. Junk Publishers, Dordrecht.
Estrada, A. and Coates-Estrada, R. 1988. Tropical rainfor-
est conversion and perspectives in the conservation of
wild primates (Alouatta and Ateles) in Mexico. Am. J Pri-
matol. 19: 315-327.
Estrada, A. and Coates-Estrada, R. 1996. Tropical rainfor-
est fragmentation and wild populations of primates at
Los Tuxtlas, Mexico. Int. J Primatol. 17: 759-783.
Estrada, A., Juan-Solano, S., Ortiz, T. and Coates-Estrada,
R. 1999. Feeding and general activity patterns of a
howler monkey (Alouatta palliata) troop living in a forest
fragment at Los Tuxtlas, Mexico. Am. J. Primatol. 48:
Estrada, A., Mendoza, A., Castellanos, L., Pacheco, R., van
Belle, S., Garcia, Y. and Mufoz, D. 2002. Population of
the black howler monkey (Alouattapigra) in a fragmented
landscape in Palenque, Chiapas, Mexico. Am. J Primatol.
58: 45-55.
Gaulin, S. and Gaulin, C. 1982. Behavioral ecology of
Alouatta seniculus in Andean cloud forest, Colombia. Int.
J Primatol 3: 1-32.

Neotropical Primates 14(2), August 2007

Giraldo, P. 2003. Dieta y dispersi6n de semillas del mono
aullador Alouatta seniculus en el Santuario de Fauna y
Flora Otdn-Quimbaya. B.Sc. Thesis, Universidad del
Valle, Cali, Colombia.
Gentry, A. H. 1992. Diversity and floristic composition of
Andean forests of Peru and adjacent countries: Implica-
tions for their conservation. Mem. Mus. Hist. Nat. U. N.
M. S. M. 21: 11-29.
G6mez-Posada, C., Martinez, J., Giraldo, P. and Kattan,
G. H. Density, habitat use and ranging patterns of red
howler monkeys in a Colombian Andean forest. Neotrop.
Primates 14: 2-10.
G6mez-Posada, C., Roncancio, N. and Hincapi6, P. 2005.
Evaluaci6n de las poblaciones de mono aullador rojo
(Alloutta seniculus: Primates) en fragments de bosque en
el Valle del Cauca. Unpublished report. Corporaci6n Re-
gional del Valle del Cauca, Cali.
Hemingway, C. A. and Overdoff, D. L. 1999. Sampling ef-
fects on food availability estimates: Phenological method,
sample size and species composition. Biotropica 31:
Hernandez-Camacho, J. and Cooper, R. 1976. The non-
human primates of Colombia. In: Neotropical Primates:
Field Studies and Conservation, R. Thorington and P. H.
Helten (eds.), pp.35-69. National Academy of Sciences,
Washington, DC.
Janson, C. H. 1983. Adaptation of fruit morphology to
dispersal agents in a neotropical forest. Science 219:
Julliot, C. 1996a. Seed dispersal by red howling monkeys
(Alouatta seniculus) in the tropical rain forest of French
Guiana. Int. J Primatol. 7: 238-258.
Julliot, C. 1996b. Fruit choice by red howler monkeys in a
tropical rain forest. Am. J Primatol. 40: 261-282.
Julliot, C. and Sabatier, D. 1993. Diet of the red howler
monkey (Alouatta seniculus) in French Guiana. Int. J Pri-
matol. 14: 527-550.
Kattan, G. and Alvarez-L6pez, H. 1996. Preservation and
management of biodiversity in fragmented landscapes in
the Colombian Andes. In: Forest Patches in Tropical Land-
scapes, J. Schelhas and R. Greenberg (eds.), pp. 3-18.
Island Press, Washington, DC.
Londofo, E. 1994. Parque Regional Natural Ucumari: Un
vistazo hist6rico. In: Ucumark Un Caso Tipico de la Di-
versidad Bidtica Andina, J. 0. Rangel (ed.), pp.13-24.
Corporaci6n Aut6noma Regional de Risaralda, Pereira.
Martinez, J. 2003. Densidad poblacional, rango vital y
patr6n de actividad del mono aullador Alouatta senicu-
lus (Primates: Cebidae) en el Santuario de Fauna y Flora
Otin Quimbaya, Risaralda, Colombia. B.Sc. Thesis,
Universidad del Valle, Cali, Colombia.
Milton, K. 1980. The Foraging S- .-. of Howler Monkeys.
A Study in Primate Economics. Columbia University Press,
New York.
Milton, K. 1991. Leaf change and fruit production in six
neotropical Moraceae species. J. Ecol. 79: 1-26.
Milton, K. 1998. Physiological ecology of howlers
(Alouatta): Energetic and digestive considerations and

comparison with the Colobinae. Int. J Primatol. 19:
Palacios, E. and Rodriguez, A. 2001. Ranging pattern and
use of space in a group of red howler monkeys (Alouatta
seniculus) in a southeastern Colombian rainforest. Am. J
Primatol. 55: 233-251.
Rios, M. M., Giraldo, P. and Correa, D. 2004. Gula de
Frutos y Semillas de la Cuenca Media del Rio Otin. Pro-
grama de Colombia de Wildlife Conservation Society
and Fundaci6n EcoAndina, Cali.
Rylands, A. and Keuroghlian, A. 1988. Primate popula-
tions in continuous forest and forest fragments in Central
Amazonia. Acta Amazonica 18(3-4): 291-307.
Robinson, J. G. 1986. Seasonal variation in use of time
space by wedge capuchin monkey, Cebus olivaceus: Im-
plications for foraging theory. Smithsonian Contr. Zool.
431: 1-60.
Schwarzkopf, L. and Rylands, A. 1989. Primate species
richness in relation to habitat structure in Amazonian
rainforest fragments. Biol. Conserv. 48: 1-12.
Silver, S. C., Ostro, L. E. T., Yeager, C. P. and Horwich,
R. 1998. Feeding ecology of the black howler monkey
(Alouatta pigra) in northern Belize. Am. J Primatol. 45:
Stevenson, P. R., Castellanos, M. C., Pizarro, J. C. and Ga-
ravito, M. 2002. Effects of seed dispersal by three ateline
monkey species on seed germination at Tinigua National
Park, Colombia. Int. J. Primatol 23: 1187-1204.
Stevenson, P. R., Quifones, M. and Ahumada, J. 1998.
Annual variation in fruiting pattern using two different
methods in a lowland tropical forest at Tinigua National
Park, Colombia. Biotropica 30: 129-134.
Stevenson, P. R., Quifones, M. and Ahumada, J. 2000.
Influence of fruit availability on ecological overlap among
four Neotropical primates at Tinigua National Park, Co-
lombia. Biotropica 32: 533-544.
Yumoto, T. K., Kimura, K. and Nishimura, A. 1999. Es-
timation of retention times and distances of seed dis-
persed by two monkey species, Alouatta seniculus and
Lagothrix lagotricha, in a Colombian forest. Ecol. Res. 14:
Zar, J. H. 1996. BiostatisticalAnalysis. Third edition. Pren-
tice Hall, New Jersey.

Neotropical Primates 14(2), August 2007

Appendix. Plant species eaten by three troops of howler monkey at Otin-Quimbaya Flora and Fauna Sanctuary, in the Central Andes of
Colombia. Table shows the percentage of each species in the diet and the type of item consumed (F = mature fruits, I = immature fruits,
L = leaves and R = flowers).

Family /Species Troop C Troop D Troop E
% Item % Item % Item
Ficus killipii 58.4 L, F
Ficus cuatrecasana 5.7 L
Ficus tonduzii 9.6 L, F
Ficus aff. 3.0 L
Ficus humboldtii 2.7 L, F 3.7 L
Ficus sp. 1 15.3 L, F
Ficus sp. 2 10.5 F
Ficus sp. 3 5.2 L
Ficus sp. 4 2.2 L
Ficus sp. 5 0.7 L
Ficus sp. 6 0.5 L
Ficus sp. 7 0.5 F
Sorocea sp. 2.3 R
Morus insignis 0.5 L
Pseudolmedia sp. 3.7 L
Moraceae sp. 1 0.8 L
Moraceae sp. 2 4.9 L
Cecropia telealba 9.6 L, F 27.1 L, F 12.6 L, F
Paullinia sp. 22.0 L 3.7 L 11.9 L
Allophylus mollis 1.0 F
Macrolobium colombianum 18.3 L, R 3.7 L, R 4.1 L
Inga aff. oerstediana 2.3 L
Fraxinus chinensis 5.3 I 4.9 I
Garcinia sp. 3.0 F
Chrysochlamys colombiana 4.1 L
Chrysochlamys sp. 0.8 L
Prunus 1.7 L
Dendropanax macrophyllus 0.3 L 0.8 L 0.2 L
Otoba lehmannii 7.9 I
Wettinia kalbreyeri 0.5 I
Casearia sylvestris 1.0 L

continued on next page

Neotropical Primates 14(2), August 2007

Appendix, continued from previous page

Family /Species Troop C Troop D Troop E
% Item % Item % Item
Miconia acuminifera 0.8 F
Solanum sycophanta 2.0 L 0.2 L
Heliocarpus americanus 0.8 L 0.3 L
Sp. 1 3.7 L, F
Sp. 2 1.5 L
Sp. 3 1.0 L
Sp. 4 1.0 L
Sp. 5 0.5 L
Sp. 6 0.5 L
Sp. 7 3.0 L
Sp. 8 0.3 L
Sp. 9 4.3 L
Sp. 10 1.3 L
Sp. 11 1.0 L
Sp. 12 0.7 L
Sp. 13 0.3 L
Sp. 14 0.3 L
Total Frequencies 405 601 637
Total Species 21 27 12

Neotropical Primates 14(2), August 2007 65

Aldo M. Giudice' and Romina Pave2

1 GIBE (Grupo de Investigaciones en Biologia Evolutiva), Departamento de Ecologia, Genitica y Evolucidn, Facultad de
Ciencias Exactas y Naturales, Universidad de Buenos Aires, Pab. II, Lab. 47, 4piso, Ciudad Universitaria (C1428EHA),
Buenos Aires, Argentina, e-mail:
2Estacidn Bioldgica Corrientes. Museo Argentino de Ciencias Naturales (MACN), Av. Angel Gallardo 470, 1405, Buenos Aires,


Captive Cebus paraguayanus were observed to express spontaneously a wide variety of species-specific behaviors in different
zoos from Argentina. The monkeys were kept in old-style wire mesh cages with cement floors, as well as island enclosures.
We collected qualitative data on nine species-specific behaviors, and all the sex-age classes analyzed expressed some of these
behaviors. The results indicated that in spite of living under predictable captive conditions without environmental enrich-
ment, Cebusparaguayanus were able to react to the incidental opportunities that were presented in these environments.

Key words: Cebus paraguayanus, captivity, natural behavior, zoos


Cebus paraguayanus alojados en zool6gicos de Argentina, expresaron de manera espontinea una amplia variedad de con-
ductas especie-especificas. Los ejemplares estaban alojados en recintos de exhibici6n tradicionales, con suelo de cemento y
alambrado perimetral, inclusive en una isla. Se observaron nueve conductas especie-especificas y todas las classes sexo-etarias
expresaron algunos de ellos. Los resultados indican que a pesar de ocupar ambientes altamente predecibles y mon6tonos,
Cebusparaguayanus fue capaz de reaccionar de manera natural hacia oportunidades que brindaban los recintos de exhibici6n
donde se alojaban.

Palabras clave: Cebusparaguayanus, cautiverio, conductas naturales, zool6gicos


Capuchin monkeys, genus Cebus (Platyrrhini: Cebidae),
are omnivorous primates with a diet consisting mainly of
fruit, and in lesser quantity, insects and other plant parts
(Freese and Oppenheimer, 1981). Terborgh (1983) has
pointed out that capuchins use a destructive feeding strat-
egy; by means of their strength, they can exploit food re-
sources not available to other animals. Both in natural and
captive conditions, capuchin monkeys use a wide variety
of manipulative skills. They have been observed to crack
open hard-shelled fruit (i.e.: Izawa and Mizuno, 1977;
Izawa, 1979), hunt different vertebrates and invertebrates
(i.e.: Isawa, 1979; Robinson and Janson, 1987; Rose, 1997;
Ferreira etal., 2002), use tools like stones and branches (i.e:
Anderson, 1990; Visalberghi, 1990; Ottoni and Mannu,
2001; Fragaszy et al., 2004a; Moura and Lee, 2004), tap
scan for presence of hidden animals (Isawa, 1978; Phillips
et al., 2003), and fur rub with different plant and animal
materials (Baker, 1996; Gilbert et al., 1998).

Cebus paraguayanus (also called Cebus apella paraguayanus
Fisher, 1829, or Cebus libidinosus paraguayanus) finds its
southern limit of distribution in Argentina (Mudry, 1990;
Martinez et al., 2002). Its natural habitat is shrinking and,
in addition, commercial traffic is affecting its populations
(Giudice and Mudry, 1995), and is increasing their vul-
nerability to local extirpation (Diaz and Ojeda, 2000). In
Argentina, C. paraguayanus is one of the Neotropical pri-
mates most frequently found in zoos (Giudice, 2000). Zoos
in Argentina can be defined as "old-style zoos," following
Crockett (1998), with conditions that lack furnishings and
bedding materials. Here we use "old style" zoos to refer to
zoos that have enclosures of either old or modern architec-
ture but that lack the physical, biological and social stimuli
necessary to favor the acclimatization of the animals. These
zoos often practice highly deficient management proce-
dures. In extremely predictable enclosures, with little en-
vironmental complexity, animals find it difficult to move
freely, are unable to choose when and what to eat, with
whom to associate with, where to sleep and where to shel-

Neotropical Primates 14(2), August 2007

ter. Under these circumstances, behavioral problems arise
and it seems unlikely that a normal range of species-specific
behaviors will be expressed (IPS, 1993; Mateos Montero,

While several studies of capuchin monkeys have been
conducted in research laboratories (i.e.: Visalberghi and
Trinca, 1989; Visalberghi and Fragaszy, 1995; de Waal,
2000; Brosnan and de Waal, 2003), there is limited pub-
lished information about capuchin behavior in zoos. Ob-
servations of species-specific behaviors recorded for Cebus
in zoos are mostly anecdotal (C. olivaceus: Urbani, 1999;
C apella: Mendes et al., 2000; Urbani and Urquiza-Haas,
2002). The groups of C. paraguayanus studied here in the
"old-style zoos" from Argentina were made up of mon-
keys that were: a) illegally captured and then confiscated,
b) pets highly habituated to humans, c) obtained from in-
stitutional exchange or d) born in zoos. Our study aim was
to examine to what extent captive C paraguayanus express
species-specific non-social behaviors. This study clarifies
how C paraguayanus behave in the captive conditions of
old-style zoos, and it gives insight into the degree of behav-
ioral plasticity found in this Neotropical primate in an en-
vironment with no human-directed enrichment protocol.

Materials and Methods

Study sites, subjects and husbandry
Data were collected in four zoos from Argentina: Jardin
Zool6gico de Buenos Aires (JZBA), Jardin Zool6gico de
La Plata (JZLP), Estaci6n de Cria de Animales Silvestres
de Berazategui, Buenos Aires province (ECAS), and Est-
aci6n Zool6gica Experimental de Santa Fe (EZE). The
enclosures, constructed for public viewing, varied in size
from 34 m3 to 127 m3. They contained wire-meshed walls
and roofs, soil or cement floors, indoor and outdoor rooms,
crossbars, ropes, platforms, dry tree trunks, and some ref-
uges. We also present data from a 100 m2 island enclosure
at JZBA. The composition of the groups of C paraguayanus
in each zoo is summarized in Table 1.

The monkeys were fed once a day around 09:00 with veg-
etables, fleshy fruit, bird eggs, and, except at EZE, meat
and bread. Both food and water were placed on trays on the
floor. None of the groups observed were presented with any
routines of environmental enrichment.

Table 1. Sex-age classes of the studied colonies of Cebus
Zoos Adult Adult Juveniles Infants Total
males females
JZBA 6 5 6 2 19
JZLP 5 3 8
ECAS 1 7 4 12
EZE 5 3 4 12
Total 17 18 14 2 51

Behavioral observations
The observations were collected between 1994 and 2002
during the course of other studies by AMG (1994-1998,
see Giudice, 2000), and RP (1994/2002, see Pave, 2003).
The total observation time was 1232 hours (Table 2). Re-
search projects had similar study goals, and in all the ob-
servation periods, scan and ad libitum sampling were used
(Altmann, 1974) in 30 minute blocks. Data were collected
between 9:00 and 19:00.

In this study, nine species-specific behaviors (as defined in
Table 3) are described and analyzed qualitatively. For hunt-
ing activity, we were able to obtain hourly occurrence rates
across all the study periods, and we calculated these as the
frequency of the hunting events divided by the total obser-
vation time per group.


Hunting behavior
We estimated an hourly frequency of hunting behav-
ior at the different zoos: 0.85 events per hour at JZBA
(n= 164 events), 0.56 events per hour at ECAS (n= 206
events), 0.52 events per hour atJZLP (n= 130 events), and
0.40 events per hour at EZE (n= 109 events). Note that
these frequencies do not take group size into account. More
than 98% of hunting activity was directed toward inverte-
brates, mainly flies. Adults and juveniles of each sex hunted
flying insects and even a three-month old infant was ob-
served in hunting attempts. In the JZBA cage enclosure, an
adult female hunted and consumed a house sparrow (Passer
domesticus). At ECAS, an adult female hunted a rufous
hornero (Furnarius rufus). Also at ECAS, the final phase
of hunting a duck (Fam. Anatidae) was observed. The duck
was grabbed through the wire mesh, and while the duck
body remained outside the cage, each capuchin approached
the prey, took a piece of it and ate its share away from other
individuals. At EZE, the capture of a hornero (Furnarius
sp.) was also observed; the bird got into one of the enclo-
sures and was immediately chased by the alpha male of the
group until it was captured through the wire mesh. The
monkey killed the bird by biting its head and then ate parts
of the head and carcass, which he also shared with an adult
and two juvenile males in the group. At the JZBA cages, an
adult male scattered food outside of the enclosure, appar-
ently in order to attract pigeons (Columba livia) for capture

Table 2. Study periods in four zoos of Argentina.
Study sites and Study periods Observation hours
lodging conditions
JZBA (Cages) 1994-1995 192
JZBA (Island) 1995-1996 50
ECAS (Cages) 1995-1997 366
JZLP (Cages) 1996 250
JZLP (Cages) 1998 100
EZE (Cages) 1999, 2002 274

Neotropical Primates 14(2), August 2007

when they came close enough to reach through the wire
mesh. In addition, a juvenile male unsuccessfully hunted a
kitten. The small cat was handled and pressed against the
mesh, as the juvenile tried to pulled it into the enclosure.
At JZBA island, an adult female handled an approximately
20 cm long aquatic turtle (Trachemys sp.); she repeatedly
hit the turtle against the floor, examined it, bit it, and fi-
nally lost interest and left it.

Tool use
Capuchins were able to attain food out of their reach by
using sticks, and occasionally, leaves and petioles that they
stuck through the mesh wire. At EZE, an adult female took
a stick from the adjoining cage, broke it up in two pieces
and used one of them to acquire a piece of meat from that
cage. The same monkey also used dry leaves that she had
removed from her cage roof to try to acquire meat from the
adjoining cage as well as a peanut from outside the cage
(note that at EZE the monkeys did not receive meat or pea-
nuts in their diet). Similarly, at ECAS, an adult male and
female used branches to reach a slice of bread and grapes
that were outside the cage and out of their reach. At ECAS
and EZE, sticks and leaves were used to explore the soil
or the grass growing outside the enclosures. Monkeys used
sticks to dig in the ground and at the bark of dry trunks, to
explore the holes of trunks or the wall, and to explore the
water depth in the pond at the JZBA island. In the JZBA
cages, an adult female was observed to use a little piece of
straw from a broom to remove splinters from her hand.

Capuchins also used objects to kill animals. At EZE, an
adult male killed a 25 cm long non-venomous snake by
using the brim of a metal container. This monkey struck the

brim of the container several times over the snake's head;
then he bit the head and continued eating the rest of the
snake. Other types of tool use included striking two objects
against one another. At EZE, an adult male used sticks and
stones to crack open fruit of the Pind6 palm (Syagrus sp.).
At JZBA, this activity took place at preferential locations
in two cages: the objects were placed in a deep-set square
metallic grate, which was used as an anvil. At EZE on two
occasions monkeys used sticks like a lever to separate the
wire mesh wall from the cage floor.

This activity includes the handling and consumption of
plant resources accessible from the enclosure. At EZE,
monkeys consumed bermuda grass (Cynodon dactylon),
which grew outside the front of the enclosures and, in some
cases, the capuchins consumed leaves and twigs either with
or without buds of poplar (Populus deltoides), plane tree
(Platanus acerifolia), and tipu tree (Tipuana tipu). A tala
tree (Celtis tala) next to one of the enclosures at ECAS had
branches overhanging the cage roof that were not accessible
to the monkeys. In autumn, monkeys were seen eating fruit
from this tree when it fell into the enclosure. In one cage
at JZBA in the early spring, some foliage from an ombu
tree (Phytolacca dioica) fell on the roof of the enclosure,
and monkeys consumed the inflorescences. Once, keepers
at EZE put some soil outside the cages and near the wire
mesh, and adults and juveniles consumed grass leaves and
roots present in the new soil.

Manipulative events
A frequent observation was that the capuchins hit apples,
potatoes, walnuts, or almonds several times against some

Table 3. Cebusparaguayanus behaviors recorded in the study.
Behavioral units Definition
Hunting behavior To obtain animal products by means of an active search process; prey is caught with the hunter's limbs and mouth
and immediately consumed.
Tool use To use an unattached environmental object to alter more efficiently the form, position or condition of another
object, another organism or the user itself (Beck, 1980). A tool is employed as a functional extension of the animal's
body to bridge between itself and its target (Visalberghi, 1990).
Foraging To direct behavior toward a potential food source not included in the zoo diet. Actions include directed search,
processing and consuming, I .. .. 1990).
Manipulative events To use hands and mouth to alter the form, position or condition of another object, which involved two or more
coordinated acts. For example, digging in the ground as a necessary step toward acquiring a hidden piece of food
was registered as a manipulative event, but picking up a piece of food lying on the ground was not (Jalles-Filho,
Tap scanning To give soft hits to a surface with the fingertips of one hand. Tapping lightly and rapidly (Phillips et al., 2003).
Fur rubbing To apply a foreign substance over the body pelage using hands, feet and tail, in a highly energetic way (Baker,
Drinking behavior To introduce hand and arm in the water source, soak their hairs and then lick the water that drains off the arm or
hand. Dipping-and-licking technique (Wrangham, 1981).
Behavioral To rest keeping arms and legs flexed and close to the body while the back is bent, to conserve the heat during
thermoregulation periods of low ambient temperature (curled posture). And to rest extending the back and all members, to dissipate
heat under hot conditions (stretched posture) (Bicca-Marques and Calegaro-Marques, 1998).
Vertical flight response To climb suddenly to the enclosure roof and to remain expectant (Giudice, 2000). In the wild, capuchins react to
aerial predators moving upwards in the trees and scanning (Rose et al., 2003).

Neotropical Primates 14(2), August 2007

hard surface, like the floor or the mesh, in order to eat the
bits stuck to the surface or to the food. At EZE, another
common food processing strategy was that adult and ju-
venile males rubbed dry leaves, grass, small sticks, food, or
other items between their hands or against the enclosure

Tap scanning
This activity is used by the capuchins in natural conditions
to detect the presence of vertebrates or invertebrates hidden
inside tree trunks. At EZE three adults and one juvenile
were observed to tap scan on a dry tree trunk several times.
They were also observed to tap scan on some sticks. In some
cases, after tap scanning, they bit the tree bark. Also, these
animals explored the holes of dry trunks with their fingers
and in some cases, they employed a technique observed in
natural conditions; they ripped up the bark of the tree with
their mouths and hands and then licked the surface.

Fur rubbing
At EZE, an adult male took some water with bleach from
the recently washed drinking dish and used his hand to
rub it over his back. On another occasion, an adult male
smelled the enclosure floor, rubbed his cheek on the same
place, and then was imitated by one of the juveniles. At
JZLP, two adult males were observed to rub onion over
their bodies.

Drinking behavior
At ECAS a keeper left a bucket with water outside the en-
closure, next to the mesh. The monkeys could not reach
the bucket with their mouths, but they introduced their
hands into the water, and then drank the water dripping
from their arms.

Behavioral thermoregulation
In each of the zoos, energy-conserving or energy-dissipat-
ing postures were observed. In winter, animals rested using
the curled posture in the sunny places of the enclosure. In
summer, monkeys adopted the stretched posture leaning
their ventral region on the floor or on thick crossbars, espe-
cially in shady places. These behaviors were observed in all
the sex-age classes studied.

Vertical flight response
At EZE, in response to calls emitted by a group member
or by monkeys from adjoining cages, the capuchin group
would suddenly climb to the upper part of the enclosure,
or run into the indoor room. They also reacted daily in this
way in response to the presence of a zookeeper near the
enclosures. This vertical flight response was observed upon
detection of southern crested caracaras (Polyborus plancus)
flying overhead, after vocalizations emitted by dogs and
captive flamingos (Phoenicopterus sp.), and on two occa-
sions in response to spider monkeys (Ateles paniscus) emit-
ting loud vocalizations in adjoining cages. At ECAS verti-
cal flight behavior was observed at least once a day. A call
made by a capuchin group member was the most frequent

stimulus that triggered this response. The individual that
made the call would climb rapidly up the mesh wire to the
cage roof, and this behavior was immediately imitated by
other group members. In all cases, after a few seconds on
the roof or in the indoor room, the monkeys returned to
the floor.


In the four study populations of C. paraguayanus kept in
old-style zoo enclosures, we observed the normal expres-
sion of nine species-specific behaviors, and all the sex-age
classes analyzed expressed at least some of these behaviors.
Monkeys hunted flying insects and different vertebrates,
mainly birds. The hunting of invertebrates and vertebrates
has been recorded often in capuchin monkeys in natural
and semi-free ranging conditions (Isawa, 1979; Freese and
Oppenheimer, 1981; Janson, 1985; Brown et al., 1986;
Robinson and Janson, 1987; Rose, 1997; Ferreira et al.,
2002; Sampaio and Ferrari, 2005). In one case, in our study
an adult male scattered food outside the cage, apparently to
attract pigeons as potential prey. This may suggest a certain
cognitive ability, and is in line with similar observations in
other captive capuchins, documented to attract ducklings
and fishes (McGrew and Marchant, 1997; Mendes et al.,
2000). This observation also highlights the ability of capu-
chins to take great advantage of the opportunities that are
presented in the zoos and to develop alternative foraging
strategies to meet them.

Tool use was observed in a variety of contexts. Branches
and leaves were spontaneously used like arm extensions to
obtain food that was out of reach, or to explore distant
or hidden surfaces. As observed in wild Cebus (Moura and
Lee, 2004), individuals in this study used objects to dig in
the ground. An adult female used a small piece of straw
from a broom to remove splinters from her hand. Monkeys
also cracked fruits with sticks and stones using the same ac-
tions as described by Izawa and Mizuno (1977). An adult
male used the brim of a metal container to kill a non-ven-
omous snake. Although Boinski (1988) reported the use
of a branch by wild Cebus capucinus to attack a venomous
snake, the case observed here with a potential prey item
can be considered a prey processing technique equivalent
to the use of objects as hammers to crack open oysters (see
Fernandez, 1991), and to the cases described above with
fruits and reported in wild Cebus (i.e.: Izawa and Mizuno,
1977; Anderson, 1990; Moura and Lee, 2004). Our mon-
keys often hit different foods against hard surfaces and
then ate the bits; this food processing technique has been
observed in captive (Urbani and Urquiza-Hass, 2002) and
wild Cebus (Moura and Lee, 2004). In one zoo (JZBA), in-
dividuals used a square metallic grate as an anvil to facilitate
the cracking open of fruits. This anvil technique has been
reported in several Cebus studies (i.e.: Ottoni and Mannu,
2001; Fragaszy et al., 2004a). Perhaps the most interesting
tool use was by an adult female, who took a stick from the
adjoining cage, broke it up in two pieces, and used one of

Neotropical Primates 14(2), August 2007

them to obtain meat from that cage. The sequence is inter-
esting for various reasons. First, it shows the ability of ca-
puchins to solve problems. It also shows the capacity of the
female to modify the stick before using it like a tool. Simi-
lar actions have been documented in Cebus apella libidino-
sus (Moura and Lee, 2004) and Pan troglodytes (Boesch and
Boesch, 1990; Pruetz and Bertolani, 2007).

Captive capuchins employed tap-scanning to detect
the presence of hidden animals inside trunks and sticks.
They used their fingers to explore inside holes of differ-
ent substrates and in some occasions they ripped up bark
to obtain something or just to lick the surface. All these
species-specific behaviors show that capuchins continue to
use sensory cues to guide their search for prey and display
the manual dexterity and extractive foraging characteristic
of capuchin monkeys (Izawa, 1978, 1979; Phillips et al.,
2003), even when in captive conditions with minimal en-
richment. The captive monkeys displayed fur rubbing and
dipping and licking behaviors. Monkeys in this study fur
rubbed with a water-bleach solution and with onion over
their bodies. Fur rubbing using several plant and animal ma-
terials has been observed in wild and captive Cebus (Baker,
1996; Gilbert et al., 1998), and it has also been reported
in owl monkeys, Aotus (Zito et al., 2003). Dipping and
licking behavior allows for access to drinking water even
when it cannot be directly obtained by the mouth, and it
has been observed in Old World Primates (Cercopithecus
aethiops: Wrangham, 1981; Procolobus badius temminckii:
Starin, 2002) as well as in wild and captive Neotropical pri-
mates (Cebus capucinus: Freese, 1978; C olivaceus: Urbani,
1999; Alouatta caraya: Bicca-Marques, 1992; Giudice and
Mudry, 2000; Saimiri sp.: Baldwin and Baldwin, 1981). In
primates behavioral thermoregulation is important to con-
serve heat during cold periods and to dissipate heat in hot
periods (Bicca-Marques and Calegaro-Marques, 1998). In
winter, animals rested huddling in sunny places using the
curled posture. In summer, C. paraguayanus rested using
the stretched posture especially in shady places (Paterson,
1986). These strategies have been observed in wild capu-
chins (Fragaszy et al, 2004b) and other Neotropical pri-
mates (Alouatta caraya: Zunino, 1989; Bicca-Marques and
Calegaro-Marques, 1998; A. palliata: Mufioz et al, 2002).

We also observed the vertical flight response in the cap-
tive groups. In wild conditions, capuchins produce differ-
ent alarm calls toward aerial and terrestrial predators and
they react differently to each type of call (Fedigan, 1993;
Di Bitetti, 2001; Rose et al., 2003). The same antipreda-
tor responses were expressed daily by our C. paraguayanus
groups. They moved upwards or downwards in the cages
in response to the detection of caregivers or flying raptors
or in response to vocalizations of dogs and flamingos. The
only species of raptor that the individuals detected and gave
alarm calls for was a southern crested caracara (Polyborus
plancus). Rose et al. (2003) reported that Cebus capucinus
in Costa Rica often alarm-called for Polyborus plancus but
they did not observe predation attempts by this raptor.

Interestingly, vertical flight response was observed in capu-
chins after loud vocalizations emitted by spider monkeys
(Ateles paniscus) kept in adjoining cages. Other interac-
tions observed between Cebus and Ateles were initiated by
the capuchins and included both affiliation, in the form
of gentle touching (n= 2 events), and aggression, includ-
ing tail and limb pulling, chases over the wire mesh, and
pinches (n= 9 events). Although interactions have been
observed between Cebus capucinus and Ateles .,T .,, in
Costa Rica (Rose et al., 2003), this is the first report of
of Cebus responding to Ateles alarm calls. Inter-specific re-
sponses to alarm calls do exist in other primates (Saguinus
labiatus / S. fuscicollis: Buchanan-Smith, 1990; Saguinus
mystax / S. fuscicollis: Heymann, 1990; Cercopithecus mitis /
C ascanius: Cords, 1990).

While the present study is about non-social behaviors, two
social behaviors observed in the study deserve mention.
First, food sharing occurred subsequent to the hunting of
a bird (Furnarius sp.). This cooperative behavior has been
reported in free-ranging conditions for C. capucinus (Perry
and Rose, 1994; Rose, 1997) and in semi-free-ranging
conditions for C. apella (Ferreira et al., 2002). In the ob-
served case the hunter, the alpha male, shared part of the
prey by allowing other group members to take scraps that
fell on the floor. This sort of food transfer is the most
common type observed by Rose (1997) and Ferreira et al.
(2002). Second, the attempt at hunting by a three month
old infant may have been a result of local or object enhance-
ment, as the infant had the opportunity to watch adults
hunting for the same prey items. As previously noted in
other studies of capuchins (Perry and Rose, 1994; Ottoni
and Manu, 2001), through the observations of older ani-
mals, the infants learn by trial and error to be more profi-
cient at hunting.

Our results confirm the behavioral plasticity of the capu-
chin monkeys, their flexible foraging strategies, and their
ability to explore and manipulate the environment, includ-
ing through the use of tools. They displayed a high capacity
to take advantage of the opportunities that were presented
often inadvertently in the highly predictable conditions
studied. Taking into consideration that modern zoos should
be conservation agents not only of genetic diversity but also
of the behavioral diversity (Rabin, 2003), zoos should be
encouraged to increase the opportunities to stimulate the
expression of species-specific behaviors in capuchin mon-
keys and other species.


Part of this study was supported by a Consejo Nacional de
Investigaciones Cientificas yT&cnicas (CONICET) fellow-
ship to Aldo M. Giudice. We are grateful to the veterinar-
ian and caretakers from Jardin Zool6gico de Buenos Aires,
Jardin Zool6gico de La Plata, Estaci6n de Cria de Animales
Silvestres, and Estaci6n Zool6gica Experimental de Santa
Fe, for their help and technical support.

Neotropical Primates 14(2), August 2007


Anderson, J. R. 1990. Use of objects as hammers to open
nuts by capuchin monkeys (Cebus apella). Folia Primatol.
54: 138-145.
Altmann, J. 1974. Observational study of behavior: Sam-
pling methods. Behaviour 49: 227-267.
Baker, M. 1996. Fur rubbing: Use of medicinal plants by
capuchin monkeys (Cebus capucinus). Am. J Primatol.
38: 263-270.
Baldwin, J. and Baldwin, J. 1981. The squirrel monkeys,
genus Saimiri. In: Ecology and Behavior of Neotropical
Primates, A. F. Coimbra-Filho and R. A. Mittermeier
(eds.), pp. 277-330. Rio de Janeiro, Academia Brasilera
de Ciencias.
Beck, B.B. 1980. Animal Tool Behavior. Garland STPM
Press, New York.
Bicca-Marques, J. C. 1992. Drinking behavior in the black
howler monkey (Alouatta caraya). Folia Primatol. 58(2):
Bicca-Marques, J. C. and Calegaro-Marques, C. 1998.
Behavioral thermoregulation in a sexually and develop-
mentally dichromatic Neotropical primate, the black-
and-gold howling monkey (Alouatta caraya). Am. J Phys.
Anthropol. 106: 533-546.
Boesch, C. and Boesch, H. 1990. Tool use and tool making
in wild chimpanzees. Folia Primatol. 54: 86-90.
Boinski, S. 1988. Use of a club by wild white-faced capu-
chin (Cebus capucinus) to attack a venomous snake (Both-
rops asper). Am. J Primatol. 14: 177-179.
Brosnan, S. E and de Waal, E B. M. 2003. Monkeys reject
unequal pay. Nature 425: 297-299.
Brown, A., Chalukian, S., Malmierca, L. and Colillas, 0.
1986. Habitat structure and feeding behavior of Cebus
apella (Cebidae) in "El Rey" National Park, Argentina.
In: Current Perspectives in Primate Social Dynamics, D. M.
Taub and F. A. King (eds.), pp. 137-151. Van Nostrand
Reinhold, New York.
Buchanan-Smith, H. M. 1990. Polyspecific association of
two tamarin species, Saguinus labiatus and Saguinus fusci-
collis, in Bolivia. Am. J Primatol. 22: 205-214.
Cords, M. 1990. Mixed-species association of East African
guenons: General patterns or specific examples? Am. J.
Primatol. 21: 101-114.
Crockett, C. M. 1998. Psychological well-being of captive
nonhuman primates: Lessons from laboratory studies. In:
Environmental Enrichment for Captive Animals, D. Shep-
herdson, J. Mellen and M. Hutchins (eds.), pp. 129-152.
Smithsonian Institution Press, Washington, DC.
Diaz, G. and Ojeda, R. 2000. Libro Rojo de los Mamiferos
Amenazados de Argentina. SAREM, Buenos Aires.
Di Bitetti, M. S. 2001. Food associated calls in the tufted
capuchin monkey (Cebus apella). Docoral thesis, State
University of New York, Stony Brook, New York.
Fedigan, L. 1993. Sex differences and intersexual relations
in adult white-faced capuchins (Cebus capucinus). Int. J.
Primatol. 14(6): 853-877.

Fernandez, M. E. B. 1991. Tool use and predation of
oysters (Crassostrea rhizophorae) by the tufted capuchin,
Cebus apella apella, in brackish-water mangrove swamp.
Primates 32(4): 529-531.
Ferreira, R., Resende, B., Mannu, M., Ottoni, E. and
Izar, P. 2002. Bird predation and prey-transfer in brown
capuchin monkeys (Cebus apella). Neotrop. Primates
10 (2): 84-89.
Fragaszy, D. M. 1990. Sex and age differences in the orga-
nization of behavior in wedge-capped capuchins, Cebus
olivaceus. Behav. Ecol. 1(1): 81-94.
Fragaszy, D. M., Izar, P., Visalberghi, E., Ottoni, E. B.
and De Oliveira, M. G. 2004a. Wild capuchin monkeys
use anvils and stone pounding tools. 27th Meeting of the
American Society of Primatologists, Madison, WI.
Fragaszy, D. M., Visalberghi, E. and Fedigan, L. M. 2004b.
The Complete Capuchin: The Biology of the Genus Cebus.
Cambridge University Press, Cambridge.
Freese, C. H. 1978. The behavior of white-faced capuchins
(Cebus capucinus) at a dry season waterhole. Primates 19:
Freese, C. H. and Oppenheimer, J. R. 1981. The capu-
chin monkeys, genus Cebus. In: Ecology and Behavior
of Neotropical Primates, A. F. Coimbra-Filho and R. A.
Mittermeier (eds.), pp. 301-385. Academia Brasileira de
Ciencias, Rio de Janeiro.
Gilbert, T. M., Brown, D. A. and Boysen, S.T. 1998. Social
effects on anointing behavior in capuchins (Cebus apella).
Am. J Primatol. 45: 182.
Giudice, A. M. 2000. Anilisis del comportamiento de
Cebus apella en jardines zool6gicos. Doctoral thesis. Uni-
versity of Buenos Aires, Buenos Aires, Argentina.
Giudice, A. M. and Mudry, M. D. 1995. Mono cai (Cebus
apella) en cautiverio: Composici6n de grupos. Bol. Pri-
matol. Latinoam. 5(1): 29-33.
Giudice, A. M. and Mudry, M. D. 2000. Drinking be-
havior in the black howler monkey (Alouatta caraya).
Zoocriaderos 3(1): 11-19.
Heymann, E. W. 1990. Reactions of wild tamarins Sagui-
nus mystax and Saguinus fuscicollis to avian predators. Int.
J Primatol. 11: 327-337.
International Primatological Society, 1993. International
Guidelines: IPS Code of Practice: 1. Housing and envi-
ronmental enrichment. Prim. Rep. 35(1): 8-16.
Izawa, K. 1978. Frog-eating behavior of wild black-capped
capuchin (Cebus apella). Primates 19(4): 633-642.
Izawa, K. 1979. Foods and feeding behavior of wild black-
capped capuchin (Cebus apella). Primates 20(1): 57-76.
Izawa, K. and Mizuno, A. 1977. Palm-fruit cracking be-
havior of wild black-capped capuchin (Cebus apella). Pri-
mates 18(4): 773-792.
Jalles-Filho, E. 1995. Manipulative propensity and tool
use in capuchin monkeys. Current Anthropol. 36(4):
Janson, C. H. 1985. Aggressive competition and individu-
al food consumption in wild brown capuchin monkeys.
Behav. Ecol. Sociobiol. 18: 125-138.

Neotropical Primates 14(2), August 2007

Martinez, R. A., Giudice, A. M., Szapkievich, V. B., As-
cunce, M., Nieves, M., Zunino, G. and Mudry, M. D.
2002. Parameters modeling speciogenic processes in
Cebus apella (Primate: Platyrrhini) from Argentina. Mas-
tozool. Neotrop. 9(2): 171-186.
Mateos Montero, C. 1994. El bienestar animal, una evalu-
aci6n cientifica del sufrimiento animal. In Etologia: In-
troduccidn a la Ciencia del Comportamiento, J. Carranza
(ed.), pp.493-527. Publicaciones de la Universidad de
Extremadura. Ciceres, Espana.
Mendes, E D., Martins, L. B. R., Pereira, J. A. and Mar-
quezan, R. E 2000. Fishing with a bait: A note on be-
havioural flexibility in Cebus apella. Folia Primatol. 71:
McGrew, W. C. and Marchant, L. F. 1997. Using the tools
at hand: Manual laterality and elementary technology in
Cebus spp. and Pan. Int. J Primatol. 18: 787-810.
Moura, A. C. de A. and Lee, P. C. 2004. Capuchin stone
tool use in Caatinga Dry Forest. Science 306: 1909.
Mudry, M. D. 1990. Cytogenetic variability within and
across populations of Cebus apella in Argentina. Folia
Primatol. 54: 206-216
Mufioz, D., Garcia del Valle, Y., Franco, B., Estrada, A.
and Magafia, A. M. 2002. Estudio del patr6n de activi-
dad general de monos aulladores (Alouatta palliata) en
el Parque Yumki, Tabasco, Mexico. Neotrop. Primates 10
(1): 11-17.
Ottoni, E. B. and Mannu, M. 2001. Semifree-ranging
tufted capuchins (Cebus apella) spontaneously use tools
to crack open nuts. Int. J. Primatol. 22 (3): 347-358.
Paterson, J. D. 1986. Shape as a factor in primate thermo-
regulation. In: Current Perspective in Primate Social Dy-
manics, D. M. Taub and F. A. King (eds.), pp.228-242.
Van Nostrand Reinhold, New York.
Pave, R. 2003. Estudio del comportamiento de Cebus apella
en condiciones de cautiverio. Bachelor's thesis, University
of Litoral, Santa Fe, Argentina.
Perry, S. and Rose, L. M. 1994. Begging and transfer of
coati meat by white-faced capuchin monkeys, Cebus ca-
pucinus. Primates 35: 409-415.
Phillips, K. A., Grafton, B. W. and Haas, M. E. 2003. Tap-
scanning for invertebrates by capuchins (Cebus apella).
Folia Primatol. 74: 162-164.
Pruetz, J. D. and Bertolani, P. 2007. Savanna chimpanzees,
Pan troglodytes verus, hunt with tools. Current Biol. 17:
Rabin, L. A. 2003. Maintaining behavioral diversity in
captivity for conservation: Natural behaviour manage-
ment. Anim. 'j .- 12: 85-94.
Rose, L. 1997. Vertebrate predation and food-sharing in
Cebus and Pan. Int. J Primatol. 18: 727-765.
Robinson, J. and Janson, C. H. 1987. Capuchins, squirrel
monkeys and atelines: Socioecological convergence with
Old World primates. In: Primate Societies, B. B. Smuts,
D. L. Cheney, R. M. Seyfarth, R. W. Wrangham and
T. T. Struhsaker (eds.), pp.69-82. University of Chicago
Press, Chicago.

Sampaio, D. T. and Ferrari, S. F. 2005. Predation of an
infant titi monkey (Callicebus moloch) by a tufted capu-
chin (Cebus apella). Folia Primatol. 76: 113-115.
Starin, E. D. 2002. Drinking observations in Temminck's
red colobus. Folia Primatol. 73: 137-139.
Rose, L. M. 1997. Vertebrate predation and food-sharing
in Cebus and Pan. Int. J. Primatol. 18: 727-765.
Rose, L. M., Perry, S., Panger, M. A., Jack, K., Manson,
J. H., Gros-Louis, J., Mackinnon, K. C. and Vogel, E.
2003. Interspecific interactions between Cebus capucinus
and other species: Data from three Costa Rican sites. Int.
J Primatol. 24(4): 759-796.
Terborgh, J. 1983. Five New World Primates: A Study
in Comparative Ecology. Princeton University Press,
Urbani, B. 1999. Spontaneous use of tools by wedge-ca-
pped capuchin monkeys (Cebus olivaceus). Folia Primatol.
70: 172-174.
Urbani, B. and Urquiza-Haas, T. 2002. Corte innovador
de frutas por el mono capuchino pardo (Cebus apella) en
cautiverio. Lab. Prim. Newsl. 41(4).
Visalberghi, E. 1990. Tool use in Cebus. Folia Primatol. 54:
Visalberghi, E. and Trinca, L. 1989. Tool use in capuchin
monkeys: Distinguishing between performing and un-
derstanding. Primates 30: 511-521.
Visalberghi, E. and Fragaszy, D. 1995. The behaviour of
capuchin monkeys, Cebus apella, with novel food: The
role of social context. Anim. Behav. 49: 1089-1095.
de Waal, F. B. M. 2000. Attitudinal reciprocity in food
sharing among brown capuchin monkeys. Anim. Behav.
60: 253-261.
Wrangham, R. W. 1981. Drinking competition in vervet
monkeys. Anim. Behav. 29: 904-910.
Zito, M., Evans, S. and Weldon, P. J. 2003. Owl monkeys
(Aotus spp.) self-anoint with plants and millipedes. Folia
Primatol. 74: 159-161.
Zunino, G. E. 1989. Habitat, dieta y actividad del mono
aullador negro (Alouatta caraya) en el noreste de Argen-
tina. Bol. Primatol. Latinoam. 1(1): 74-97.

Neotropical Primates 14(2), August 2007


Claudia R. Silva

Os primatas do genero Cebus Erxleben 1777 possuem pro-
nunciada destreza manual, c6rebro grande com quantidade
elevada de circunvolug6es, alta sociabilidade, forte capaci-
dade de manipulagao e dieta onivora (Defler, 1979; Freese
e Oppenheimer, 1981; Auricchio, 1995; Sussman, 2000;
Defler, 2004). Estas caracteristicas presumivelmente garan-
tem a estes primatas a capacidade de explorer o ambiente
de forma diferenciada. Macacos-prego sao generalistas e fle-
xiveis no uso de recursos alimentares atrav5s de grande va-
riedade de habitats (Fragaszy et al., 1990). Sua plasticidade
evolutiva 6 elevada, tornando-os aptos a aproveitar oportuni-
dades ecol6gicas diferentes sem apresentar adaptac6es muito
especificas a ambientes particulares (Auricchio, 1995). Cebus
apella (sensu Silva Junior, 2002 = Cebus apella apella sensu
Hill, 1960) possui uma ampla distribuigao geogrifica, ocor-
rendo na Amaz6nia centro oriental, em pelo menos cinco
pauses, Brasil, Venezuela, Suriname e as Guianas (Silva Jdnior,
2002). Esta esp6cie 6 amplamente distribuida no Estado do
Amapi, ocorrendo desde os mangues altos da costa litoranea
at6 as florestas de terra firme. Os macacos-prego tem hibi-
tos generalistas, com dieta baseada principalmente em frutas
e artr6podes (Terborgh, 1983; Robinson e Janson, 1987).
Flores, brotos, ovos, aves, pequenos anfibios, rdpteis e ma-
miferos tambdm sao consumidos, embora menos freqiien-
temente (Freese e Oppenheimer, 1981). Os macacos-prego
sao conhecidamente de hibitos diurnos, no entanto Rimoli
(2001) registrou deslocamento noturno entire Cebus nigritus
enquanto acompanhava o grupo at6 sua area de dormir. Se-
gundo o autor, os macacos se mantiveram em atividade por
cerca de uma hora ap6s anoitecer.

Esse trabalho relata observagoes de um bando de macaco-
prego (Cebus apella) se alimentando a noite, no Parque Na-
cional Montanhas do Tumucumaque, Estado do Amapai,
Brasil. 0 Parque do Tumucumaque localiza-se na regiao no-
roeste do Estado do Amapi. Possui uma area de 3.867.000
ha de floresta de terra firme, bem conservados. 0 clima 6
considerado tropical dmido, com uma curta estagao seca,
culminando nos meses de outubro e novembro. Cinco ex-
pedigoes, de cerca de 20 dias de duracao, foram efetuadas
ao Parque Tumucumaque para inventariar a fauna de verte-
brados, crusticeos e plants superiores pelo Projeto Inven-
tArios Biol6gicos no Corredor de Biodiversidade do Amapi.
Na primeira delas, no period de 11 a 29 de setembro de
2004, foi explorada uma area situada na confluencia dos
rios Amapari e Anacuf (5229'32"W, 0135'45"N, Fig. 1).

Nesta expedigao durante o censo de mamiferos, um bando
de macacos-prego (C apella) foi observado em atividade,

vocalizando e se deslocando, na copa de um piquii-amarelo
(Caryocar villosum), apesar da noite escura. 0 piquid 6 uma
irvore caracteristica da floresta de terra firme, ocorre em
toda a Amaz6nia, principalmente na regiao do estuario
do rio Amazonas (Lorenzi, 1992). Esta irvore alcanga ate
45 metros de altura, pode ser vista compondo o dossel ou
como emergente, e floresce durante os meses de agosto, se-
tembro e outubro (Lorenzi, 1992; Ribeiro, 1999). Os ma-
cacos foram observados na mesma arvore por duas noites
consecutivas, 17 e 18 de setembro de 2004. Corn o auxilio
de um bin6culo (Bushmaster 10 x 50) e de uma lantern
de foco amplo, foi possivel verificar que na copa do piquii,
a aproximadamente 30 m de altura, os macacos se alimen-
tando de flores. As observagoes se deram no period das
20:00 as 24:00 do primeiro dia, e das 19:00 as 24:00 do dia
seguinte. Na primeira noite de observagao, foi possivel re-
gistrar apenas dois individuos se alimentando. As observa-
goes se iniciaram Xs 20:00, quando um juvenile se alimentou
por 15 minutes e depois um adulto, por aproximadamente
20 minutes. Embora o bando fosse maior, nao foi possi-
vel verificar o tamanho do bando nem quantos individuos
estavam se alimentando. No entanto, os macacos se man-
tiveram em atividade, vocalizando e se movimentando no
alto da arvore, at6 as 21:45. N6s permanecemos embaixo
da irvore, no entanto nao foi verificada atividade alguma
at6 as 24:00 quando abandonamos as observagoes. Na se-
gunda noite, o primeiro registro efetuado foi de um juvenile
as 20:25, se alimentando por 15 minutes e, em seguida, foi
registrado um adulto se alimentando por aproximadamen-
te 25 minutes. Nesta noite, os animals apresentaram menos
atividade (movimentagao e vocalizagao) do que na anterior,
provavelmente devido a presenga dos observadores desde o
incio da noite.

Estudos sobre padrao de uso espacial de primatas sao con-
sistentes com a hip6tese de que macacos de vida livre co-
nhecem a localizagao de suas arvores de alimento e se deslo-
cam eficientemente entire elas (Sigg e Stolba, 1981; Garber,
1989). Di Bitetti (2000) estudando Areas de dormir de
Cebus nigritus, encontrou padres na escolha das irvores
onde passariam a noite, sendo irvores de grande porte e
altas em floresta madura as escolhidas. Os macacos podem
distribuir-se em irvores proximas nao estando todo o bando
em uma mesma irvore. Provavelmente, os macacos obser-
vados no Parque do Tumucumaque se deslocaram para a
irvore de alimentagao durante o dia e permaneceram ate
o anoitecer. Muitos estudos com primatas tem mostrado
que o olfato pode ser considerado na escolha do alimen-
to (Laska e Hudson, 1993; Ueno, 1994a, b; Laska et al.,
1996; Hubener e Laska, 1998; Drapier et al., 2002). Uma
caracteristica das flores de piquidi o forte odor, o que pode
auxiliar a sua localizagao. E provivel que durante a noite
os macacos estivessem utilizando do olfato para a locali-
zagao das flores na copa da irvore. As flores do piquid sao
utilizadas na alimentagao de virias esp6cies de mamiferos.
As cameras-trap utilizadas durante os inventirios realizados
no Corredor de Biodiversidade do Amapi registraram tatu-
galinha (Dasypus novemcinctus) e veado-mateiro (Mazama

Neotropical Primates 14(2), August 2007

Figura 1. Local onde foi realizada a 1- expedigao de inventirios biol6gicos ao Parque Tumucumaque corn a localizagao do ponto de
observagao do bando de macaco-prego (C. apella).

americana) se alimentando das flores de piquid (Caryocar
villosum) durante o dia.

Durante a expedigao ao rio Amapari cinco esp6cies de
primatas foram encontradas em simpatria, C. apella, Sai-
miri sciureus, Alouatta seniculus, Ateles paniscus e Saguinus
midas e duas outras esp6cies foram relatadas por barquei-
ros, eximios conhecedores da regiao: Chiropotes satanas
e Pithecia pithecia. Sendo Ateles paniscus, Cebus apella e
Saimiri sciureus as esp6cies mais comumente encontradas
durante as procuras ativas desenvolvidas na irea. Em duas
outras oportunidades, bandos de Saimiri sciureus foram
registrados em atividade noturna (vocalizando/deslocan-
do). No entanto, devido is dificuldades de observagao de
primatas durante a noite, nao foi possivel verificar o com-
portamento dos grupos atraves de observagao de longa


Este trabalho contou corn o suporte da Conservagao Inter-
nacional do Brasil e do Instituto de Pesquisas Cientificas
do Estado do Amapi e teve apoio do IBAMA-AP. Agradego
ao Dr. Jos6 da Silva Junior e a um revisor an6nimo pelos
comentirios no manuscrito. Agradego tambdm a Domin-
gos Ramos da Costa Vidal por ter me auxiliado nas ob-

servagoes, ao Dr. Enrico Bernard e Ana Carolina Moreira
Martins por terem encontrado o bando durante coleta de
morcegos. Cliudia Funi produziu o mapa.

Claudia R. Silva, Instituto de Pesquisas Cientificas e Tec-
nol6gicas do Estado do Amapi (IEPA), e-mail: ap@gmail.com>.


Auricchio, P. 1995. Primatas do Brasil. Sao Paulo, Terra
Brasilis, 168p.
Defler, T. R. 1979. On the ecology and behavior of Cebus
,bifbol's in eastern Colombia. Primates 20: 475-502.
Defler, T. R. 2004. Primates of Colombia. Bogoti, Conser-
vation International, 550pp.
Drapier, M. e Thierry, B. 2002. Social transmission of
feeding techniques in Tonkean macaques? Int. J. Prima-
tol. 23: 105-122.
Di Bitetti, M., Vidal, E. M. L., Baldovino, M. C. e Be-
nesovsky, V. 2000. Sleeping site preferences in tufted
capuchin monkeys (Cebus apella nigritus). Am. J. Prima-
tol. 50:257-274.
Fragaszy, D. M., Visalberghi, E. e Robinson, J. G. 1990.
Variability and adaptability in the genus Cebus. Folia Pri-
matol. 54: 114-118.

L- I d.Ob-.9A. 500 (1 $00 1000 1500 2000 2500GSW01S r~r~~O

GwgraphitCodinae Symm
acs WOS1994 Feniro' 2007

Neotropical Primates 14(2), August 2007

Freese, C. H. e Oppenheimer, J. R. 1981. The capuchin
monkeys, genus Cebus, p. 331-389.
Em: A. E Coimbra-Filho e R.A. Mittermeier (Ed.). Ecolo-
gy and behavior of Neotropical primates. Rio de Janeiro,
Academia Brasileira de Ciencias, 496pp.
Garber, P. A. 1989. The role of spatial memory in primate
foraging patterns: Saguinus mystax and Saguinus fuscicol-
lis. Am. J. Primatol. 19: 203-216.
Hubener, F e Laska, M. 1998. Assessing olfactory per-
formance in an old world primate: Macaca nemestrina.
Physiol. Behav. 64: 521-527.
Hill, 0. C. 1960. Primate comparative anatomy and tax-
onomy. Vol. IV Cebidae, Part A. Edinburgh University
Laska, M. e Hudson, R. 1993. Discriminating parts
from the whole: Determinants of odor mixture percep-
tion in squirrel monkeys, Saimiri. Comp. Physiol. 173:
Laska, M., Hudson, R. e Alicke, T. 1996. A study of long-
term odor memory in squirrel monkeys (Saimiri sciureus).
J. Comp. Psychol. 110: 125-130.
Lorenzi, H. 1992, Arvores Brasileiras. Nova Odessa: Editora
Plantarum. 384pp.
Ribeiro, J. E. L. da S. 1999. Flora da Reserva Ducke: Guia
de Identificaado das plants Vasculares de Terra-firme na
Amazdnia Central. Manaus: INPA. 816pp.
Rimoli, J. 2001. Ecologia de macacos-pregos (Cebus
apella nigritus, Goldfuss, 1809) na Estagao Biol6gica
de Caratinga, MG: Implicagoes para a Conservagao de
Fragmentos da Mata Atlhntica. Tese de Doutoramento,
Centro de Ciencias Biol6gicas, Universidade Federal
do Para.
Robinson, J. G. e Janson, C. H. 1987. Capuchins, squirrel
monkeys and Atelines: socioecological convergence with
Old World Primates. Em: Primate Societies, Smuts, B.B.
et al. (eds.). Chicago: Chicago Society Press.
Sigg, H. e Stolba, A. 1981. Home Range and Daily March
in a Hamadryas Baboon Troop. Folia Primatol. 36(1-2):
Silva Jdnior, J. S. 2002. Taxonomy of capuchin monkeys,
Cebus Erxleben, 1777. Neotrop. Primates 2(1): 29.
Sussman, R. W. 2000. Primate ecology and social structure:
New World monkeys. Needham Heights, Pearson Custom
Publishing, 207pp.
Terborgh, J. 1983. Five new world primates: a study in
comparative ecology. Princeton University Press, New
Ueno, Y. (1994a). Olfactory discrimination of eight food
flavors in the capuchin monkey (Cebus apella): Com-
parison between fruity and fishy odors. Primates 35:
Ueno, Y. (1994b). Olfactory discrimination of urine odors
from five species by tufted capuchin (Cebus apella). Pri-
mates 35: 311-323.


Tiago Soares Bortolini
Jilio Cisar Bicca-Marques


Tool use, the use of a free object in the environment as a
functional extension of one's own body (Beck, 1980), has
been reported in invertebrates, fish, birds, and mammals,
including primates (Alcock, 1989). However, tool use is
not common or widespread in nonhuman primates. It has
been observed in a small number of species including chim-
panzees, bonobos, gorillas, orangutans, some macaques
and baboons, and capuchin monkeys (van Schaik et al.,
1999). Captive capuchin monkeys (Cebus spp.) were long
reported to use tools in a variety of contexts (Visalberghi,
1990; Urbani, 1999). Recently, however, a growing body
of evidence shows that semi-captive and free-ranging capu-
chins in several populations and species use tools, includ-
ing the use of stones as hammers and anvils to crack and
open nuts (Ottoni and Mannu, 2001; Fragaszy et al., 2004;
Moura and Lee, 2004; Waga et al., 2006). Tool-making is
a cognitively complex process that involves an intentional
modification of the tool for improving its efficiency (Beck,
1980). Reports of non-human primate tool-making have
been restricted to the great apes (Boesch and Boesch, 1990;
Fontaine et al., 1995; Tomasello and Call, 1997; Schick
et al., 1999; van Schaik et al., 2003) with the exception
of a few experimentally induced cases in captive capuchins
(Westergaard and Suomi, 1994, 1995; Westergaard et al.,
1995). Here we report a case of spontaneous tool-making
by a captive capuchin monkey.


A group of capuchin monkeys (Cebus sp.) composed of an
adult male, two adult females and three immature males
living in an enriched enclosure 7.0 m long x 8.7 m wide
x 2.9 m high at the Sapucaia do Sul Zoological Park, State
of Rio Grande do Sul, Brazil, was opportunistically (ad
libitum) observed and video-taped in January and Febru-
ary 2007. The enclosure included sand on the floor, trees,
stones, and perches for the monkeys. For enrichment pur-
poses, food was concealed inside ice cubes, PVC pipes, and


On 12 January 2007, an adult female (putative Cebus
nigritus) was observed banging a twig with a piece of stone
against a larger stone, licking/chewing and likely extracting
something from it with her mouth. She was then observed
probing an unseen structure (probably a hole in the enclo-
sure's drinking fountain) with the modified twig (Fig. 1).
This sequence of events occurred very rapidly. The latency

Neotropical Primates 14(2), August 2007

Figure 1. Adult female capuchin monkey (putative Cebus nigritus) using a hammer stone and an anvil stone to pound on a twig that was
then inserted into a crevice (photos from video frames).

.- V. Y 1 I
Figure 2. Adult female using a stone to crack open an ice cube
with food inside.

between the end of banging and the start of probing was
3 to 4 seconds, during which time the female moved from
the banging site to the probing site. After this observation,
the group was monitored for 15 days and no additional
cases of tool-making were observed. The capuchins, how-
ever, often used stones as hammers to crack nuts and other
foods, including ice cubes containing food (Fig. 2).


Although we do not know what happened immediately
prior to this behavioral sequence and could not see whether
the female acquired anything as a result of probing, the
speed at which this sequence of events occurred is highly
suggestive of a causal understanding during object ma-
nipulation and seems to qualify as a case of spontaneous
tool-making. Future research will focus on confirming the
capuchin monkeys' capability to make tools, an ability
that would suggest less cognitive difference than is pres-
ently thought to exist between capuchins and the great apes
(Visalberghi, 1990, 1997; Urbani and Garber, 2002). In
addition to suggesting that capuchin monkeys understand
cause and effect relationships during object manipulation,
these findings strengthen the argument that the mainte-
nance of captive animals in enriched environments is an

important strategy to allow the expression of the species'
fullest behavioral repertoire. A previous study of the same
group (200 hours during 2002-2003) in this enclosure,
but with minimum enrichment (paved ground, a single
swing and no feeding enrichment), failed to record any case
of tool use (D.B. Montano, personal observation). Envi-
ronmental enrichment serves an important function in im-
proving capuchin monkeys' welfare by reducing boredom
and eliciting tool use.


We thank the staff of the Ndcleo de Zoologia of the Sapu-
caia do Sul Zoological Park, especially Renato Petry Leal
and Marcelo Linck, for the permission to conduct this
study. We also thank Paul A. Garber and Bernardo Urbani
for reviewing an earlier version of this manuscript.

Tiago Soares Bortolini, Instituto de Biociencias, Uni-
versidade Federal do Rio Grande do Sul, Avenida Bento
Gongalves 9500, Campus do Vale, Predio 43323 Sala 115,
Porto Alegre, RS 91501-970, Brazil, e-mail: gmail.com>, and J6lio CUsar Bicca-Marques, Laborat6rio
de Primatologia, Faculdade de Biociencias, Pontificia Uni-
versidade Cat61lica do Rio Grande do Sul, Avenida Ipiranga
6681 Predio 12A, Porto Alegre, RS 90619-900, Brazil,
e-mail: .


Alcock, J. 1989. Animal Behavior. 4th ed. Sinauer,
Beck, B. B. 1980. Animal Tool Behavior. Garland, New
Boesch, C. and Boesch, H. 1990. Tool use and tool making
in wild chimpanzees. Folia Primatol. 54: 86-99.
Fontaine, B., Moisson, P. Y. and Wickings, E. J. 1995. Ob-
servations of spontaneous tool making and tool use in a
captive group of western lowland gorillas (Gorilla go: A//l
go: JlaI. Folia Primatol. 65: 219-223.
Fragaszy, D., Izar, P., Visalberghi, E., Ottoni, E. B. and
Oliveira, M. G. 2004. Wild capuchin monkeys (Cebus

Neotropical Primates 14(2), August 2007

libidinosus) use anvils and stone pounding tools. Am. J
Primatol. 64: 359-366.
Moura, A. C. A. and Lee, P. 2004. Capuchin stone tool use
in Caatinga dry forest. Science 306: 1909.
Ottoni, E. B. and Mannu, M. 2001. Semifree-ranging
tufted capuchins (Cebus apella) spontaneously use tools
to crack open nuts. Int. J. Primatol. 22: 347-358.
Schick, K. D., Toth, N., Garufi, G., Savage-Rumbaugh,
E. S., Rumbaugh, D. and Sevcik, R. 1999. Continuing
investigations into the stone tool-making and tool-using
capabilities of a bonobo (Pan paniscus). J Archaeol. Sci.
26: 821- 832.
Tomasello, M. and Call, J. 1997. Primate Cognition. Oxford
University Press, New York.
Urbani, B. 1999. Spontaneous use of tools by wedge-
capped capuchin monkeys (Cebus olivaceus). Folia Prima-
tol. 70: 172-174.
Urbani, B. and Garber, P. A. 2002. A stone in their hands...
Are monkeys tool users? Anthropologie 40: 183-191.
van Schaik, C. P., Deanes, R. 0. and Merrill, M. Y. 1999.
The conditions for tool use in primates: Implications
for the evolution of material culture. J Hum. Evol. 36:
van Schaik, C. P., Ancrenaz, M., Borgen, G., Galdikas, B.,
Knott, C. D., Singleton, I., Suzuki, A., Utami, S. S. and
Merrill, M. 2003. Orangutan cultures and the evolution
of material culture. Science 299: 102-105.
Visalberghi, E. 1990. Tool use in Cebus. Polia Primatol. 54:
Visalberghi, E. 1997. Success and understanding in cogni-
tive tasks: A comparison between Cebus apella and Pan
troglodytes. Int. J Primatol. 18: 811-830.
Waga, I. C., Dacier, A. K., Pinha, P. S. and Tavares, M. C.
H. 2006. Spontaneous tool use by wild capuchin mon-
keys (Cebus libidinosus) in the Cerrado. Folia Primatol.
77: 337-344.
Westergaard, G. C. and Suomi, S. J. 1994. The use and
modification of bone tools by capuchin monkeys. Curr.
Anthropol. 35: 75-77.
Westergaard, G. C. and Suomi, S. J. 1995. The manufac-
ture and use of bamboo tools by monkeys: Possible im-
plications for the development of material culture among
east Asian hominids. J Archaeol. Sci. 22: 677- 681.
Westergaard, G. C., Greene, J. A., Babitz, M. A. and
Suomi, S. J. 1995. Pestle use and modification by tufted
capuchins (Cebus apella). Int. J Primatol. 16: 643-651.


Luisa Xavier Lokschin
Rodrigo Cambard Printes
Juliane Nunes Hallal Cabral
Gerson Buss


Urban growth affects ecosystems in several ways, leaving
them more vulnerable (Alberti and Marzluff, 2004). In
Porto Alegre, the combined effects of human presence in-
cluding deforestation, hunting and other indirect effects are
reducing howler's area distribution with consequences still
unknown (Lokschin et al., 2005). Human density within
a primates' geographical area should be considered by the
World Conservation Union (IUCN) in the evaluation of
species status (Harcourt and Parks, 2003). The southern
brown howler monkey (Alouatta guariba clamitans, Ca-
brera 1940) is considered an endangered species in Rio
Grande do Sul (Marques, 2003); in Brazil and globally it is
considered near threatened (Machado et al., 2005; Rylands
et al., 2006).

There are many species of Neotropical primates living
close to urban areas, including: Alouatta clamitans (Buss,
1996), Alouatta caraya (Codenotti et al., 2002), Callicebus
nigrifrons (Oliveira et al., 2003), Saguinus leucopus (Poveda
and Sinchez-Palomino, 2004) and Saguinus bicolor (Vas-
concelos et al., 2005). Problems and threats linked to ur-
banization, such as danger from vehicles when crossing
roads, predation by dogs and electric hazard, are already
documented for A. clamitans (Printes, 1999; Alonso et al.,
2005), C. jacchus (Menezes, 2005) and S. bicolor (Vascon-
celos et al., 2005). Ecosystems close to urban areas are im-
portant for wildlife (Dickman, 1987) and measures must
be taken to guarantee their existence. Howler monkeys (A.
g. clamitans) utilize areas of forests close to urban develop-
ments and are suffering from contact with several electric
hazards. Here we describe a way to mitigate the occurrence
of such accidents around Porto Alegre.


Porto Alegre is the capital city of the state of Rio Grande
do Sul, Brazil (Fig. 1), with a population of 1.4 million
(IBGE, 2006). Approximately 10% of the municipal area
is natural semi-deciduous seasonal forest, influenced by At-
lantic rainforest (Brack et al., 1998; Velez et al., 1998). The
southernmost area of the municipality (Fig. 2) is a rural
landscape containing a number of small villages. The most
important natural areas are also in this zone, which is also
the most important area for howler monkeys (Alouatta
guariba clamitans) (Romanowski et al., 1998; Lokschin
et al., 2005). Lami Biological Reserve, the only biological

Neotropical Primates 14(2), August 2007

reserve of the city, is located in this region (3014'13,2"S,
51005'43,4"W). In its buffer zone there are some forest
fragments and a district named Lami, which is an urban
area holding around 2,700 people (Porto Alegre, 2006),
with many native trees still remaining around the houses.
Most of the power lines in Brazil are aerial. Since 1999, the
'Programa Macacos Urbanos'(Urban Monkeys Program) has
been documenting howler monkey mutilations and deaths
caused by electric hazards in Lami (Printes, 1999). We con-
ducted a survey to identify the critical areas where electric
cables, responsible for the majority of accidents, should be
insulated. These areas were identified based on previously
documented howler distribution (Romanowski, 1998), the
presence of vegetation, known accidents and reports from
local people. Areas where electric cables pass through tree
branches (especially those of Ficus organensis), and ripar-
ian forests were prioritized for cable insulation. Areas where
howlers died or were mutilated were plotted on a Global
Positioning System (GPS) and photographed. String bridg-
es were also installed on three critical points, based on as-
sessment of risk of animals being electrocuted (Figure 2, a,
b and c).

Results and Discussion

From 2000 to 2006 eight howlers were electrocuted and
three of them died as a result of injuries sustained. This
number is lower than that found by Printes (1999) who re-
corded three cases in one year in the same locality. But our
results may have underestimated the total number of deaths
and mutilations caused by electric discharge since not all
accidents are reported or recorded. When animals attempt

to use cables as a bridge or as a base (Fig. 3), they support
two of their limbs on two different cables causing a short
circuit. Electric current, passing through the animals' body,
may cause burns, hemorrhage and cardiac arrest, which can
eventually kill them. Low tension cables (127-380 v) are
the main cause of accidents, due to their frequent use by
howlers as bridges between forest fragments. Howlers are
not the only victims of such accidents in the region. Birds
with large wing spans are also susceptible; these include
the southern screamer (Chauna torquata Oken, 1816) and
striped owl (Rhinoptynx clamator Vieillot, 1807).

The first step to reduce electric hazards was the installation
of a bridge in 1999. It consisted of two parallel vertical
strings. This model of bridge had no success and was re-
placed by another one, similar to a 'ship ladder' but in a
horizontal orientation (Figures 4 and 5). The 'steps of the
ladder' are used by howlers in a horizontal plane (the same
way ladders are used by humans vertically). We reported
accidents to the relevant authorities every time we became
aware of their occurrence and requested the insulation of
the cables, at least in critical areas. As nothing was done by
2003 a legal strategy was adopted; in March 2003, a legal
case was presented based on National Legislation for Envi-
ronmental Crimes (no 9605/98), citing known cases. This
resulted in Civil Inquiry no 21/03 and the legal authorities
judged that the Electric Energy State Company (CEEE)
should insulate cables in critical areas. The insulation work
started in 2004 costing approximately US$ 30,000. The
'Programa Macacos Urbanos' was ordered by the court to
designate priority areas, identifying those in which cables
presented a higher risk of accidents. We decided that close

Figure 1. Porto Alegre, RS, Brazil. (300S, 51'W)

Neotropical Primates 14(2), August 2007

collaboration with local communities was vital for the suc-
cessful implementation of any conservation activities. The
first step taken to engage with local communities was to
use about 10 local residents to monitor the occurrence of
howler electrocutions pre-installation and use of bridges
post-installation. Thus, the presence of houses and people's
availability to collaborate was a decisive point in choosing
areas for new bridges. Two other bridges were installed in
critical areas in 2003 and 2004, with local people report-
ing occasional use by howlers. In March 5, 2006, we saw a
group of howlers crossing one of these bridges (Fig. 2c and
Fig. 5). There seems to be a seasonal pattern to the use of
bridges by howlers, probably linked to the availability of
food resources.

We suggest that bridges should be considered a comple-
mentary activity to the insulation of electric cables. In ad-

Figure 2. Southernmost area of Porto Alegre, and the neighbor-
ing city of Viamao, Itapua district, a, b and c are critical points
where bridges for howler monkeys were installed in Lami, Porto
Alegre, RS, Brazil.

edition to insulation, the three low tension cables must be
braided forming only one cable, thereafter reducing the
possibility of animal use. Since the first bridge was installed
and critical areas were isolated accidents have become rare.
One accident was recorded in 2005 in an already insulated
area, which had its terminal poles exposed. Since then, the
CEEE arranged to insulate all terminals. In 2006, another
howler was hurt on high tension cables, in an area already
requested to be isolated. The CEEE does not have a way to
isolate this type of cables, so pruning was requested. These
two last cases suggest that prioritized areas are actually
being used by howlers and that they are exposed to danger.
If cables are not insulated, they offer potential risks to the
animals. Nowadays, to reduce accidents, CEEE has taken
the responsibility for keeping cables insulated and trees cut.
This legal decision in favor of wild animals was the first one
in the country and might set a precedent in Environmental

Figure 4. String bridge, 'ship ladder' model, before being installed
in Lami, Porto Alegre. (Photo by L. X. Lokschin).

Figure 3. Hov leI mi nkc i J. ;,,'m:.:;.:',: .:../: .,:,,..), using cables
as a bridge, at Lami, Porto Alegre. (Photo by L. X. Lokschin).

Figure 5. String bridge where howlers were seen crossing between
forest fragments at Lami, Porto Alegre. (Photo by G. Buss).

Neotropical Primates 14(2), August 2007

Justice. The 'Programa Macacos Urbanos'will keep supervis-
ing selected areas around Porto Alegre, monitoring risks to
howler monkeys from power lines and installing bridges in
critical points. In 2006, a howler died from electrocution
crossing low tension cables at Itapua Village (3017'00"S,
5101'19"W), in the neighboring municipality of Viamao,
20 km from Lami (Fig. 2). This resulted in a preliminary
study of other cases at Itapua Village that revealed at least
five other deaths caused by contact with electric cables. The
first one was in 1995. At present, we are mapping critical
points where cables must be insulated at Itapua.

Conservation Consequences

In 2002, a Municipal Law (No 9.971) was created regu-
lating the use of 'ecological' power lines in Porto Alegre.
These lines can be either underground or aerial, with an
insulating or semi-insulating cover, and built in a com-
pact form. With appropriate monitoring and commu-
nity participation the enforcement of this law (Decree no
14.196/03) should guarantee a lower impact of cables on
wild howler populations. However, the application of this
law is not retroactive; areas where cables were installed
before 2003 are still dangerous and should be monitored.
Other municipalities should create laws concerning power
lines and their impact on flora and fauna; we suggest that
researchers should collaborate in this long process. Brazil-
ian researchers should meet in a forum to discuss primates
in urban areas. The aim of this forum should be getting to
know common problems and standardizing proposals and
actions to be taken.


We would like to thank the cooperation of the "Ndcleo
Amigos da Terra Brasil" (Friends of Earth), managers of
the Lami Biological Reserve, Eng. Nelson (CEEE), Andrd
Chein Alonso, Jos6 Samir Borges, (Fundagno Zoobotnnica
do Rio Grande de Sul), Gleide Marsicano, Comando de
Policiamento Ambiental, Dr. Helena Romanowski, Dr.
Liane Printes, Dr. Fernanda Michalski and MSc. Darren
Norris for suggestions on the manuscript. Special thanks to
Enio Mancuso (in memorial).

Luisa Xavier Lokschin, Rodrigo Cambari Printes, Juliane
Nunes Hallal Cabral and Gerson Buss, Programa Maca-
cos Urbanos, Departamento de Zoologia Instituto de
Biociencias Universidade Federal do Rio Grande do Sul
(UFRGS), Av. Bento Gongalves, n 09500 prddio 43435,
sala 218, CEP 90501-970, Porto Alegre, RS, Brasil, e-mail:
, ,
, .


Alberti, M. & Marzluff, J. M. 2004. Ecological resilience in
urban ecosystems: Linking urban patterns to human and
ecological functions. Urban Ecosystems 7: 241-264.

Alonso, A. C.; Cabral, J. N. H.; Lokschin, L. X.; Marsicano,
G.; Roll, A.; Buss, G.; Printes, R. C.; Santos, M. E;
Jardim, M. M. A. & Romanowski, H. P. 2005. Program
Macacos Urbanos: Manejo de bugio-ruivo (Alouatta
guariba clamitans Cabrera 1940) em Porto Alegre. Livro
de resumes do XI Congresso Brasileiro de Primatologia: 67.
Brack, P.; Rodrigues, R. S.; Sobral, M.; Leite, S. L. de C.
1998. Arvores e arbustos na vegetagao natural de Porto
Alegre, Rio Grande do Sul, Brasil. Iheringia Ser. Bot.
51(2): 139-166.
Buss, G. 1996. Urban Monkeys Alouatta fusca in the
municipality of Porto Alegre. Neotrop. Primates 4(2):
Codenotti, T. L.; Silva, V. M.; Albuquerque, V. J.; Camargo,
E. V.; Silveira, R. M. M. 2002. Distribuigao e situaiao
atual de conservagao de Alouatta caraya (Humboldt,
1912) no Rio Grande do Sul, Brasil. Neotrop. Primates
10 (3): 132-140.
Dickman, C. R. 1987. Habitat fragmentation and
vertebrate species richness in an urban environment. J.
AppI. Ecol. 24: 337-351.
Harcourt, A. H. & Parks, S. A. 2003. Threatened primates
experience high human densities: adding an index of
threat to the IUCN Red List criteria. Biol. Conserv. 109:
IBGE, 2006. Institute Brasileiro de geografia e estatistica.
Website: Accessed on March
30, 2006.
Lokschin, L. X.; Paim, E P.; Cabral, J. N. H.; Rossato, R.
S.; Setubal, R. B.; Alonso, A. C.; Silva, F. E.; Buss, G. &
Romanowski, H. P. 2005. Program Macacos Urbanos:
Ocorrenciae distribuigao do bugio-ruivo (Alouattaguariba
clamitans Cabrera, 1940) em Porto Alegre, RS, Etapa 2 -
resultados preliminares. Livro de resumes do XI Congresso
Brasileiro de Primatologia: 120. SBPr- PUC RS.
Machado, A. B. M.; Martins, C. S. & Drummond, G. M.
2005 Lista da fauna brasileira ameagada de extinygo.
Fundagao Biodiversitas. Belo Horizonte, MG, Brasil,
Marques, A. A. 2003. Primatas. In. Livro da
Fauna Ameacada de Extincdo no Estado do Rio Grande
do Sul. Fontana, C; Bencke, G. & Reis, R. E. (eds),
pp. 499-506. Edipucrs, Porto Alegre, RS, Brasil.
Menezes, M. 0. T. 2005. Influencia da urbanizagao sobre
uma populagao de C .-' jacchus (Linnaeus, 1758) de
vida livre no Campus do PICI (UFC), Fortaleza, Ceari.
Livro de resumes doXI Congresso Brasileiro dePrimatologia:
129. SBPr-PUCRS.
Oliveira, R. C. R., Coelho, A. S. & Melo, F R. 2003.
Estimativa de densidade e tamanho populacional de
saui (Callicebus nigrifrons) em um fragmento de mata
em regeneracao, Vigosa, Minas Gerais, Brasil. Neotrop.
Primates 11(2): 91-94.
Porto Alegre, 2006. Censo populacional de Porto Alegre.
Website: default.php?reg=46&p_secao=43> Accessed on March
28, 2006.

Neotropical Primates 14(2), August 2007

Poveda, K. & Sinchez-Palomino, P. 2004. Habitat use
by the White-footed tamarin, Saguinus leucopus: a
comparison between a forest-dwelling group and an
urban group in Mariquita, Colombia. Neotrop. Primates
12(1): 6-9.
Printes, R. C. 1999. The Lami Biological Reserve, Rio
Grande do Sul, Brazil and the danger of power lines
to howlers in urban reserves. Neotrop. Primates 7(4):
Romanowski, H. P.; Dornelles, S. da S.; Buss, G.; Brutto,
L. E G.; Jardim, M. M. de A; Printes, R. C.; Fialho, M.
de S. 1998. Bugio-ruivo: o ronco ameagado. In: Atlas
Ambiental de Porto Alegre, Menegat, R. (ed), pp. 62- 63.
UFRGS / PMPA / INPE, Porto Alegre, RS, Brasil.
Rylands, A.B., Bampi, M.I., Chiarello, A.G., da Fonseca,
G.A.B., Mendes, S.L. & Marcelino, M. 2003. Alouatta
guariba. In: IUCN 2006. 2006 IUCN Red List of
Threatened Species. Website .
Accessed May 5, 2006.
Vasconcelos, C. M.; Subird, R. J. & Kluczkovski Jr.,
A. 2005. Projeto piloto de reintegracao de grupos de
sauim-de-coleira, Saguinus bicolor, ao habitat natural
em Manaus, Amazonas, Brasil. Livro de resumes do XI
Congress Brasileiro de Primatologia: 176. SBPr-PUC
Velez, E. M.; Meira, J. R. & Oliveira, P. L. 1998. Avaliadao
dos morros com base no uso do solo. In: Atlas Ambiental
de Porto Alegre, Menegat, R. (ed), pp. 81 Editora da
Universidade. UFRGS/PMPA/INPE, Porto Alegre, RS,


Sidn S. Waters and Oscar Ulloa


Black howler monkeys (Alouattapigra) and Yucatan spider
monkeys (Ateles ,.. yucatanensis) are the only non-
human primate species found in Belize. Black howler
monkeys occupy the most restricted range of any other
species in the genus Alouatta (Wolfheim, 1983), and
are listed as Endangered in the IUCN Red List (IUCN,
2006). The Yucatan spider monkey is listed as Vulner-
able in the IUCN Red List (IUCN, 2006). Both species
are threatened with ongoing habitat loss and degradation
(IUCN, 2006). Howler monkeys are the focus of the
Community Baboon Sanctuary Conservation Program in
an area along the Belize River (Horwich, 1990) and have
been reintroduced in the Cockscomb Wildlife Reserve
(Horwich et al., 1993). Data have been collected on black
howler monkey group size and ranging behavior in two
areas of Belize (Horwich et al., 2001; Pavelka, 2003), but
no countrywide survey of the species has taken place since
the 1980s (Horwich and Johnson, 1986). Spider monkeys
are vulnerable to habitat fragmentation because they occur

in low numbers, have low fecundity rates, and rely on ripe
fruit, a patchily distributed food resource (Meffe and Car-
roll, 1994). Basic information on this species in Belize is
lacking. A known area of spider monkey distribution in
the Chiquibul protected area (comprised of the Chiquibul
Forest Reserve and the Chiquibul National Park) is heav-
ily frequented by illegal collectors of xate palm leaves
(Chamaedorea sp.). This activity has been prevalent since
1998 (Anon, 2005) and up to 1,000 illegal xate collectors
have been reported to camp and hunt in the area while
harvesting the leaf (Friends for Conservation and Devel-
opment, 2005). This must give cause for concern for the
species even in a protected area.


We undertook a countrywide assessment of human/
wildlife conflict among subsistence farmers in Belize from
March to May 2006 (the results of which will be pub-
lished elsewhere). As part of the survey we asked questions
about the presence of howler monkeys, spider monkeys
and Baird's tapir (Tapirus bairdii). The latter two spe-
cies were chosen because they are amongst the first spe-
cies to disappear from over-exploited forests (Bodmer et
al., 1997) and are, as such, more vulnerable to popula-
tion fragmentation and eventual extinction. Black howler
monkeys were included in the list because they often in-
habit similar riparian habitat to that of the Baird's tapir.
Using a structured questionnaire, we surveyed all districts
of Belize for evidence of crop raiding by wild animals. The
villages surveyed were all outside protected areas and were
selected because they were predominantly dependent on
subsistence agriculture. Villages and communities whose
livelihoods depended on employment in intensive agricul-
ture such as cattle ranching, and sugar cane, citrus and
banana plantations were excluded. The questionnaire was
administered at every sixth house in a village to a person
who worked on his/her farm. The respondents' farms or
gardens were typically situated outside the villages sur-
veyed and were visited when possible. When the question-
naire was completed, and if the respondent had not already
mentioned the species as a crop raider, the respondent was
asked about the presence of primates and tapirs in the area.
GPS locations for each species were recorded if presence
was reported by at least two respondents independently of
one another in the village surveyed or, if the animals were
directly observed, or if howler monkey vocalizations were
heard. Howler vocalizations could occur at any time of the
day or night. Early morning walks were taken in all forests
where the focal species were reported as occurring. A GIS
map was generated from these data.


A total of 168 people were interviewed during the survey
and 14.9% reported that howler monkeys were present near
their farms or gardens. Reports of the presence of black
howler monkeys came from all districts except Corozal,

Neotropical Primates 14(2), August 2007

and these were verified in 43% of cases either by sighting
or hearing the monkeys. The GIS map of these locations
can be seen in Figure 1. Black howler monkeys were not
reported as a crop raider by any of the survey respondents.
This species was heard frequently in the watershed of the
Temash River in southwestern Toledo and was also heard in
undisturbed coastal forest in the southeast of Belize. Groups
were also commonly reported and heard in the northern
part of Cayo District where riparian forest is still common
(Fig. 1). Spider monkeys were reported by respondents on
seven occasions from three districts, including reliable re-
ports from Orange Walk (Fig. 1) in the northwest of the
country, where a group of about 10 animals was reported
as living in a forest on the edge of a working quarry on the
western border with Guatemala. A respondent in Toledo
stated that a group of about eight spider monkeys raided
his pineapple crop when it was ripe. All other reports of the
species were recorded in Cayo District (Fig. 1).


Neither primate species were reported as a serious crop
raider in Belize. Unsurprisingly, black howler monkeys
are widespread in the villages participating in the Com-
munity Baboon Sanctuary Conservation Program (Hor-
wich, 1990). They also are present in the upper watershed
of the Temash River in the southwest of Toledo District,
where much of the forest was damaged by Hurricane Iris
in 2001. Additionally, we believe our auditory detection of

10 0 10 20 30 40 60 60 Hjbmtn*


* DW fo"

*. SpdUtrdonteI
SSeize OaMt BWuiI

Figure 1. GIS map showing the distribution of black howler mon-
keys and spider monkeys in Belize.

a group in undisturbed coastal forest in the southeast of
Belize is a new report for this species. This area of forest
is presently undisturbed as the inhabitants of the nearby
village rely on fishing for their livelihood. The groups re-
ported from the upper Temash River and in Cayo Dis-
trict all occur in unprotected areas of forest increasingly
utilized for logging and agricultural purposes. Reports of
spider monkeys were less frequent and came from only
three districts, Orange Walk, Cayo and Toledo, but the
survey team was unable to verify their presence. This
may be because there are fewer spider monkeys, and/or
because spider monkeys are more difficult to locate be-
cause they are more quiet and wary. The spider monkey
population in Belize needs further surveys on its areas of
distribution, along with research on abundance, density,
connectivity and demographics in order to determine ap-
propriate conservation actions. This report on the distri-
bution of both species of non-human primates in Belize
is not meant to be definitive but is a useful first step in
identifying populations outside protected areas that need
active conservation management. These areas include the
Cayo District for both species and the upper watershed of
the Temash River in the southwest of Toledo District for
black howler monkeys. These populations would benefit
from closer scrutiny to ascertain their long-term sustain-
ability and their suitability for a potential conservation
effort involving local stakeholders. Although there are
far fewer reports of the presence of spider monkeys, they
may be more common than this survey demonstrates,
particularly in Belize's extensive protected areas' network,
but determining this will require further investigation.


We are very grateful to the Royal Zoological Society
of Scotland (Edinburgh Zoo) for fully funding this
survey and to the Government of Belize, Department
of Natural Resources for granting permission to under-
take the work. We thank the farmers of Belize for their
patience and time answering our questionnaire. We
thank our field assistant, David Barclay for his help and
good humor, and Dr. John Morris of the Government
of Belize Institute of Arc' i,..1..;,- for help while in the
field. Isabel Gonzalez provided assistance with the map
and, along with Arnaud Desbiez and Maria Elisa Hob-
belink, provided helpful comments on an earlier version
of this manuscript.

Sign S. Waters, 14 Lindsay Gardens, Tredegar, Gwent
NP22 4RP UK, e-mail: ,
and Oscar Ulloa, Department of Forestry, 24/25 Unity
Blvd., Belmopan City, Belize.


Anon. 2005. Xatd in Belize, a Growers Guide. Belize Bo-
tanic Gardens, Ya'axche' Conservation Trust, Belize, and
the Natural History Museum, London, UK. Website:

Neotropical Primates 14(2), August 2007

http://www.belizebotanic.org/xate_manual.pdf. Accessed
25 October 2006.
Bodmer, R. E., Eisenberg, J. and Redford, K. 1997. Hunt-
ing and the likelihood of extinction of Amazonian mam-
mals. Cons. Biol. 11: 460-466.
Friends for Conservation and Development. 2005. Pro-
ceedings of the Chiquibul Stakeholders' Planning Work-
shop. 33pp. Website: http://www.eco-index.org/search/
pdfs/970report_l.pdf. Accessed 25 October 2006.
Horwich, R. H. 1990. How to develop a community sanc-
tuary: An experimental approach to the conservation of
private lands. Oryx 24: 95-102.
Horwich, R. H. and Johnson, E. D. 1986. Geographic dis-
tributions of the black howler (Alouattapigra) in Central
America. Primates 27: 53-62.
Horwich, R. H., Koontz, R., Saqui, E., Saqui, H., and
Glander, K. E. 1993. A reintroduction program for the
conservation of the black howler monkey (Alouattapigra)
in Belize. Endangered Species UPDATE 10: 1-6.
Horwich, R. H., Brockett, R. C. James, R. A. and Jones,
C. B. 2001. Population growth in the Belizean black
howling monkey (Alouatta pigra). Neotrop. Primates 9:
IUCN. 2006. IUCN Red List of Threatened Species. IUCN,
Gland, Switzerland. Website: http://www.iucnredlist.org.
Accessed 28 July 2006.
Meffe, G. K. and Carroll, C. R. 1994. Principles of Conser-
vation Biology. Sinauer Associates, Sunderland, MA.
Pavelka, M. S. M. 2003. Group, range and population size
in Alouatta pigra at Monkey River, Belize. Neotrop. Pri-
mates 11: 187-191.
Wolfheim, J. H. 1983. Primates of the World: Distribution,
Abundance and Conservation. University of Washington
Press, Seattle.


Anna B. S. Biintge
Lennart W Pyritz


Sympatry of two species belonging to the same genus
occurs rarely in Neotropical primates. It has been observed
regularly for Cebus apella and Cebus ai/b> ois (Terborgh,
1983), and for Saguinusfuscicollis and Saguinus sp. (Hey-
mann and Buchanan-Smith, 2000). In both cases, the
species occupy different ecological niches, through the
use of different forest strata or individual foraging strat-
egies. For species of the genus Alouatta there are several
limited cases of sympatry, for example Alouatta palliata
and Alouatta pigra in Tabasco, Mexico and at the south-
ern Belize-Guatemala border (Horwich and Johnson,
1986), Alouatta caraya and Alouatta guariba in southern
Brazil (Jdlio Cesar Bicca-Marques, pers. comm.) and the El

Pifialito Provincial Park in Misiones, Argentina (Di Bitetti,
2004), and A. palliata and Alouatta seniculus in northwest-
ern Colombia (Hernandez-Camacho and Cooper, 1976).
Here we report for the first time a clearly sympatric occur-
rence of the two howler monkey species Alouatta sara and
A. caraya.

Study area and Methods

The Rio Yacuma is a small tributary of the Rio Mamore,
approximately 10 km upstream of the village of Santa
Rosa (1410'S, 6652'W, Fig. 1). The Rio Yacuma flows
through the alluvial plain of the Rio Mamord at an eleva-
tion of approximately 150 m a.s.l. Heavy floods during the
rainy season transform the region into a vast swamp. Mean
annual temperature is 26C, and mean annual precipita-
tion is approximately 1,800 mm (Montes de Oca, 1997;
Navarro and Maldonado, 2002). The region is part of the
biogeographic sector of the Moxos lowlands, characterized
by tree savannahs and Varzea forests along the watercourses
(Navarro and Maldonado 2002). Observations were made
during a boat trip between April 10 and 12.

Results and Discussion

In April 2006, while travelling northeast of the city of
Rurrenabaque in the Beni Department in northern Bolivia
(Fig. 1), we observed two distinct howler species, A. caraya
and Alouatta sara (taxonomy following Groves, 2001), for-
aging and resting in close proximity (at a minimum distance
of approx. 100 m and in the range of vision of each other
on the bank of a river). The two species were clearly distin-
guished by the different coloration of the fur (black in male
and yellowish in female A. caraya, red in A. sara). Presum-
ably, the Rio Yacuma is not a natural barrier to the disper-
sal of the Alouatta species as both species were seen in the
gallery forest on the left and right bank of the river. Water
levels decline considerably during the dry season, probably
enabling the howler monkeys to cross the river. Groups of
A. caraya were observed three times. The observations in-
cluded: an adult male and an adult female; an adult male
and two adult females, one carrying a baby; and three adult
females with a male. Alouatta sara was seen two times. Once
we observed a group of three individuals-two adult males
and an adult female-foraging in a Cecropia concolor tree.
On another occasion, we saw a single adult male of A. sara
resting in the tree canopy. Several minutes of howling of
Alouatta groups were heard repeatedly in the morning and
in the late afternoon on both sides of the river.

In Bolivia, A. caraya has been observed at two localities in
the Santa Cruz Department and at various localities in the
Beni Department including the mouth of the Rio Yacuma.
Distributional notes on A. sara in Bolivia include localities
in the Beni, Cochabamba, La Paz, Pando, and Santa Cruz
departments. There have been no reports for the central
Beni or Rio Yacuma region, though (Anderson, 1997). Both
Alouatta species observed are assessed as Least Concern on

Neotropical Primates 14(2), August 2007

Figure 1. Location of the study area (black box) at the Rio Yacuma in the Beni Department, Northern Bolivia.

a global level by IUCN-The World Conservation Union
(2004) although declines have occurred in many parts due
to hunting pressure and habitat loss. The sympatric occur-
rence of the two Alouatta species at the Rio Yacuma raises
the following questions: 1) Is the sympatry of A. caraya
and A. sara a result of ecological differences (i.e., the use of
different forest strata, different foraging strategies or activ-
ity patterns)?, 2) Is interspecific territoriality the same as
within species territoriality?, and 3) Is the observed sym-
patry a recent phenomenon (i.e., caused by habitat loss
and hunting pressure in the surrounding area) or has it
existed for a longer time?


We thank our Bolivian guide Bismarck Vaca from Rur-
renabaque for his support and patience in the field. Dr.
Eckhard W. Heymann from the German Primate Centre
(DPZ) in Gbttingen, Dr. Michael Kessler, and Moritz
Rahlfs gave important comments on the manuscript.

Anna B. S. Biintge, Primate Genetics Working Group,
German Primate Centre (DPZ), Kellnerweg 4, 37077
Gbttingen, Germany, e-mail:
and Lennart W. Pyritz, Ecological Department of the
Johann-Friedrich-Blumenbach-Institute for Zoology and
An rl-...p..1.. ;., University of Gbttingen, Berliner Str. 28,
37073 Gbttingen, Germany, e-mail: gmx.net>.


Anderson, S. 1997. Mammals of Bolivia: Taxonomy and
Distribution. Bulletin of the American Museum of Nat-
ural History 231.
Di Bitetti, M. 2004. A comparative study of two howler
monkey species living in sympatry (Alouattaguariba and

Alouatta caraya) in El Pifalito Provincial Park, Argentina.
Cleveland Metroparks Zoo/Cleveland Zoological Society,
Scott Neotropical Fund: conservation/grants/scott/2004/diBitetti.asp>.
Groves, C. P. 2001. Primate Taxonomy. Smithsonian Insti-
tution Press, Washington, DC.
Hernindez-Camacho, J. and Cooper, R. W 1976. The
nonhuman primates of Colombia. In: Neotropical Pri-
mates. Field Studies and Conservation, R.W. Thorington
Jr. and P. G. Heltne (eds.), pp.32-69. National Academy
of Sciences, Washington, DC.
Heymann, E. W., Buchanan-Smith, H. M. 2000. The be-
havioural ecology of mixed-species troops of callitrichine
primates. Biol. Rev. 75: 169-190.
Horwich, R.H., Johnson, E.D. 1986. Geographical dis-
tribution of the black howler (Alouatta pigra) in Central
America. Primates 27: 53-62.
Montes de Oca, I. 1997. Climatologia. In: C-... i Recur-
sos Naturales de Bolivia. Montes de Oca, I., pp.125-142.
Universidad Nuestra Sefora de La Paz, La Paz.
Navarro, G. 2002. Provincias biogeograficas del Beni y del
Pantanal. In: C.. F' Ecoldgica de Bolivia. Vegetacidn
y Ambientes Acudticos, Navarro, G., Maldonado, M.,
pp.157-193. Fundaci6n Simon I. Patifo, Departamento
de Difusi6n, Cochabamba.
Terborgh, J. 1983. Five New World Primates. Princeton
University Press, Princeton.
World Conservation Union 2004. IUCNRedList of Threat-
ened Species. A Global Species Assessement. World Conser-
vation Union, Gland.

Neotropical Primates 14(2), August 2007


Fernanda Almeida Cunha, Maria Aparecida Lopes,
Sidnei de Melo Dantas, Nivia Aparecida Silva do Carmo,
Suleima do Socorro Bastos da Silva

O caiarara ka'apor (Cebus kaapori Queiroz, 1992) tem uma
Area de distribuigao geografica relativamente pequena, se
comparada is de outros primatas amaz6nicos. Esta esp6cie
tem sido encontrada sempre em baixissimas densidades em
seu habitat natural. Os fatores determinantes de sua distri-
buigao e as causes de sua raridade local ainda permanecem
desconhecidos. A area de ocorrencia original do caiarara
ka'apor coincide corn a regiao de ocupagao humana mais
antiga da Amaz6nia, marcada por grandes desmatamentos e
intense fragmentagao de habitat. A soma destes fatores faz
com que C. kaapori seja listada como uma das poucas esp6-
cies de mamiferos amaz6nicos criticamente em perigo de ex-
tingao (Ferrari e Queiroz, 1994; Lopes e Ferrari, 1996; Car-
valho Jr., 2003; Silva Jr. e Queiroz, no prelo; SECTAM, em
preparacao). A area de ocorrencia originalmente identificada
deste primata 6 a Amaz6nia Oriental, a oeste do Estado do
Maranhao entiree os rios Gurupi e Pindare), com limits entire
a Floresta Amaz6nica e a Zona dos Cocais (Queiroz, 1992).
Posteriormente, esta area foi ampliada a noroeste do Mara-
nhao e leste do Pari, entire os rios Tocantins e Grajad, atra-
v6s de observagces diretas e informagoes de habitantes locais
durante inventirios faunisticos realizados na regiao (Lopes,
1993; Ferrari e Lopes, 1996; Lopes e Ferrari, 1996; Silva Jr. e
Cerqueira, 1998; Carvalho et al., 1999, Silva Jr., 2001; Silva
Jr. e Queiroz, no prelo). Por6m, os limits da distribuigao
geogrifica de C kaapori ainda nao estao bem definidos. 0
limited oeste da area de distribuigao da espcie parece coincidir
corn a margem direita do rio Tocantins, mas os limits norte,
leste e sul ainda estao indeterminados. Os limits orientais e
meridionais da distribuigao original podem ter sido retrai-
dos devido a grande perda de habitat natural conseqiiente da
ocupagao humana (Silva Jr. e Queiroz, no prelo).

A Usina Hidrel6trica de Tucurui (UHE-Tucurui), construi-
da entire 1984 e 1985, situa-se a cerca de 300 km ao sul de
Belkm, Pard (3043'-5015'S, 49012'-5000'0) (Eletronorte,
1985). Em 2002, foram criadas a Area de Protegao Ambien-
tal, a APA Lago de Tucurui, que abrange toda a Area do en-
torno do reservat6rio e duas Reservas de Desenvolvimento
Sustentivel (Pucurui Ararao e Alcobaga). Com o estabeleci-
mento de duas Zonas de Protegao da Vida Silvestre (antigas
Areas de soltura chamadas Base 3 e Base 4) dentro da APA
em 2004, completou-se o mosaico de unidades de conserva-
gao do Lago de Tucurui (Eletronorte, 2006). A fauna da Area
foi inventariada antes da realizagao do empreendimento e os
animals capturados durante o enchimento do reservat6rio
foram translocados para Areas de soltura ao redor do lago
(Eletronorte, 1985). Desde entao, estudos sobre a biota local
foram realizados pela ELN em parceria com instituig6es

como a Universidade Federal do Pard (UFPA) e o Museu
Paraense Emilio Goeldi (MPEG) (por exemplo, Ferrari et
al., 2002; Santos, 2002; Ferrari, 2003; Silva, 2003; Sam-
paio, 2004; Camargo, 2005; Vieira, 2005; Veiga, 2006).

Desde 2005, o MPEG e instituigoes colaboradoras UFPA
e Institute Nacional de Pesquisas da Amaz6nia (INPA) -
avaliam a situacao das comunidades de anfibios, rnpteis,
aves e mamiferos na APA Lago de Tucurui em convenio
corn a ELN atraves do projeto "Avaliacao e monitoramen-
to das comunidades de vertebrados na Area de influencia
do reservat6rio da UHE Tucurui". A fauna de mamiferos
esti sendo inventariada pelo m6todo de transeccao linear
em nove pontos de amostragem (quatro na margem direita
e cinco na margem esquerda). No period de janeiro de
2005 a agosto de 2006, mais de 900 km foram percorri-
dos (473,5 km na margem direita e 457,6 km na margem
esquerda) e 39 espcies de mamiferos foram registradas
(28 na margem direita e 35 na esquerda). Ate recentemen-
te, C. kaapori nao havia sido registrada na regiao de influ-
encia da UHE-Tucurui, apesar de esta estar dentro da Area
de distribuigao geogrifica proposta para a espcie (Masca-
renhas e Puorto, 1988, Lopes e Ferrari, 1996; Ferrari et al.,
2002; Silva Jr. e Queiroz, no prelo). Durante as atividades
de censo e monitoramento de fauna do projeto supracitado,
foram feitas tres observagoes da espcie em floresta continue
de terra fire na margem direita do lago. Na primeira oca-
siao, um animal solitirio foi observado. Na segunda, um
individuo foi visto juntamente com um grupo de Cebus
apella. Na terceira, foram gravadas vocaliza6oes caracteris-
ticas da espcie, mas nao foi possivel estimar o tamanho e a
composigao do grupo.

Apesar de existirem diversas unidades de conservagao
dentro da Area de ocorrencia de C kaapori, apenas na Re-
serva Biol6gica do Gurupi, no Estado do Maranhao, sua
presenga havia sido registrada. A falta de observagoes ante-
riores de C kaapori na drea de influencia da UHE-Tucurui
e o pequeno ndmero de observagoes realizadas em mais
de 900 km de trilha percorridos corroboram a hip6tese de
sua raridade natural (Lopes e Ferrari, 1993, 1996; Ferrari e
Queiroz, 1994; Ferrari e Lopes, 1996; Silva Jr. e Cerqueira,
1998; Carvalho Jr. et al., 1999; Carvalho Jr., 2003; Silva
Jr. e Queiroz, no prelo).


As observagoes foram feitas durante trabalhos de campo do
projeto "Avaliacao e monitoramento das comunidades de
vertebrados na Area de influencia do reservat6rio da UHE
Tucurui", com apoio financeiro da Eletronorte.


Brasil, ELETRONORTE. 1985. Plano de enchimento do
reservatdrio:fauna. Relatdriofinal. Secretaria Executiva de
Ciencia, Tecnologia e Meio Ambiente, Centrais Elktricas
do Norte do Brasil (ELETRONORTE), Belkm.

Neotropical Primates 14(2), August 2007

Brasil, ELETRONORTE. 2006. Mosaico de Unidades de
Conservacao do Lago de Tucurui Centrais Eltricas do
Norte do Brasil (ELETRONORTE), Belkm.
Brasil, SECTAM. (em preparacao). Biota Pard. Relatdrio
Anual. Secretaria Executiva de Ciencia, Tecnologia e
Meio Ambiente (SECTAM), Belkm.
Camargo, C. C. 2005. Ecologia comportamental de Alou-
atta belzebul (Linnaeus, 1766), naAmaz6nia Oriental sob
alteracao antr6pica de habitat. Dissertagao de Mestrado,
Universidade Federal do Pari, Belkm.
Carvalho Jr, 0. de. 2003. Primates in a forest fragment in
eastern Amazonia. Neotrop. Primates 11(2): 100-103.
Carvalho Jr, 0. de, Pinto, A. C. B. e Galetti, M. 1999.
New observations on Cebus kaapori in eastern Brazilian
Amazonia. Neotrop. Primates 7 (2): 41- 43.
Ferrari, S. F. 2003. Manejo das populagoes do cuxid, Chi-
ropotes satanas, um primata amaz6nico ameacado de ex-
tingao, na area de influencia da Usina Hidreltrica de Tu-
curui, Pard. Relat6rio tecnico a Fundagao 0 Boticirio de
Protegao a Natureza.
Ferrari, S. E, Ghilardi Jr., R., Lima, E. M., Pina, A. L. C.
B. e Martins, S. S. 2002. Mudangas a long prazo nas
populagoes de mamiferos da airea de influencia da Usina
Hidreltrica de Tucurui, Pard. Em: Resumos: XXIV Con-
gresso Brasileiro de Zoologia, pp. 540-541. Universidade
Federal de Santa Catarina, Itajaf, 17 a 22 de fevereiro,
Ferrari, S. E. e Lopes, M. A. 1996. Primate populations in
eastern Amazonia. Em: Adaptative Radiations of Neotro-
pical Primates, M. Norconk, A. L. Rosenberger e P. A.
Garber (eds.), pp.53-67. Plenum Press, New York.
Ferrari, S. F. e Queiroz, H. L. 1994. Two new Brazilian
primates discovered, endangered. Oryx 28: 31-36.
Lopes, M. A. 1993. Distribuigao, ecologia e conservagao
do cuxid-preto, Chiropotes satanas satanas (Cebidae, Pri-
mates) na Amaz6nia Oriental. Dissertagao de Mestrado,
Universidade Federal do Pari, Belem.
Lopes, M. A. e Ferrari, S. F 1993. Primate conservation in
eastern Brazilian Amazonia. Neotrop. Primates 1(4): 8-9.
Lopes, M. A. e Ferrari, S. F. 1996. Preliminary observations
on the Ka'apor capuchin Cebus kaapori Queiroz, 1992
from eastern Brazilian Amazonia. Biol. Conservation 76:
Mascarenhas, B. M. and Puorto, G. 1988. Nonvolant
mammals rescued at the Tucurui dam in the Brazilian
Amazon. Primate Conservation 9: 91-3.
Queiroz, H. L. 1992. A new species of capuchin monkey,
genus Cebus Erxleben, 1777 (Cebidae: Primates),
from eastern Brazilian Amazonica. Goeldiana Zoologia
Sampaio, D. T. 2004. Ecologia e comportamento de maca-
cos-prego (Cebus apella) num fragmento de floresta Ama-
z6nica. Dissertagao de Mestrado, Universidade Federal
do Pari, Bel6m.
Santos, R. R. 2002. Ecologia de cuxids (Chiropotes satanas)
na Amaz6nia Oriental: perspectives para a conservagao
de populagoes fragmentadas. Dissertagao de Mestrado,
Universidade Federal do Pari, Bel6m.

Silva Jr., J. S. 2001. Especiacao nos macacos-prego e caia-
raras, genero Cebus Erxleben, 1777 (Primates, Cebidae).
Tese de Doutoramento, Universidade Federal do Rio de
Janeiro, Rio de Janeiro.
SilvaJr., J. S. e Cerqueira, R. 1998. New data and a histori-
cal sketch on the geographical distribution of the Ka'apor
capuchin, Cebus kaapori Queiroz, 1992. Neotrop. Prima-
tes 6(4): 80-121.
Silva Jr., J. S. e Queiroz, H. L. (prelo). Cebus kaapori (Quei-
roz, 1992). Em: Livro vermelho das species ameafadas de
extinfao da fauna brasileira, A. B. M. Machado, G. M.
Drummond e A. P. Paglia (eds.), Fundagao Biodiversitas,
Belo Horizonte.
Silva, S. S. B. 2003. Comportamento alimentar de cuxid-
preto (Chiropotes satanas) na airea de influencia do reserva-
t6rio da usina hidreltrica de Tucurui-PA. Dissertagao de
Mestrado, Universidade Federal do Pari, Bel6m.
Veiga, L. M. 2006. Ecologia e comportamento do cuxid-
preto (Chiropotes satanas) na paisagem fragmentada da
Amaz6nia Oriental, Brasil. Tese de Doutoramento, Uni-
versidade Federal do Pari, Belem.
Vieira, T. M. 2005. Aspectos da ecologia do cuxid de Uta
Hick, Chiropotes utahickae (Hershkovitz, 1985), com
enfase na exploracao alimentar de esp6cies arb6reas da
ilha de Germoplasma, Tucurui-PA. Dissertagao de Mes-
trado, Universidade Federal do Pari, Belem.

Fernanda Cunha, Museu Paraense Emilio Goeldi, Belem,
PA, e-mail: , Maria Apare-
cida Lopes, Departamento de Biologia, Universidade
Federal do Pari, Belem, PA, e-mail: ,
Sidnei Dantas, Museu Paraense Emilio Goeldi, Belem, PA,
e-mail: , Nivia Carmo, Museu
Paraense Emilio Goeldi, Belem, PA, e-mail: museu-goeldi.br>, Suleima Silva, Museu Paraense Emilio
Goeldi, Belem, PA, e-mail: .


Thor Hanson

Tree bark has been reported as a minor dietary component
in many primate species, from western lowland gorillas
(Gorilla go l// go //,I) (Goldsmith, 1999) to red-handed
howler monkeys (Alouatta belzebul discolor) (Pinto and Setz,
2004). Barbary macaques (Macaca sylvanus) commonly
strip bark from cedar (Cedrus atlantica) trees in Morocco,
where research suggests they are seeking water or scarce
nutrients present in the cambium tissue (see Camperio-
Ciani et al., 2004). There is also mounting evidence of
primate self-medication, the use of medicinal plants to
treat a range of ailments from ectoparasites to intestinal
worms (reviewed in Huffman, 1997). Increased consump-
tion of known medicinal plants has been associated with
seasonal trends of nematode infection in chimpanzees (Pan

Neotropical Primates 14(2), August 2007

troglodytes), while non-nutritive bark and wood are used as
suspected purgatives by numerous great ape populations
(Huffman et al., 1997). In the Neotropics, Gottlieb et al.
(1996) noted a close higher-order similarity in the taxo-
nomic groups of plants utilized by humans and primates,
and also suggested a correlation between the choice of cer-
tain plant types and the medicinal properties of their sec-
ondary compounds.

White-faced capuchin monkeys (Cebus capucinus) inhabit
a wide range of low- to mid-elevation forest types from
Honduras to Ecuador (Reid, 1997). Their diet and behav-
ior are well documented at several locations (e.g., Oppen-
heimer, 1968; Buckley, 1983). They are known to remove
tree bark in search of insects (Rose, 1994) and to occasion-
ally ingest the bark of at least two tree species, Rhizophora
mangle and Byrsonima crassifolia (Buckley, 1983). Several
authors have watched C. capucinus engage in fur-rubbing,
the topical application of plants with known ethnome-
dicinal qualities in a possible attempt to treat ectoparasites
or other skin conditions (Oppenheimer, 1968; Buckley,
1983; Baker, 1996). Self-medication for intestinal parasites
has not been studied, but C. capucinus hosts a wide range
of intestinal worms and pathogens (Thatcher and Porter,
1968) and the barks of both R. mangle and B. crassifolia
are well-documented medicinals. B. crassifolia is a common
rural remedy for diarrhea (Heinrich, 2003) and has shown
efficacy in treating Giardia (Peraza-Sanchez et al., 2005),
while R. mangle is being studied as a treatment for gastro-
intestinal ulcers (e.g., Sanchez-Perera et al., 2004). Carapa
guianensis Aubl. (Meliaceae) is a widespread Neotropical
canopy tree, characteristic of lowland forests from Belize
south to the Amazon basin (Fournier, 2002). Its bark, seeds
and leaves have a wide range of ethnobotanical uses, from
fever reduction to the treatment of ulcers and skin condi-
tions (Schultes and Raffauf, 1990; Fournier, 2002). Most
notably, the bark tissue is used in a tea to expel intestinal
worms and parasites (Schultes and Raffauf, 1990).

On 7 March 2006, I observed an adult male C. capucinus
consuming bark from the trunk of a mature C. guianensis
(-90 cm dbh) in the southwestern portion of La Reserva
Indio-Maiz, Nicaragua (1056'18.4"N, 8419'54.3"W).
At least eight other members of the troop were foraging
nearby, but only a single individual was engaged in bark
stripping. Balancing in the crown of a small sub-canopy
tree, he used his weight to lean the tree against the straight
bole of an adjacent C. guianensis, approximately 10 m
above the ground. He then grasped at the smooth trunk
and used his teeth to strip off long pieces of bark, chewing
first the bark strips and then directly chewing the exposed
inner bark and cambium of the trunk itself. In more than
ten minutes of feeding he completely removed the bark
from an area approximately 0.75 m by 0.5 m, leaving the
cambium and sapwood exposed. The tree bore no signs of
insect infestation and the individual appeared to be con-
suming the inner bark and cambium specifically.

Consumption of C. guianensis bark by C. capucinus has
not been previously reported and authors of recent studies
in Panama and Costa Rica have not observed this behav-
ior (L. Fedigan, pers. comm.; E. Wehnke, pers. comm.).
It remains to be learned whether the event described here
is a local habit of the Indio-Maiz population, or a more
widespread behavior that is simply unusual and rarely seen.
Given the documented ethnobotanical uses of C guianensis
bark, as well as two other tree barks eaten by C capucinus,
this behavior should be looked for in other populations as a
potential new example of self-medication in primates.

Acknowledgment: This work was supported in part by NSF
IGERT Grant No. 0114304.

Thor Hanson, Department of Forest Resources, University
of Idaho, P.O. Box 441133, Moscow, Idaho 83844, USA,
e-mail: .


Baker, M. 1996. Fur rubbing: Use of medicinal plants by
capuchin monkeys (Cebus capucinus). Am. J Primatol.
38: 263-270.
Buckley, J. S. 1983. The feeding behavior, social behavior
and ecology of the white-faced capuchin monkey, Cebus
capucinus, at Trujillo, Northern Honduras, Central
America. Doctoral thesis, University of Texas, Austin.
Camperio-Ciani, A., Martinoli, L., Capiluppi, C., Arhou,
M. and Mouna, M. 2001. Effects of water availability
and habitat quality on bark stripping behavior in Bar-
bary macaques. Conserv. Biol. 15: 259-265.
Fournier, L. A. 2002. Carapa guianensis. In: Tropical Tree
Seed Manual, USDA Forest Service Agriculture Hand-
book 721, J. A. Vozzo (ed.), pp.360-363. USDA Forest
Service, Washington DC.
Goldsmith, M. L. 1999. Ecological constraints on the for-
aging effort of western gorillas (Gorilla gol, lla goaul/a) at
Bai Kokou, Central African Republic. Int. J Primatol.
20: 1-23.
Gottlieb, 0. R., Borin, M. R. and Bosisio, B. M. 1996.
Trends of plant use by humans and nonhuman primates
in Amazonia. Am. J Primatol. 40: 189-195.
Heinrich, M. 2003. Ethnobotany and natural products:
The search for new molecules, new treatments of old dis-
eases or a better understanding of indigenous cultures?
Cur. Topics Med. Chem. 3: 141-155.
Huffman, M. A. 1997. Current evidence for self-medica-
tion in primates: A multidisciplinary perspective. Ybk.
Phys. Anthrop. 40: 171-200.
Huffman, M. A., Gotoh, S., Turner, L. A., Hamai, M. and
Yoshida, K. 1997. Seasonal trends in intestinal nema-
tode infection and medicinal plant use among chimpan-
zees in the Mahale Mountains, Tanzania. Primates 38:
Oppenheimer, J. R. 1968. Behavior and ecology of the
white-faced capuchin monkey (Cebus capucinus) on

Neotropical Primates 14(2), August 2007

Barro Colorado Island. Ph.D. dissertation, University of
Illinois, Urbana.
Peraza-Sanchez, S. R., Poot-Kantdn, S., Toores-Tapia, L.
W., May-Pat, E, Sima-Polanco, P. and Cedillo-Rivera, R.
2005. Screening of native plants from Yucatan for anti-
Giardia lamblia activity. Pharm. Biol. 43: 594-598.
Pinto, L. P. and Setz, Z. E 2004. Diet of Alouatta belzebul
discolor in an Amazonian rain forest of Northern Mato
Grosso State, Brazil. Int. J Primatol. 25: 1197-1211.
Reid, E 1997. Field Guide to the Mammals of Central
America and Southwest Mexico. Oxford University Press,
Sanchez-Perera, L. M., Batista, N. Y., Rodriguez, A. and
Bulnes, C. 2004. Gastric and duodenal antiulcer effects
of Rhizophora mangle. Pharma. Biol. 42: 225-229.
Rose, L. M. 1994. Sex differences in diet and foraging
behavior in white-faced capuchins (Cebus capucinus).
Int. J. Primatol. 15: 95-114.
Schultes, R. E. and Raffauf, R. E 1990. The Healing
Forest: Medicinal and Toxic Plants of the Northwest Ama-
zonia (Historical, Ethno- and Economic Botany, Vol. 2).
Dioscorides Press, Portland.
Thatcher, V. E. and Porter, J. A. 1968. Some Helminth
parasites of Panamanian primates. Trans. Am. Micr. Soc.
87: 186-196.


Marcelo Marcelino de Oliveira
Marcos de Souza Fialho
Jflio Cesar Bicca-Marques

Foi realizada entire os dias 17 e 18 de abril de 2007, em
Sao Luis (MA), a I Reuniao do Comite Internacional para
Conservagao e Manejo dos Primatas Amaz6nicos. Partici-
param desta reuniao Onildo Marini Filho e Juciara Pelles
(IBAMA/Coordenacao de Especies da Fauna), Marce-
lo Marcelino, Marcos Fialho e Juliana Gongalves Ferreira
(IBAMA/Centro de Protegao de Primatas Brasileiros), Bea-
triz Gomes e Roberto Veloso (IBAMA/Superintendencia do
Maranhao), Juliana Shiraishi (IBAMA/Coordenacao Geral
de Unidades de Conservagao), Fabiano Costa (IBAMA/Co-
ordenacao Geral de Florestas Nacionais e Reservas Equiva-
lentes), Wilson Spironello (Instituto Nacional de Pesquisas
da Amaz6nia), Marcelo Gordo (Universidade Federal do
Amazonas), Helder Queiroz (Instituto de Desenvolvimento
Sustentivel Mamiraui), Jos6 de Souza e Silva Jdnior (Museu
Paraense Emilio Goledi), Fernanda Marques (Wildlife Con-
servation Society), Deyse Campista (Sociedade de Zool6gi-
cos do Brasil), Alcides Pissinatti (Centro de Primatologia
do Rio de Janeiro), Anthony Rylands (Center for Applied

Biodiversity Science/Conservation International), DAlia
Rizel Nogueira (Jardim Zool6gico de Belo Horizonte), Jdlio
Cesar Bicca-Marques (Pontificia Universidade Cat61lica do
Rio Grande do Sul), Jean Philippe Boubli (Universidade de
Auckland), Cibele Bonvicino (Fundagao Instituto Oswaldo
Cruz), Rosana Subird (Secretaria de Defesa do Meio Am-
biente de Manaus), Mariluce Messias (Fundagao Univer-
sidade Federal de Rond6nia), Lilian Pinto (Universidade
Estadual de Campinas), Liza Veiga (Universidade Federal
do Pari), Ricardo Santos (Universidade Estadual do Mara-
nhao) e Adrian Barnett (Universidade de Roehampton). 0
Comite foi instituido pelo Ibama no final de 2005, como
6rgao de assessoramento para identificacao e proposigao de
medidas de manejo e conservagao para as especies de prima-
tas amaz6nicos presents na lista official da fauna brasileira

Foi definida a elaboracao de dois pianos de agao para 2008
com o objetivo de estabelecer medidas que sejam efetiva-
mente dteis para a conservagao das esp6cies. 0 primeiro
plano de agao contemplari o sauim-de-Manaus (Saguinus
bicolor), esp6cie considerada "criticamente em perigo" e
com irea de ocorrencia restrita ao municipio de Manaus e
arredores. 0 segundo plano contemplard quatro esp6cies: o
guariba (Alouatta ululata) e o caiarara (Cebus kaapori) "cri-
ticamente em perigo", o cuxid-preto (Chiropotes satanas)
"em perigo" e o cuxid (Chiropotes utahickae) "vulnerivel".
A situacao de C kaapori 6 especialmente preocupante pela
raridade de populagoes remanescentes na natureza em con-
seqiiencia da perda de habitat em sua drea de ocorrencia
nos estados do Maranhao e Pard e da capture illegal para o
com&rcio de animals de estimacao e caga.

Como prioridades de pesquisa foram definidos (1) inven-
tirios para as areas no sul dos estados do Pard e Amazonas
e no alto Solim6es visando ampliar o conhecimento acerca
dos limits de distribuigao de algumas das esp6cies amea-
gadas de extinygo que deverno ser alvo de pianos de agno a
partir de 2009, tais como os uacaris (Cacajao calvus calvus
e Cacajao calvus rubicundus), (2) estudos sobre as especies
"dados deficientes" que podem estar sob risco de extingao
em decorrencia de sua presenga em dreas sob pressao de
desmatamento ou de instalagao de usinas hidrel6tricas ou
cuja ocorrencia no territ6rio brasileira 6 incerta, tais como
Saguinus fuscicolis cruzlimai e Saguinus fuscicolis crandalli.
0 Comit& tambdm recomendou que sejam mantidas po-
pulagoes em cativeiro de Cebus kaapori, Ateles marginatus,
Saguinus bicolor e Chiropotes satanas como "poupangas ge-
n&ticas" para possiveis futures programs de reintrodugao.

Houve especial preocupagao com a necessidade de uma
melhor orientagao para a destinacao de primatas resgatados
de areas inundadas em projetos de implantagao de usinas
hidrel6tricas. Um document com sugest6es seri elaborado
pelo Comite para a Diretoria de Fauna e Recursos Pesquei-
ros do Ibama. Devido a relevincia do tema, o Comite re-
comendou que o impact sobre as populagoes de primatas
seja alvo de pesquisas de long prazo nas areas que serao

Neotropical Primates 14(2), August 2007

afetadas por futures projetos hidrel6tricos. Como exemplo,
citou-se a operaiao de resgate de fauna da Usina Hidre-
16trica de Tucuruf, na qual foram resgatados e soltos nas
margens do reservat6rio 19.496 guaribas-de-mao-vermelha
e 2.580 macacos-pregos, cujo impact sobre a estrutura da
comunidade 6 desconhecido.

Em sua pr6xima reuniao, o Comite deveri discutir a influ-
encia da caga sobre as populagoes de primatas amaz6nicos e
a identificaiao de medidas que possam reduzir sua pressao,
bem como o efeito do aquecimento global sobre algumas
esp6cies ameagadas, especialmente aquelas que habitam
os ambientes de virzea. Modelagens matemaiticas apon-
tam para uma redugao dristica das populagoes de algumas
destas espcies em decorrencia das mudangas climiticas.

Marcelo Marcelino de Oliveira e Marcos de Souza Fialho,
IBAMA-Centro de Protegio de Primatas Brasileiros, Praga
Anthenor Navarro 5, Varadouro, Joao Pessoa, Paraiba
58010-480, Brasil, e-mail: .

Jilio C6sar Bicca-Marques, Laborat6rio de Primatologia,
Faculdade de Bioci&ncias, Pontificia Universidade Cat61lica
do Rio Grande do Sul, Avenida Ipiranga 6681 Prndio 12A,
Porto Alegre, Rio Grande do Sul 90619-900, Brasil, e-mail:


Joseph W. Kemnitz, Ph.D.

The Wisconsin National Primate Research Center
(WNPRC) is pleased to announce the establishment of the
Lawrence Jacobsen WNPRC Conservation Research Award.
The award supports studies in applied conservation biology
that protect non-human primate species and their habitat.
The WNPRC at the University of Wisconsin-Madison has
a long history of supporting work in primate conservation
that has significantly impacted the survival of many pri-
mate species and the protection of forested habitat. The late
J. Stephan Gartlan began the WNPRC's pioneering primate
conservation studies in Cameroon and today the center
continues to make significant strides in helping to protect
the world's most threatened primates and their habitats. The
award is available to students and/or faculty working in the
field of primate conservation. Applicants must be affiliated
with a university, college or non-governmental organization
(NGO). Students and/or researchers from all countries are
welcome to apply. Preference will be given to those working
directly with a nonhuman primate species that IUCN lists
as threatened or endangered. For more information (includ-
ing the online application) see edu/jacobsen>. Grant applications for this year should be
received no later than August 1, 2007. Questions about the
application can be directed to Joseph W. Kemnitz, Ph.D.,
e-mail: .


La Estaci6n Biol6gica Corrientes y el Museo Argentino
de Ciencias Naturales invitan al curso te6rico de posgra-
do de Ecologia y Comportamiento de Primates, Visidn Cld-
sica y Nuevos Enfoques, el cual se llevard a cabo del 15 al
23 de Septiembre de 2007 en las instalaciones de la Esta-
ci6n Biol6gica Corrientes. El curso consiste en la lectura y
discusi6n de articulos sobre primatologia en inglks, por lo
cual es requisite indispensable el buen manejo del idioma.
Ademis durante el curso se realizard una salida al campo. El
curso cubrird los siguientes temas: Introducci6n del Orden
primates, diversidad, patrons de organizaci6n social, sis-
temas de apareamiento; Modelos actuales que explican
la organizaci6n social de primates no-humanos: modelo
socioecol6gico, modelo de constrefimientos ecol6gicos,
modelo de interacciones cooperatives; Estado actual de la
discusi6n te6rica en el campo de la Primatologia; Ejemplo
de trabajos de campo en diferentes lugares del mundo y
Argentina; Efectos de la deforestaci6n sobre los patrons
comportamentales de los primates no humans; Fragmen-
taci6n de habitat, efectos de la alteraci6n del habitat en la
relaci6n parisito/hospedador; Contribuci6n de la gen&tica
molecular en la conservaci6n, consecuencias gen&ticas de
poblaciones pequefas; Corredores biol6gicos, traslocaci6n
y conservaci6n ex situ; Diversidad de primates en Argen-
tina y su estado de conservaci6n. Para mayor informaci6n
visitar: .
Cualquier duda consultar con Martin Kowalewski, e-mail:


O Program Trainee em Meio Ambiente da Fundacao
O Boticirio de ProtegAo a Natureza chega a sua quinta
ediiAo. Ap6s acompanhar por virios anos a realidade am-
biental no Brasil e a atuaiAo dos profissionais na drea de
conservagio da natureza, a Fundagao 0 Boticirio langou,
em 2003, o Programa Trainee em Meio Ambiente. Com
a intencAo de contribuir para o desenvolvimiento das
instituiy6es participants do program, a Fundagio 0
Boticario ofrece is ..-: i,-i 1,:. da sociedade civil da
irea de conservagio da natureza e empresas que reali-
zem aS6es de meio ambiente selecionadas, um program
de formaiAo de profissionais que protagonizem solug6es
para a conservagio da natureza. Os trainees e tutores
indicados pelas instituiy6es participam durante um ano
de encontros presenciais com a equipe da Fundagio 0
Boticirio, al6m de desenvolverem um projeto institucio-
nal. As inscriy6es para a turma de 2008 estio abiertas.
Para saber mais, acesse o site da Fundagio 0 Boticirio
ou entire em contact:

Neotropical Primates 14(2), August 2007


Na iltima d6cada o Curso Latino Americano de Biologia da
Conservagao e Manejo da Vida Silvestre vem contribuindo
para a formacao de profissionais comprometidos com a con-
servagao da diversidade socioambiental da Am&rica Latina.
Durante as cinco semanas de capacitagao os participants
compartilham experiencias com renomados profissionais
que atuam nas diferentes esferas da Biologia da Conserva-
gao. 0 conteddo programitico deste curso trata primeira-
mente de uma abordagem conceitual te6rica, seguido de
metodologias e ferramentas iteis em estudos de campo e
por l1timo, estudos de caso nos quais os alunos vivenciarao
a realidade de um program de conservagao na regiao do
Pontal do Paranapanema. As aulas serao ministradas na sede
do IPE, em Nazar6 Paulista- SP e no municipio de Teodoro
Sampaio -SP, no period de 06 de novembro a 12 de dezem-
bro de 2007. Mais informac6es: .


Dr. Barbara M. Thiers, Director, New York
Botanical Garden (NYBG)

This Catalogue will contain information from more than
90,000 specimens housed in The New York Botanical
Garden Herbarium that represent vascular plant species oc-
curring in Central and Northeastern Brazil. Specimens from
the Distrito Federal and the following states are included:
Ceari, Goias, Maranhao, Mato Grosso, Mato Grosso do
Sul, Minas Gerais, Piaul, and Tocantins. In addition, a dig-
ital image of one or two specimens of each species have been
captured and attached to the database record. The collec-
tion locality of any specimen with geographic coordinates
can be viewed on a map. The information in this catalogue
is intended to provide a means to assess the geographic dis-
tributions of species in this region and the diversity and en-
demism of the flora. This Catalogue will present one of the
jewels of the NYBG Herbarium-the set of about 42,000
specimens collected by New York Botanical Garden staff
members, Howard S. Irwin and William R. Anderson, and
several collaborators during a series of expeditions to the Bra-
zilian Planalto funded by the National Science Foundation
from 1964-1975 and carried out in collaboration with the
University of Brasilia. At the time of the Planalto program,
the flora of the Planalto was relatively unknown, and many
new species were discovered, most represented by types in
the NYBG Herbarium. Creating this Catalogue is expected
to take three years, after which it will be updated regularly
with records of new specimens from the region accessioned
at the NYBG Herbarium and with any changes to records al-
ready in the Catalogue. For more information go to: sciweb.nybg.org/science2/hcol/planalto/index.asp>.


The Conservation Breeding Specialist Group (CBSG) an-
nounce the launch of its new website, which is the result of
another collaboration between CBSG, Linda Malek, and
Evenson Design Group-the same design firm that has
donated the design of the Annual Reports. Visit the new
website at where you will find all
the same reports, newsletters and other documents, and
new information on our activities.



The IUCN/SSC Reintroduction Specialist Group (RSG)
and Lincoln Park Zoo will host the 1st International Wild-
life Reintroduction Conference April 15-16, 2008 in Chi-
cago, IL, USA. The theme of the conference will be "Re-
introduction programs: Applying science to conservation".
Twenty speakers and 40 posters will be selected for presen-
tation in addition to 12 already invited speakers. Registra-
tion is limited to 275 participants. More information about
the conference theme and topics is available at the website
. We hope you will join us for
this exciting event.


The objectives of the Amazon Protected Areas Network
(RANPA) are to update the information about protected
areas in the Amazon basin, using a standard process; to es-
tablish an agreement to exchange experiences and infor-
mation between the managers of the regional protected
areas; to use standards of metadata to describe, evaluate
and predict the state of the protected areas according to
international standards; and to identify the areas that need
financial support in order to improve their role in the con-
servation of the biodiversity in the Amazon region. The
services that RANPA provides are a satellite image bank,
a fire alert system, and databases that include complete
information of the Amazon protected areas, publications
and maps. The RANPA net has the support of organiza-
tions from Venezuela, Perd, Colombia, Ecuador, Guyana
and Brazil. For more information about RANPA visit
or contact Alvaro Espinel (Senior
Environmental Information Specialist, Sustainable Devel-
opment Department, Organization of American States) at


IPS Research Grants: Alison Behie (Canada) "The roles of
nutrition, stress and parasites in determining population
density in black howlers"; Sarah Carnegie (Canada) "Re-
productive strategies and hormonal patterns in wild female
white-faced capuchins"; Marietta Dindo (UK) "Investigat-
ing primate social learning and culture"; Kathelijne Koops
(UK) "Elementary technology of foraging and shelter in the
chimpanzees of Nimba Mountains, Guinea"; Marie Pele
(France) "What is the influence of psychological traits in the
ability of macaques to delay gratification?"; Fiona Stewart
(UK) "The evolution of shelter: Modelling human origins
through field study of chimpanzee nest building"; Michael
Wasserman (USA) "The role of phytoestrogens in the feed-
ing ecology of red colobus monkeys". Charles Southwick
Conservation Education Commitment Award: Jerry Akpara-
wa, CERCOPAN. Lawrence Jacobsen Education Development
Award: Christos Astaras (University of Goettingen) "Raising
Awareness About Drill Conservation Status Among Youth
at Korup Region, Southwest Cameroon"; Marina Cords
(Columbia University) "Kakamega Environmental Educa-
tion Program: Building a Conservation Education Center at
Kibiri"; Damodar Gaire (Institute of Forestry, Nepal) "Cre-
ation of Community Awareness on Primate Conservation
among the School Students and Indigenous People in the
Buffer Zone of Bardia National Park, Nepal". IPS Captive
Care Grants: Sagan Friant "An Investigation of the Gastro-
intestinal Parasites in Wild and Captive Cercopithecine Pri-
mates of Southern Nigeria"; Natalia Ceballos-Mago "Survey
of Margarita capuchin monkey, Cebus apella margaritae
and other monkey species in captivity on Margarita Island,
Venezuela" Doug Cress "Pan African Sanctuary Alliance
(PASA) Population Project / Chimfunshi, Zambia". IPS
Conservation Grants: Abby Baird "RAPID Development
of playback for rapid population assessment of the critically
endangered brown-headed spider monkey (Atelesfusciceps)
in Ecuador"- Catherine Cooke "An assessment of primate
species abundance and habitat use in Sette Cama and south
Loango Park, Gabon"; Antje Engelhart "Reproductive bi-
ology of wild Sulawesi-crested black macaques (Macaca
nigra) in the Tangkoko-Batuangus Nature Reserve"- Nguyen
Man Ha "Primate Survey, with special focus on Hatinh
langur (T .'.-'..... laotum hatinhensis), red-shanked douc
(Ti .-'. nemaeus) and white cheeked gibbon (Nomascus
leucogenys) in Huong Hoa district, Quang Tri province, Viet-
nam"; Stacy Lindshield "Bridging Conservation and Devel-
opment: Applied Primate Conservation in the Talamanca
Region, Costa Rica"; Alexandre Nascimento "Black-faced
lion tamarin (Leontopithecus caissara) Conservation Program:
Implementing Action Plan through fiIl. ng data gaps Felicia
Ruperti "Population density and habitat preferences of the
Sahamalaza sportive lemur (Lepilemur sahamalazensis)".
Martha J. Galante Award: Martin Kowalewski.

Neotropical Primates 14(2), August 2007


Faces na Floresta (Faces in the Forest: The Endangered Muriq-
ui Monkeys of Brazil), by Karen B. Strier. Portuguese trans-
lation by Luiz Roberto Mendes Gongalves and Thaiis Costa;
preface by Miriam Leitao. Rio de Janeiro: Sociedade para
a Preservagao do Muriqui -"Preserve Muriqui", 2007.
ISBN: 978-85-7650-101-5. If interested in obtaining a
copy of the Portuguese version, contact: hotmail.om.br>.

English version: Karen Strier, 1999. Harvard University
Press. 170 pp. ISBN: 978-0674290082. The woolly spider
monkey, or muriqui, is one of the most threatened pri-
mate species in the world. Because of deforestation in their
natural habitat (the Atlantic coastal forests of southeastern
Brazil) the muriquis are confined to less than three per-
cent of their original range. As of 1987, there were only a
dozen forest fragments known to support a total muriqui
population of about 500. As of 1998, at least 20 forests are
known to support at least 1000 muriquis. This book traces
the natural history of the muriqui from its scientific disco-
very in 1806 to its current, highly endangered status. Karen
Strier provides a case study of this scientifically important
primate species by balancing field research and ecological
issues. Through her accessible presentation, readers gain
a broad understanding of primate behavior and tropical

Life in the Cerrado, a South American Tropical Seasonal
Ecosystem, by Gerhard Gottsberger and I. Silberbauer-
Gottsberger. 2005. The book describes the Cerrado (one
of the top biodiversity hotspots in the world), a large scale
South American ecosystem, its formation and origin, its
plants and their adaptations, their rhythms of life, and
their interactions with animals. The Central Brazilian Cer-
rado should be considered as a distinct vegetation type,
distinguished from other topographically similar Central
and South American vegetation types by its ecology, spe-
cies composition and floristic diversity. Cerrado occurs
frequently in savanna-like forms, but also as forest (closed
arboreal canopy), woodland (open arboreal canopy), scrub
and open grassland forms. Floristic similarities of Cerrado
and Central and South American savannas and savanna-
like vegetation are the result of a common origin of all these
vegetation types and also testify to the floristic exchange
between Neotropical savannas and Cerrado during the
Tertiary and Quaternary. In the first volume is a detailed
examination of its vegetation, its structure, dynamics and
presumed origin. Emphasis is on the adaptational features
of plants in relation to their physical environment, in par-
ticular climate, soil conditions and fire. Further, the utility
to humans of Cerrado plants is discussed, as well as the
influence of humans upon this ecosystem. We also discuss

Neotropical Primates 14(2), August 2007

some of the issues associated with conservation politics. In
the second volume, pollination and seed dispersal phenom-
ena are described, and emphasis is placed on how these pro-
cesses, essentially plant-animal relationships, are critical to
the maintenance and regeneration of this ecosystem. Con-
tents: Vol. 1. Origin, Structure, Dynamics and Plant Use;
Vol. 2. Pollination and Seed Dispersal. For more informa-
tion and ordering of the book contact: Prof. Dr. Gerhard
Gottsberger .

The Mammals of Costa Rica: A Natural History and Field
Guide, by Mark Wainwright and Oscar Arias. 2007. Cor-
nell University Press. ISBN: 978-0801473753. From the
raucous mantled howler monkeys and the charismatic
white-nosed coatis to the elusive jaguar, The Mammals of
Costa Rica offers authoritative accounts of the fascinating
creatures of the Neotropics. With more than four hundred
spectacular illustrations and a wealth of detailed informa-
tion drawn from firsthand observation, new research, and
synthesis of the scientific literature, this book describes all
of Costa Rica's readily identifiable terrestrial and freshwater
mammals. The clear and entertaining text is perfectly suited
to meet the needs of naturalists, students, and researchers,
as well as both experienced and first-time visitors to Costa
Rica and the American tropics. The mammal descriptions
include key identification features, range maps, vocaliza-
tions, local folklore and mythology, and comprehensive
information about natural history and conservation. The
color illustrations show not only the mammals themselves
but also their tracks, foods, and skulls. Also included are
illustrations of numerous other animals and plants with
which the mammals have close ecological links. By present-
ing mammals in a broader context, The Mammals of Costa
Rica provides an entry point into a general study of tropical
ecology and conservation.

Feeding Ecology in Apes and Other Primates. Ecological, Phys-
iological and Behavioural Aspects, edited by G. Hohmann,
M. M. Robbins and C. Boesch. 2006. Cambridge Univer-
sity Press. 540 pp. ISBN: 978-0521858373. The book fo-
cuses on evolutionary perspectives of the complex interac-
tions between the environment, food sources, physiology
and behaviour in primates. This highly interdisciplinary
volume provides a benchmark to assess dietary alterations
that affected human evolution by putting the focus on the
diet of hominid primates. It also offers a new perspective
on the behavioral ecology of the last common ances-
tor by integrating corresponding information from both
human and non-human primates. The potential of inno-
vations of applied biotechnology are also explored to set
new standards for future research on feeding ecology, and
new information on feeding ecology in humans, apes and
other primates is synthesized to help refine or modify cur-
rent models of socioecology. By taking a comparative view,
this book will be interesting to primatologists, anthropolo-
gists, behavioral ecologists and evolutionary biologists
who want to understand better non-human primates, and
the primate that is us. Contents: 1. Variability of the feed-

ing ecology of eastern gorillas M. M. Robbins, J. Bosco
Nkurunungi and A. McNeilage; 2. Sympatric western go-
rilla and mangabey diet D. M. Doran-Sheehy, N. F Shah
and L. A. Heimbauer; 3. Effects of fruit scarcity on forag-
ing strategies of sympatric gorillas and chimpanzees J.
Yamagiwa and A. K. Basabose; 4. Chimpanzee feeding
ecology and comparisons with sympatric gorillas in the
Goualougo Triangle, Republic of Congo D. Morgan and
C. Sanz; 5. Frugivory and gregariousness of Salonga bono-
bos and Gashaka chimpanzees G. Hohmann, A. Fowler,
V. Sommer and S. Ortmann; 6. Feeding ecology of savanna
chimpanzees at Fongoli, Senegal J. D. Pruetz; 7. Food
choice in Tai chimpanzees: are cultural differences present?
- C. Boesch, Z. B. Gond Bi, D. Anderson and D. Stahl;
8. The effects of food size, rarity, and processing complexity
on white-faced capuchins' visual attention to foraging con-
specifics S. Perry and J. C. Ordofiez; Part II. 9. Primate
foraging adaptations: two research strategies S. A. Alt-
mann; 10. The predictive power of socioecological models:
a reconsideration of resource characteristics, agonism, and
dominance hierarchies A. Koenig and C. Borries; 11.
Hunger and aggression in capuchin monkeys C. Janson
and E. Vogel; 12. How does food availability limit the
population density of white-bearded gibbons? A. J. Mar-
shall and M. Leighton; 13. Influence of fruit availability
on Sumatran orangutan sociality and reproduction S. A.
Wich, M. L. Geurts, T. M. Setia and S. S. Utami-Atmoko;
14. Central place provisioning: the Hadza as an example
- FW. Marlowe; Part III. 15. Estimating the quality and
composition of wild animal diets S. Ortmann, B. J. Brad-
ley, C. Stolter and J. U. Ganzhorn; 16. The possible ap-
plication of novel marker methods for estimating dietary
intake and nutritive value in primates R. W. Mayes; 17.
Energy intake by wild chimpanzees and orangutans N. L.
Conklin-Brittain, C. D. Knott and R.W. Wrangham; 18.
The role of sugar in diet selection in redtail and red colo-
bus monkeys L. Danish, C. A. Chapman, M. B. Hall,
K. D. Rode and C. O'Driscoll; 19. Primate sensory systems
and foraging behavior N. J. Dominy, P. W. Lucas and
N. S. Noor.

New Perspectives in the Study of Mesoamerican Primates:
Distribution, Ecology, Behavior, and Conservation, edited
by Alejandro Estrada, Paul A. Garber, Mary Pavelka, and
Leandra Luecke. 2006. Springer, New York. xvi + 600pp.
ISBN-10: 0-387-25854-X (hardback, US$139.00). The
ninth volume in the series Developments in P '..I
Progress and Prospects, Series Editor Russell H. Tuttle, Uni-
versity of Chicago, Chicago, Illinois. A timely overview
of the taxonomy and biogeography of approximately 21
primates of the genera Saguinus, Saimiri, Cebus, Aotus,
Alouatta and Ateles which occur in Mesoamerica and
northwestern Colombia, and of the behavior, ecology and
conservation of Mesoamerican Cebus, Alouatta and Ateles.
Following a summary review by the editors ("Overview
of the Mesoamerican primate fauna, primate studies and
conservation concerns", pp.1-22), the book has a further
22 chapters divided into five parts (each with a short intro-

Neotropical Primates 14(2), August 2007

duction) as follows. Part 1. Taxonomy and Biogeography.
Chapter 2. Taxonomy and distributions of Mesoamerican
primates A. B. Rylands, C. P. Groves, R. A. Mittermeier,
L. Cortes-Ortiz and J. J. H. Hines, pp.29-79; Chapter
3. The biogeographic history of Mesoamerican primates
- S. M. Ford, pp.81-114. Part 2. Population Responses
to Disturbance. Chapter 4. Demographic features of Al-
ouatta pigra populations in extensive and fragmented for-
ests S. Van Belle and A. Estrada, pp.121-142; Chapter
5. Population structure of black howlers (Alouatta pigra)
in southern Belize and response to Hurricane Iris M. S.
M. Pavelka and C. A. Chapman, pp.143-163; Chapter 6.
The effects of forest fragment age, isolation, size, habitat
type, and water availability on monkey density in a tropi-
cal dry forest H. N. DeGama-Blanchet and L. M. Fedi-
gan, pp.165-188; Chapter 7. Forest fragmentation and its
effects on the feeding ecology of black howlers (Alouatta
pigra) from the Calakmul area in Mexico A. Rivera and
S. Calmd, pp.189-213; Chapter 8. Intestinal parasitic in-
fections in Alouatta pigra in tropical rainforest in Lacan-
dona, Chiapas, Mexico: Implications for behavioral ecol-
ogy and conservation K. E. Stoner and A. G. di Pierro,
pp.215-240. Part 3. Behavior and Ecology. Chapter 9. Av-
erage body weight for mantled howling monkeys (Alouatta
palliata): An assessment of average values and variability,
K. E. Glander, pp.247-263; Chapter 10. An exploratory
analysis of developmental plasticity in Costa Rican man-
tled howler monkeys (Alouatta palliata palliata) C. B.
Jones, pp.265-285; Chapter 11. Travel patterns and spatial
mapping in Nicaraguan mantled howler monkeys (Alouat-
tapalliata) P. A. Garber and P. E. Jelinek, pp.287-309;
Chapter 12. Use of landmark cues to locate feeding sites in
wild capuchin monkeys (Cebus capucinus): An experimen-
tal field study P. A. Garber and E. Brown, pp.311-332;
Chapter 13. Leap, bridge or ride? Ontogenetic influences
on positional behavior in Cebus and Alouatta M. E Be-
zanson, pp.333-348; Chapter 14. Food choice by juvenile
capuchin monkeys (Cebus capucinus) in a tropical dry forest
- K. C. MacKinnon, pp.349-365; Chapter 15. Why be
alpha male? Dominance and reproductive success in wild
white-faced capuchins (Cebus capucinus) K. M. Jack and
L. M. Fedigan, pp.367-386; Chapter 16. Post-conceptive
mating in white-faced capuchins, Cebus capucinus: Hor-
monal and sociosexual patterns of cycling, noncycling, and
pregnant females S. D. Carnegie, L. M. Fedigan and T.
E. Zeigler, pp.387-409. Part 4. Conservation and Man-
agement Policies. Chapter 17. Growth of a reintroduced
spider monkey (Ateles ._ ..r 'i i population on Barro Col-
orado Island, Panama K. Milton and M. E. Hopkins,
pp.417-435; Chapter 18. Primates in agroecosystems:
Conservation value of some agricultural practices in Meso-
american landscapes A. Estrada, J. Saenz, C. Harvey, E.
Naranjo, D. Munioz and M. Rosales-Meda, pp.437-470;
Chapter 19. Primate populations in the protected forests
of Maya archaeological sites in southern Mexico and Gua-
temala A. Estrada, S. van Belle, L. Luecke and M. Ro-
sales-Meda, pp.471-488; Chapter 20. Mapping primate
populations in the Yucatan Peninsula, Mexico: A first as-

sessment J. C. Serio-Silva, V. Rico-Gray and G. Ramos-
Fernandez, pp.489-511; Chapter 21. A metapopulation
approach to conserving the howler monkey in a highly
fragmented landscape in Los Tuxtlas, Mexico S. Mandu-
jano, L. A. Escobedo-Morales, R. Palacios-Silva, V. Arroyo-
Rodriguez, and E. M. Rodriguez-Toledo, pp.513-538;
Chapter 22. Quantifying fragmentation of black howler
(Alouatta pigra) habitat after Hurricane Iris (2001), south-
ern Belize S. M. Alexander, M. S. M. Pavelka and N. M.
Bywater, pp.539-560. Part 5. Synopsis and Perspectives.
Chapter 23. New perspectives in the study of Mesoameri-
can primates: Concluding comments and conservation
priorities P. A. Garber, A. Estrada and M. S. M. Pavelka,
pp.563-584. There are indices for species and for subjects.
Available from: Springer, 233 Spring Street, New York, NY
10013, USA, Tel.: 1-212-460-1500 or 1-800-SPRINGER,
Fax: 1-212-460-1575, e-mail: .
More information available on the Springer website at


Aguiar, L. M., Mellek, D. M., Abreu, K. C., Boscarato,
T. G., Bernadi, I. P., Miranda, J. M. D. and Passos,
E C. 2007. Sympatry between Alouatta caraya and
Alouatta clamitans and the rediscovery of free-ranging
potential hybrids in southern Brazil. Primates 48(3):
Anzures-Dadda, A. and Manson R. H. 2007. Patch- and
landscape-scale effects on howler monkey distribution and
abundance in rainforest fragments. Anim. Conser. 10(1):
Asensio, N., Cristobal-Azkarate, J., Dias, P. A. D., Vea, J. J.
and Rodriguez-Luna E. 2007. Foraging habits of Alouatta
palliata mexicana in three forest fragments. Folia Prim.
78(3): 141-153.
Asensio, N., Arroyo-Rodriguez, V. and Cristobal-Azkarate,
J. 2007. Feeding encounters between a group of howler
monkeys and white-nosed coatis in a small forest frag-
ment in Los Tuxtlas, Mexico. J. Tropical Ecol. 23(2):
Aureli, F and Schaffner, C. M. 2007. Aggression and con-
flict management at fusion in spider monkeys. Biology
Letters 3(2): 147-149.
Bezerra, B. M., Souto, A. S. and Schiel, N. 2007. Infanti-
cide and cannibalism in a free-ranging plurally breeding
group of common marmosets (C .-' jacchus). Am. J
Prim. 69(8): 945-952.
Boinski, S., Fragaszy, D. M., Ehmke, E. and Kauffman, L.
2007. Wild brown capuchins in Suriname exploit com-
binatorial manipulation for multiple functions. Am. J.
Phys. Anthropol. (Suppl 44): 77.
Bovendorp, R. S. and Galetti, M. 2007. Density and popu-
lation size of mammals introduced on a land-bridge island
in southeastern Brazil. Biol. Invasions 9(3): 353-357.
Camargo, C. C. and Ferrari, S. E 2007. Interactions be-
tween tayras (Eira barbara) and red-handed howlers

Neotropical Primates 14(2), August 2007

(Alouatta belzebul) in eastern Amazonia. Primates 48(2):
Campos, F, Manson, J. H. and Perry, S. 2007. Urine wash-
ing and sniffing in wild white-faced capuchins (Cebus
capucinus): testing functional hypotheses. Int. J Prim.
28(1): 55-72.
Caselli, C. B. and Setz, E. Z. 2007. Seasonality in long
calls by titi monkeys (Callicebus nigrifrons) in the Atlan-
tic Forest of southeast Brazil. Am. J Prim. 69(Suppl 1):
Clarke, M. R., Ford, R., Arden, D. and Glander, K. E.
2007. Survey of the howler population of La Pacifica,
2006. Am. J Prim. 69(Suppl 1): 55-56.
Cortes-Ortiz, L., Duda, T. E Jr., Canales-Espinosa, D.,
Garcia-Orduna, E, Rodriguez-Luna, E. and Bermin-
gham, E. 2007. Hybridization in large-bodied New
World primates. Genetics. Pgs: online.
Cristobal-Azkarate, J., Chavira, R., Boeck, L., Rodriguez-
Luna, E. and Vea, J. J. 2007. Glucocorticoid levels in free
ranging resident mantled howlers: a study of coping strat-
egies. Am. J Prim. 69(8). Pgs: 866-876.
Cunningham, E. and Janson, C. 2007. Integrating infor-
mation about location and value of resources by white-
faced saki monkeys (Pithecia pithecia). Anim. Cognition
Pgs: online(i-12).
De laTorre, S., Snowdon, C. T. and Yepez, P. 2007. Precious
variability: behavior, ecology, and conservation of pygmy
marmosets in Ecuador. Am. J Prim. 69(Suppl 1): 36.
Digweed, S. M., Fedigan, L. M. and Rendall, D. 2007.
Who cares who calls? Selective responses to the lost calls
of socially dominant group members in the white-faced
capuchin (Cebus capucinus). Am. J Prim. 69(7): 829-835.
Dirzo, R., Mendoza, E. and Ortiz, P. 2007. Size-related dif-
ferential seed predation in a heavily defaunated neotropi-
cal rain forest. Biotropica 39(3): 355-362.
Di Fiore, A., Fernandez-Duque, E. and Hurst, D. 2007.
Adult male replacement in socially monogamous equa-
torial saki monkeys (Pithecia aequatorialis). Folia Prim.
78(2): 88-98.
Di Fiore, A. and Suarez, S. A. 2007. Route-based travel and
shared routes in sympatric spider and woolly monkeys:
cognitive and evolutionary implications. Anim. Cogni-
tion. Pgs: online (1-13).
Garber, P. A. 2007. Primate locomotor behavior and ecol-
ogy. Primates in Perspective. Campbell C. J., Fuentes, A.,
MacKinnon K. C, Panger M., Bearder S. K., Editors.
New York: Oxford Univ Press. 2007. Pgs: 543-560.
Gunst, N., Boinski, S. and Fragaszy, D. M. 2007. Studying
how social content aids acquisition for foraging skills in
wild brown capuchins (Cebus apella) in Suriname. Am. J.
Prim. 69(Suppl 1): 76.
Hankerson, S., Pitts, M., Dietz, J. and Raboy, B. 2007. A
test of exploitative and interference competition in wild
golden-headed lion tamarins (Leontopithecus chrysomelas).
Am. J Prim. 69(Suppl 1): 57.
Huck, M., Roos, C. and Heymann, E. W. 2007. Spatio-ge-
netic population structure in mustached tamarins, Sagui-
nus mystax. Am. J Phys. Anthropol. 132(4): 576-583.

Janson, C. H. 2007. Experimental evidence for route in-
tegration and strategic planning in wild capuchin mon-
keys. Anim. Cognition Pgs: online(i-16).
Jones, C. B. and Van Cantfort, T. E. 2007. Multimo-
dal communication by male mantled howler monkeys
(Alouattapalliata) in sexual contexts: a descriptive analy-
sis. Folia Prim. 78(3): 166-185.
Kitchen, D. M., Kenney, S. and Wilson, R. E. 2007. Do
male black howler monkeys (Alouatta pigra) assess the
fighting ability of individual rivals during inter-group en-
counters? Am. J Phys. Anthropol. (Suppl 44): 144.
Kowalewski, M. and Garber, P. A. 2007. Mating promiscu-
ity, energetic, and reproductive strategies in black and
gold howler monkeys (Alouatta caraya). Am. J Phys. An-
thropol. (Suppl 44): 147.
Laska, M., Bauer, V. and Hernandez Salazar, L. T. 2007.
Self-anointing behavior in free-ranging spider monkeys
(Ateles. '....., i in Mexico. Primates 48(2): 160-163.
Lawrence, J. M. 2007. The pair bond in brown titi mon-
keys (Callicebus brunneus): male and female reproductive
interests. Am. J Phys. Anthropol. (Suppl 44): 151-152.
Lindshield, S. M. 2007. The density and distribution of
Ateles ..'r in a mosaic landscape at El Zota Biological
Field Station, Costa Rica. Master Abstracts 45(2): 603.
Ludwig, G., Aguiar, L. M., Miranda, J. M. D., Teixeira,
G. M., Svoboda, W. K., Malanski, L. S., Shiozawa,
M. M., Hilst, C. L. S., Navarro, I. T. and Passos, F C.
2007. Cougar predation on black-and-gold howlers
on Mutum Island, Southern Brazil. Int. J Prim. 28(1):
Luecke, L. G., Martinez-Mota, R. and Estrada, A. 2007.
Censuses of parapatric howler monkeys (Alouatta pallia-
ta and Alouatta pigra) in Pantanos de Centla Biosphere
Reserve, Tabasco, Mexico. Am. J Phys. Anthropol. (Suppl
44): 160.
Lynch Alfaro, J. W., Gutierrez-Espeleta, G., Olson, L. E.,
Ross, V., Neitzel, S., Sukraw, K., Drown, D. M. and
Alfaro, M. E. 2007. Patterns ofmtDNA genetic diversity
in capuchin monkeys at regional and local scales. Am. J
Phys. Anthropol. (Suppl 44): 161.
Lynch Alfaro, J. W. 2007. Subgrouping patterns in a
group of wild Cebus apella nigritus. Int J Prim. 28(2):
Matthews, L. J., Schmitt, C. A. 2007. Courtship behaviors
of genus Cebus: a test case for inferences from phylogeny.
Am. J Phys. Anthropol. (Suppl 44): 166.
McLaughlin, M. V., Fernandez-Duque, E. 2007. Spa-
tial distribution of feeding and sleeping trees in territo-
rial owl monkeys (Aotus azarai). Am. J Phys. Anthropol.
(Suppl 44): 169-170.
Mendes Pontes, A. R., Normande, I. C., Fernandes, A. C.
A., Rosas Ribeiro, P. F and Soares M. L. 2007. Fragmen-
tation causes rarity in common marmosets in the Atlan-
tic forest of northeastern Brazil. Biodiv. Conserv. 16(4):
Monteiro, R. V., Dietz, J. M., Beck, B. B., Baker, A. J.,
Martins, A. and Jansen, A. M. 2007. Prevalence and
intensity of intestinal helminths found in free-ranging

Neotropical Primates 14(2), August 2007

golden lion tamarins (Leontopithecus rosalia, Primates,
Callitrichidae) from Brazilian Atlantic forest. Veterina.
Pathol. 145(1-2): 77-85.
Mourthe, I. M., Guedes, D., Fidelis, J., Boubli, J. P.,
Mendes, S.L. and Strier, K.B. 2007. Ground use by
northern muriquis (Brachyteles .':.... .-.. .4. J. Prim.
69(6): 706-712.
Nunez-Iturri, G. and Howe, H. E 2007. Bushmeat and
the fate of trees with seeds dispersed by large primates
in a lowland rain forest in western Amazonia. Biotropica
39(3): 348-354.
Peker, S., Pave, R., Zunino G. E. and Kowalewski,
M. M. 2007. Observation of a birth in wild black and
gold howler monkeys (Alouatta caraya). Am. J. Phys. An-
thropol. (Suppl 44): 186.
Peres, C. A. and Palacios E. 2007. Basin-wide effects of
game harvest on vertebrate population densities in Ama-
zonian forests: implications for animal-mediated seed
dispersal. Biotropica 39(3): 304-315.
Porter, L. M. and Garber, P. A. 2007. Niche expansion of
a cryptic primate, Callimico goeldii, during polyspecific
associations. Am. J. Phys. Anthropol. (Suppl 44): 191.
Possamai, C. B., Young, R. J., Mendes, S. L. and Strier,
K. B. 2007. Socio-sexual behavior of female northern
muriquis (Brachyteles .',;..i. ..-.'.. "i Am. J Prim. 69(7):
Pozo-Montuy, G. And Serio-Silva, J. C. 2006. Move-
ment and resource use by a group of Alouatta pigra in
a forest fragment in Balancan, Mexico. Primates. 48(2):
Rakhovskaya, M. V., Fernandez-Duque, E. and Di Fiore,
A. 2007. The effects of demographic and ecological fac-
tors on territory size and ranging patterns of Argentine-
an owl monkeys (Aotus azarai). Am. J. Phys. Anthropol.
(Suppl 44): 195.
Righini, N. and Martinez-Mota, R. 2007. Spatial proxim-
ity and association patterns in four groups of wild black
howler monkeys (Alouatta pigra) in Southern Mexico.
Am. J. Phys. Anthropol. (Suppl 44): 199.
Rocha, V. J., Aguiar, L. M., Ludwig, G., Hilst, C. L. S.,
Teixeira, G. M., Svoboda, W. K., Shiozawa, M. M.,
Malanski, L. S., Navarro, I. T., Marino, J. H. F. and
Passos, F. C. 2007. Techniques and trap models for
capturing wild tufted capuchins. Int J. Prim. 28(1):
Rodman, P. S. and Bossuyt, E J. 2007. Fathers and step-
fathers: familial relations of old and new males within
groups of Callicebus brunneus in southeastern Peru. Am.
J Phys. Anthropol. (Suppl 44): 201.
Rodrigues, M. R. and Lindshield, S. L. 2007. Scratching
the surface: observations of tool use in wild spider mon-
keys. Am. J. Phys. Anthropol. (Suppl 44): 201-202.
Schmitt, C. A., Di Fiore, A., Link, A., Matthews, L. J.,
Montague, M. J., Derby, A. M., Hurst, D., Carrillo,
G., Sendall, C., Field, M. Y. and Fernandez-Duque E.
2007. Comparative ranging behavior of eight species of
primates in a western Amazonian rainforest. Am. J. Phys.
Anthropol. (Suppl 44): 208-209.

Sendall, C., Fernandez-Duque, E. and Di Fiore, A. 2007. A
preliminary study of mate-guarding in wild titi monkeys
(Callicebus discolor). Am. J. Phys. Anthropol. (Suppl 44):
Siani, J. M. and Dietz, J. M. 2007. Costs of providing
infant care in wild golden lion tamarins at Poco das Antas
Biological Reserve, Brazil. Am. J. Prim. 69(Suppl 1):
Suarez, S. S. 2007. Paternity, relatedness, and socio-re-
productive behavior in a population of wild red-bellied
tamarins (Saguinus labiatus). Dissertation. Abs. Intern.
A68(1): 244.
Taggart, A. 2007. Primate poaching in the Peruvian
Amazon. Australasian Prim. 18(1): 24.
Valero, A. and Byrne, R. W. 2007. Spider monkey rang-
ing patterns in Mexican subtropical forest: do travel
routes reflect planning?. Anim. Cognition. Pgs: online
Veiga, L. M. and Ferrari, S. F. 2007. Geophagy at ter-
mitaria by bearded sakis (Chiropotes satanas) in south-
eastern Brazilian Amazonia. Am. J. Prim. 69(7):
Verderane, M. P., Falotico, T., Resende, B. D., Labruna,
M. B., Izar, P. and Ottoni, E. B. 2007. Anting in a sem-
ifree-ranging group of Cebus apella. Int. J Prim. 28(1):
Vogel, E. R. and Janson, CH. 2007. Predicting the frequen-
cy of food-related agonism in white-faced capuchin mon-
keys (Cebus capucinus), using a novel focal-tree method.
Am. J. Prim. 69(5): 533-550.
Weghorst, J. A. 2007. High population density of black-
handed spider monkeys (Ateles ._... 7..r, i in Costa Rican
lowland wet forest. Primates 48(2): 108-116.
Welter, B. J., Koenig, W., Pietsch, M. and Adams, R. P.
2007. Feeding selectivity by mantled howler monkeys
(Alouatta palliata) in relation to leaf secondary chem-
istry in Hymenaea courbaril. J Chemical. Ecol. 33(6):
Wheeler, B. C. 2007. The function of alarm calling among
wild tufted capuchin monkeys (Cebus apella). Am. J. Phys.
Anthropol. (Suppl 44): 248.
Williams, S. H., Vinyard, C. J., Glander, K. E., Teaford,
M. E, Deffenbaugh, M. and Thompson, C. L. 2007.
EMG telemetry in free-ranging primates: pilot data from
howling monkeys (Alouattapalliata) at La Pacifica, Costa
Rica. Am. J. Prim 69(Suppl 1): 121-122.
Wolovich, C. K., Perea-Rodriguez, J. P. and Fernandez-Du-
que, E. 2007. Food sharing as a form of paternal care in
wild owl monkeys (Aotus azarai). Am. J Phys. Anthropol.
(Suppl 44): 252.
Wright, K. A. 2007. The relationship between locomotor
behavior and limb morphology in brown (Cebus apella)
and weeper (Cebus olivaceus) capuchins. Am. J. Prim.
69(7): 736-756.
Zucker, E. and Clarke M. 2007. Proximities to specific
adult males by adult female howling monkeys (Alouat-
ta palliata) in Costa Rica. Am. J. Prim. 69(Suppl 1):

Neotropical Primates 14(2), August 2007


Abstracts from the 2nd Congress of the European Federa-
tion for Primatology. Charles University, Prague, Septem-
ber 3-7, 2007.

Addessi, E., A. Mancini, L. Crescimbere and E. Visalberghi.
Do capuchin monkeys (Cebus apella) deal with tokens as
they do with real food?, p.4.
Albiach-Serrano, A., E Guillkn-Salazar and J. Call. Inhi-
bition of a prepotent response in mangabeys (Cercocebus
torquatus lunulatus), p.4.
Alvergne, A., D. Caillaud, M. Charpentier, E. Huchard,
L. Martinez and Raymond M. Inter-specific kin recogni-
tion: Are humans able to associate family relatives among
other primate species?, p.4.
Amici, E and J. Call. Response facilitation in the Great
Apes, p.5.
Amici, F., F. Aureli and J. Call. Inhibitory skills are associ-
ated with high levels of fission-fusion dynamics, p.5.
Andreoli, M., C. Spiezio and D. Grassi. Chimpanzee re-
lationships in captivity: all-male groups and mixed-sex
groups, p.6.
Antonacci, D., I. Norscia and E. Palagi. Aggressive and af-
finitive behaviours around the mating period in berenty
sifakas (Propithecus verreauxi): preliminary results, p.6.
Arora, N., D. Perwitasari-Farajallah, J. Pamungkas and
M. Kriitzen. Genetic identification of confiscated oran-
gutans for their release in the wild, p.6.
Asensio, N., A. H. Korstjens and E Aureli. Ranging costs in
wild spider monkeys, p.7.
Assahad, G., C. Neumann, K. Hammerschmidt, D. Per-
witasari-Farajallah and A. Enhelhardt. Rank dependent
differences in loud call frequency and structure in Su-
lawesi crested black macaques (Macaca nigra), p.7.
Bernede, L., A. S. Beresford, K. A. I. Nekaris and A. Guna-
wardene. Home range use by the red slender loris (Loris
tardigradus tardigradus) in Masmullah proposed forest
reserve, Sri Lanka, p.8.
Bissonnette, A. and C. Van Schaik. Competitive ability as
predictor of coalition success in barbary macaques, p.8.
Bottin, G., M. C. Huynen and T. Savini. Preliminary re-
sults on activity budget, feeding ecology and ranging
behavior of wild pigtailed macaques (Macaca nemestrina
leonina) in Khao Yai national park, Thailand, p.9.
Brauch, K., M. Heistermann, A. Engelhardt and J. K.
Hodges. Male reproductive success in free-ranging bar-
bary macaques: influence of male rank and female direct
mate choice, p.9.
Brijuer, J., J. Call and M. Tomasello. Inequity aversion in
great apes, p. 10.
Breuer, T., E. J. Strokes, R. Parnell, A. Robbins and M. M.
Robbins. Male life history patterns and reproductive suc-
cess in western gorillas insights from Mbeli Bai, Repub-
lic of Congo, p. 10.
Brotcorne, E, M. C. Huynen andT. Savini. Preliminary re-
sults on the behavioral ecology of a long-tailed macaque

(Macacafascicularis) population in disturbed urban habi-
tats, Bangkok, p. 10.
Brown, K. M. and K. A. Bard. Playful expressions and their
contexts in one year old chimpanzees, p. 11.
Burkart J. M. and C. Van Schaik. Are common marmosets
prosocial? Eliminating alternative hypotheses, p. 11.
Byrne, R. W. The primate mind in the wild, p.12.
Caillaud, D., J. Benavides, N. Menard and M. Raymond.
Spatial position of individuals within western lowland
gorilla groups: what can we learn about social relation-
ships, p.12.
Carrillo-B, G. A., J. J. Bravo, S. Martin and M. C. Huynen.
Preliminary approach of the community of primates in
the protected area of Oglan Alto, Arajuno-Pastaza, Ec-
uador, p. 13.
Carvalho, S., C. Sousa and T. Matsuzawa. The Nut-Crack-
er: bridging archaeology and primatology Chimpanzee
stone tool use in Bossou and Dieck6, Guinea, p.13.
Casiraghi, N. and M. Zizzo. Artistic expression in primates,
Cheyne, S. M. Gibbon feeding ecology and diet character-
istics, p.14.
Chu, K., K. H. Jung, S. T. Lee, E. C. Song, H. K. Park, D.
I. Sinn, J. M. Kim, M. Kim and J. K. Roh. Identification
of neurosphere-forming cells in the adult rhesus monkey
brain, p.14.
Van Coillie, S., P. Galbusera, A. Roedor, W. Schempp,
J. Stevens, M. Bruford and K. Leus. The effect of inbreed-
ing on infant mortality in captive bonobos (Pan panicus)
determined from DNA analysis of hair samples, p. 15.
Cordoni, G., D. Antonacci and E. Palagi. The question of
play: a comparison between chimpanzees (Pan troglodytes)
and lowland gorillas (Gorilla goll// godl/). p.15.
Custance, D., N. Rakotomalala and H. Rasamimanana.
Using the two-action method to test social transmission
in despotic ring-tailed lemurs, p. 15.
Dameirus, L. A. and C. P. Van Schaik. Planning in the
wild? A study of the cognitive abilities of wild orangu-
tans (Pongopymaeus) at Sumatra South Aceh, Indonesia,
Donati, G., N. Baldi, V. Morelli, J. U. Ganzhorn and S. M.
Borgognini-Tarli. Proximate cues and ultimate determi-
nants of brown lemur cathemerality in a humid littoral
and in a dry gallery forest of southern Madagascar, p.16.
Van Dognen, M., B. Meuleman, S. E. Koski, Z. Pereboom
and E. H. M. Sterck. Bonobo (Panpanicus) conflict man-
agment and reconciliation during potentially stressful
situations, p.17.
Douglas, P. H., P. S. Moore, S. Wimalasuriya, A. Guna-
wardene and K. A. I. Kekaris. Microhabitat variables in-
fluencing abundance and distribution of diurnal primates
(T ': '..... vetulus vetules and Macaca sinica aurifrons)
in a fragmented rainforest in southern Sri Lanka, p. 17.
Engelhardt, A., J. K. Hodges and M. Heistermann. Post-
conception mating in wild longtailed macaques (Macaca
fascicularis), p.18.
Engelhardt, A. and D. P. Farajallah. Reproductive biology
of Sulawesi black macaques (Macaca nigra), p.18.

Neotropical Primates 14(2), August 2007

Eschmann, C., K. A. I. Nekaris, P. H. Douglas, L. P. Bir-
kett and A. Gunawardene. A comparison of calling pat-
terns of purple-faced leaf monkeys (T '-.-.'..... vetulus
vetulus and T v. nestor) in Sri Lanka's west zone, p.19.
Ey, E., K. Hammerschmidt, D. Zinner and J. Fischer. In-
fluences of environmental factors on vocal communica-
tion in baboons, p.19.
Fernandez-Lazaro, G., F J. De Miguel Agueda and A. Lo-
pez-Goya. The behaviour of the Galago senegalensis under
semi-natural conditions: Factors affecting activity and
social behaviour, p.19.
Fiore, M. and B. Chiarelli. Homebox genes and their role
in the morphogenesis and the evolution of the human
brain, p. 20.
Fistarol, L., D. Grassi and C. Spiezio. The influence of
music on behaviour: a comparative study on two species
of nonhuman primates in captivity, p.20.
Foitova, I., B. Koubkova, V. Barus and W. Nurcahyo.
Record of the gapeworm Mammomonogamus laryngeus
in semi-wild population of sumatran orangutan (Pongo
pygmaeus abelii) in Indonesia, p.21.
Forrester, G. S. A multidimensional approach to investiga-
tions of primate communication, p.21.
Fowkes, S. Effects of food presentation and diet on special-
typical foraging and tool use behaviours in five captive
orangutan populations, p.21.
Fraiser, 0. N. and F Aureli. Reconciliation, consolation
and relationship quality in chimpanzees, p.22.
Ganas, J., S. Ortmann and M. M. Robbins. Food choice
strategies of mountain gorilla groups in Bwindi Impen-
etrable National Park, Uganda, p.22.
Garcia, C., P. C. Lee and L. Rosetta. Impact of social con-
text on female reproductive function in a group of captive
olive baboons (Papio anubis), p.23.
Gaspersic, M. and J. D. Pruetz. Savanna chimpanzees (Pan
troglodytes verus) and baobab fruits (Andasonia digitata):
investigation of percussive technology among three chim-
panzee communities in southeastern Senegal, p. 23.
Govoroff, N. Looking for Chimps: A social anthropolo-
gist's point of view, p. 24.
Grueter, C. C. Home range analysis of snub-nosed mon-
keys (Rhinopithecus bieti) at South Baima Snow Moun-
tain Nature Reserve, China, p.24.
Gupta, A. K. Ecodevelopment: an effective tool for primate
conservation, p.24.
Haag, L. Responses to decreasing fruit availability and di-
etary differences in white-bearded gibbons (Hylobates al-
bibarbis) and Bornean orangutans (Pongo pygmaeus wur-
mbii), p.25.
Hadi, S., T. Ziegler and J. K. Hodges. Groups structure
and physical characteristics of simakobu monkeys (Simias
concolor) on the Mentawai Island of Siberut, Indonesia,
Hanus, D. and J. Call. Chimpanzees infer the location of a
reward based on the effect of its weight, p.26.
Harrison, M. E. Adaptive strategies for the use of fall-back
foods in apes, p.26.

Hauster, B., T. Deschner and C. Boesche. Development
of a liquid chromatography-tandem mass spectometry
method for the quantification of endogenous steroids in
primate urine, p.26.
Hemelrijk, C. K., J. Wantia and K. Isler. The more males,
the more dominant are female primates, p.27.
Hesse, B., N. Schilling, M. S. Fischer and R. Fraber. Bipe-
dalism adaptations in the human lumbar back muscula-
ture?, p.27.
Higman, J. P., C. Ross, Y. Warren, M. Heistermann and
A. MacLarnon. Reduced reproductive function in wild
baboons related to natural consumption of plant com-
pounds, p.28.
Hill, R., A. Bentley and R. Dunbar. Network scaling re-
veals consistent pattern in hierarchical animal societies,
Hrubesch, C., S. Preuschoft and C. Van Schaik. Conserva-
tive choices in an innovation and transmission experi-
ment with captive chimpanzee groups, p.28.
Huchard, E., J. Benavides, M. Raymond, L. A. Knapp and
G. Cowlishaw. Evolution of sexual swellings: size and
shape under examination in a wild population of Chacma
baboons (Papio ursinus), p.29.
Humle, T. and C. T. Snowdon. The dynamics of socially
biased learning in the acquisition of a complex foraging
task in juvenile cottontop tamarins, p.29.
Hutchinson, J. E. and A. W Fletcher. Comparing the role
of alloparenting in apes and monkeys, p.30.
Hutchinson, J. E. and A. W. Fletcher. Using multi-dimen-
sional scaling to map spatial proximities in captive gorilla
groups, p.30.
Isler, K. and C. Van Schaik. Life history pace and brain size:
from correlation to causation, p.31.
Jankowski, F Human observer-primate relationship in the
field: interactionist approach of the habituation process,
Jebavy, L., P. Kubecek, M. Kuncova and L. Libichova. En-
vironmental enrichment and welfare of laboratory ma-
caques, p.31.
Joulian, E Human and primates in perspective: a team re-
search project in Europe and West Africa, p.32.
Kamiski, J., J. Call and M. Tomasello. False belief under-
standing in children (Homo sapiens) and chimpanzees
(Pan troglodytes), p.32.
Kappeler, P. The social organisation of sifakas (Propithecus
verreauxi) at Kirindy forest, p.32.
Koops, K., T. Humle, E. H. M. Sterck and T. Matsuzawa.
Ground-nesting in chimpanzees (Pan troglodytes verus) of
the Nimba Mountains, Guinea, West Africa: New find-
ings, p.33.
Koops, K., W C. McGrew, I. C. Gilby and R. W. Wrang-
ham. Nesting height selection in male and female chim-
panzees (Pan troglodytes schweinfurthii) in Kibale forest
NP, Uganda, p. 33.
Koski, S. E. and E. H. M. Sterck. Self-regarding consolers -
chimpanzee post-conflict third-party-initiated affiliation
benefits the affiliators, p.34.

Neotropical Primates 14(2), August 2007

Krause, C., M. S. Fischer and M. Schmidt. Influences of in
vivo joint load history on the form of the shoulder joint
in the cercopithecid primates, p.34.
Latinne, A., M. C. Huynen and T. Savini. Feeding ecology
and seed dispersal of pigtail macaques (Macaca nemest-
rina) in Khao Yai National Park, Thailand, p.35.
Leblan, V. Anthropological dimensions of spatial interac-
tions between chimpanzees and humans. A case study
from the Bok6 region, northwestern Guinea, p.35.
Legrain, L., L. Van Elsacker and J. A. Iscoa. How does a
bonobo mother (Pan paniscus) manage a conflict between
her son and one of her female coalition partners?, p.36.
Lehmann, H. and J. J.Bryson. Explaining the despotic /
egalitarian continuum: a new model, p.36.
Lehner, S., J. Burkart and C. Van Shaik. Innovation in Or-
angutans, p.37.
Lein, I., S. Treue and R. Teepe. EUPRIM-NET European
Primate Network: establishing specialized infrastructures
and procedures for biological and biomedical research,
Lhota, S., T. Junek, A. A. Kubena and L. Bartos. Use of
two specialized fingers in aye-ayes (Daubentonia mada-
gascariensis), p.38.
Lisensky, D. Frequency of play and antagonism in or-
phaned chimpanzees at Chimfunshi Wildlife Orphanage
Zambia, p.38.
Locatelli, S., F Liegeois, B. Lafay, A. D. Roeder, M.W. Bru-
ford, P.Formenty, R. Noe, E. Delaporte and M. Peeters.
Prevalence and genetic diversity of simian immunodefi-
ciency virus infection in wild living red colobus monkeys
(Piliocolobus badius badius) from the Tai Forest, Cote d'
Ivoire, p.38.
Lorch, D., P. M. Kappeler, M. Heistermann and L. Walter.
Parasite mediated sexual selection in primates? The effect
of intestinal parasites on the reproductive success of wild
lemurs (Eulemurfulvus rufus), p.39.
Liihrs, M. L., M. Dammhahn, C. Fichtel and P. M. Kap-
peler. Spatial memory in grey mouse lemurs (Microcebus
murinus), p.39.
Lutter, S. Interspecies communication with gorillas reveals
advantage cognitive abilities and thought processes: lon-
gitudinal study of acquisition and usage of American Sign
Language by western lowland gorillas, p.40.
MacLarnon, A. M., S. Semple, K. Shutt, M. Heistermann
and J. P. Higham. Sociality and stress in macaques and
baboons, p.40.
Maguire, V. and K. A. Bard. Gesture use in two differen-
tially reared groups of infant chimpanzees, p.40.
Maguire, V. and K. A. Bard. The differential use of gestures
in chimpanzees based on their intended audience, p.41.
Majolo, B., A. B. Vizioli and G. Schino. Group size ef-
fects on behavior and demography in primates: a meta-
analysis, p.41.
Maldonado, A. and S. Bearder. Assessing the impact of
hunting on harvest sensitive primates within protected
areas and indigenous land in the Colombian Amazon,

Manciocco, A. and A. Vitale. Effect on human interac-
tion on the behaviour of a colony of common marmosets
(Callithtixjacchus), p.42.
Martin-Carrera, 0. and F Guillen-Salazar. Do enclosures
in European zoo's guarantee safe public primate interac-
tions?, p.42.
Martin-Ordas, G., J. Call and F Colmenares. How great
apes perform in functionally equivalent trap tube and
trap table tasks, p.43.
Marvan, R., T. Polak, H. Marsault, J. Kantorova and
V. Vancata. A comparative analysis of positional behav-
iour of captive chimpanzees (Pan troglodytes): effect of
structural change in the enclosure, p.43.
Mass, V. and P. Kappeler. The role of subordinate males in
Propithecus verreauxi in the Kirindy forest, Madagascar,
Mevis, L., C. Richter, J. Ostner and 0. Schiilke. Post-con-
flict behaviour in free-ranging male stumptail macaques
(Macaca arctoides).
Morf, N., H. Morrogh-Bernard and M. Kriitzen. The util-
ity of real time polymerase chain reaction for molecular
studies on faecal samples, p.45.
Msindai, N. J. Identifying the economic and social impacts
of crop-raiding by non-human primates on local liveli-
hoods, in Mount Rungwe, southwest Tanzania, p.45.
Nater, A. and M. Kriitzen. Quantifying microsatellite
ascertainment bias between humans and orangutans,
Neel, C., E Van-Heuverswyn, Y. Li, B. Keele, J. Takehi-
sha, Y. Bienvenue, E. M. Ngole, E. Delaporte, B. Hahn
and M. Peeters. SIV infection in great apes: a multidisci-
plinary approach for better understanding of the origin,
history and transmission of SIV in wild gorillas in Cam-
eroon, p.46.
Nekaris, K. A. I. and V. Nijman. Survey on the abundance
and conservation of Sumatran slow lorises (Nycticebus
coucang .. i in Aceh, northern Sumatra, p.74.
Neumann, C., G. Asahad, K. Hammerschmidt, D. Per-
witasari-Farajallah and A. Engelhardt. Individual and
contextual differences in loud calls of male crested black
macaques, Macaca nigra, p.47.
Njiman, V, and K. A. I. Nekaris. Alarm calls, startle behav-
iour and predator avoidance in six congeneric arboreal
primates (Cercopithecidae: Presbytis), p.48.
Noel.Lambot, F, M. C. Huynen and T. Savani. An etho-
ecological survey of macaques in Thailand: preliminary
data, p.48.
Van-Noordwijk, M. and C. P. Van-Schaik. Food transfer
among non-kin in two natural populations of orangu-
tans, p.49.
Norscia, I., S. Kaburu, E. Palagi and D. Antonacci. Feeding
ecology of Berenty sifakas (Propithecus verreauxi) in the
wet season, p.49.
Nyakatura, J. A., M. S. Fischer and M. Schmidt. Tamarins
gait plasticity on oblique arboreal substrates, p.50.
Ostner, J., and 0. Schiilke. Male social relationships in
wild Assamese macaques (Macaca assamensis), p.50.

Neotropical Primates 14(2), August 2007

Petrasova, J., K. Pomajbikova, K. J. Petrzelkova, M. Jirku,
I. Profousova, D. Modry and C. Hashimoto. Parasites of
chimpanzees in Kalinzu forest reserve, Uganda, with em-
phasis on commensal protozoans, p.50.
Petru, M., M. Spinka, S. Lhota and P. Sipek. Head rota-
tions in the play of Hanuman langurs: a description and
an analysis of function, p.51.
Pfefferle, D., K. Brauch, M. Heistermann, K. Hodges and
J. Fischer. Do female copulation calls influence mating
outcome? A study in free ranging barbary macaques
(Macaca sylvanus), p. 51.
Pieta, K. Female mate choice in chimpanzees (Pan troglo-
dytes schweinfurthii) of Kanyawara, Kibale National Park,
Pomajbikova, K., K. J. Petrzelkova, T. Tokiwa, S. Imai and
D. Modry. The occurrence of the T-. ..,.-. abrassarti
in captive gorillas, p.52.
Port, M. and P. M. Kappeler. Grooming for aid or groom-
ing to stay? Patterns of social grooming in free-ranging
redfronted lemurs (Eulemurfulvus rufus), p.52.
Preuschoft, S., S. Peterhans and C. P. Van-Schaik. Food
sharing in groups: a comparison between captive chim-
panzees and recently socialised long-term isolates, p.53.
Puga-Gonzalez, I., C. Hilderbrandt and C. Hemelrijk.
Grooming patterns in primates: a model, p.53.
Quaresmini, C., C. Spiezio and D. Grassi. Hand reference
in captive nonhuman primates: Lemur catta and Macaca
nemestrina, p.54.
Van-Reeuwijk, N. and M. Kriitzen. Can ancient DNA aid
in documenting the loss of genetic diversity in orangu-
tans?, p.54.
Richter, C., L. Mevis, 0. Schiilke and J. Ostner. Social
relationships in free-ranging male stumptail macaques
(Macaca arctoides), p.54.
Riviello, M. C., A. Wirz, E. Haddis, F Natale, P. Poto,
G. Sabbatini, M. Saporiti, G. Spinozzi, V. Truppa and
E. Visalberghi. The primate center of the CNR (Rome,
Italy): research and school, p.55.
Von-Rohr, C. R. Tail marking and taxonomic status of the
Goeldi's monkey (Callimico goeldii), p.55.
Ruperti, F S., E P. M. Rabenandrasana, S. Renaudineau,
S. K. Bearder, N. Schwitzer and C. Schwitzer. A pilot
study of the Sahamalaza sportive lemur (Lepilemur sa-
hamalazensis). Population density, habitat requirements,
ethogram and vocalisations from four forest patches of
the Sahamalaza Peninsula, northwest Madagascar, p.56.
Ruperti, F S, E. L. Nelson and M. A. Novak. Bridging sci-
ence and tourism, a preliminary study of the black and
white ruffed lemur at Monkeyland Primate Sanctuary,
South Africa, p.56.
Sabbatini, G. and E. Visalberghi. Capuchin's use of visual
and auditory stimuli to locate hidden food, p.57.
Schaffner, C. M. and E Aureli. The implications of fission-
fusion dynamics for social interactions in wild spider
monkeys, p.57.
Van-Schaik, C. and B. Hellriegel. Between-group antago-
nism: is it affected by the collective action problem?,

Setchell, J. M. Signal content of red coloration in female
mandrills, p.58.
Shanee, N., S. Shanee and A. M. Maldonado. Preliminary
information on the current status of the yellow-tailed
woolly monkey (Oreonax flavicauda) at Yambrasbamba,
Peru, p.58.
Schino, G. Primate reciprocity: general patterns and proxi-
mate mechanisms, p.59.
Schiilke, 0., C. Fischer and J. Ostner. Female social rela-
tionships in two species of Asian colobines, T ''
auratus and Presbytis melalophos, p.59.
Servais, V. and E Joulian. Eri .1. .;, between science and
practice. Different kinds of objectification of animal be-
havior (west-Africa and western-Europe examples), p.60.
Smith, A. M. S., G. P. J. Jonkers. The Barbary macaque:
working towards a solution, p.60.
Skvor, J., V. Vancata, R. Marvan, J. Trilcova, T. Polar, B.
Laznova and C. Tennie. The problem of quality and com-
parability of chimpanzee faecal simples before hormonal
analysis- variability and standarization, p.60.
Smith, A. C. and A. M. Buchanan-Smith. Food, sex and
violence: influences on the intergroup encounters in tam-
arins (Saguinus spp.), p.61.
Smith, D. A., S. Thompson and K. A. I. Nekaris. An asses-
ment of conservation initiatives and priorities, and habi-
tat risks of human land use to the primates (Colobinae,
Hylobatidae and Pongidae) of Kalimantan, p.61.
Smith, T. M. New approaches to understanding tooth de-
velopment and life history in hominoid evolution, p.62.
Spiezio, C., E Fornale and D. Grassi. Problem-solving and
social learning in cotton-top tamarins, p.62.
Spinozzi, G., C. De-Lillo, V. Truppa and G. Castorina. Per-
ceptual grouping ability in humans (Homo sapiens) and
monkeys (Cebus apella). The role of gestalt factors, p.63.
Stojan, M. and E. W. Heymann. Vigilance in mixed-species
and single-species groups of moustached tamarins (Sagui-
nus mystax) in Peruvian Amazonia, preliminary results,
Strasser, A. and J. Burkart. Manual lateralization in Cal-
lithrixjacchus increases as task complexity increases, both
for motor and cognitive complexity, p.64.
Stiickle, S. and D. Zinner. Group coordination in chacma
baboons, p.64.
Tennie, C. and J. Call. Social learning of tool-making in
great apes and human children: the loop study, p.64.
Thorn, J., S. Thompson and K. A. I. Nekaris. Modelling
slow loris (Nycticebus) distributions in Indonesia as a pre-
dictive tool for setting conservation priorities, p.65.
Toxopeus, I. B., J. A. Van-Hoof, J. Bolhuis, M. J. L. Kik
and R. Bleys. The primate brain bank, p.65.
Trilcova, J., R. Marvan, T. Polak, C. Tennie and V. Van-
cata. Preferable and predictable enrichment in captive
common chimpanzees. How affiliative and agonistic be-
havioural patterns may be elicited via pifiatas enrichment
device, p.66.
Truppa, V., G. Spinozzi, T. Stegagno and J. Fagot. Picture-
object recognition in tufted capuchin monkeys (Cebus
apella), p.66.

Neotropical Primates 14(2), August 2007

Tuma, J. and R. Marvan. Grooming body sites prefer-
ences among captive adult common chimpanzee females,
Urbani, B. The ontogeny of the postcranial skeleton in two
neotropical primates (Callimico goeldii and Saguinus fus-
cicollis), p.67.
Vaglio, S., E R. Dani, Y. Yadid, J. M. Cecchi and B. Chi-
arelli. Scent-making in mandrills (Mandrillus sphinx).
Preliminary reports of a chemical and ethological inves-
tigation, p.67.
Vaglio, S., L. Orioli, U. Mauno, S. Coxe, M. Hurley,
G. C. Faussone and B. Chiarelli. Conservation of bono-
bos (Pan paniscus) through Kyoto protocol: the establish-
ment of the "Bonobo Peace Forest" in Maringa-Lopori-
Wamba region (Democratic Republic of Congo), p.68.
Vancatova, M. Tool behaviour of western lowland gorillas
(Gorillagolla go//la,) in captivity, p.68.
Vauclair, J. The contribution of prim ir..1. ._, to the ques-
tion of the gestural origin of language, p.69.
De- Vries, M. and C. Van-Schaik. Factors influencing the
process of innovation in captive great apes (Pongo pygmae-
us abelii, Gorilla godl//a golI/a and Pan troglodytes), p.69.
De-Wandelaer, T. and J. Burkart. Is there a relationship
between gazing behaviour and social structure? An ex-
perimental study with common marmosets (C
jacchus), p.69.
Wantia, J. Bold and cautious personality types in primates:
an individual-based model, p.70.
Weingrill, T. and M. Heistermann. Factors associated with
faecal glucocorticoid levels in zoo orangutans, p.70.
Widdig, A., S. C. Alberts, P. Nuernberg and M. Krawzcak.
Do mothers promote social preference among their pa-
ternally related offspring? Testing mechanisms of paternal
kin discrimination, p.71.
Willems, E. and R. Hill. An assessment of spatial and tem-
poral variation in range use by a group of vervet monkeys
(Cercopithecus aethiops), p.71.
Van-Woerden, J. T and J. Pettersson. Effect of reproductive
state on wild female orangutans, p.71.
Ziltener, A. and C. Van-Schaik. The influence of individual
abilities and social factors on innovation among captive
chimpanzees (Pan troglodytes), p.72.
Zinner, D. and R. Hilgartner. Ecology and use of day shel-
ters in red-tailed sportive lemurs (Lepilemur ruficauda-
tus), p.72.
Zweifel, N. and M. Bastian. Geographic variation in orang-
utan diet: How important is culture?, p.73.


2nd Congress of the European Federation for Primatolo-
gy. September 3 to 7. The second congress of the European
Federation for Primatology (EFP) will be organised by the
Czech Group of Primatologists at the Faculty of Education,
Charles University in Prague. This Congress aims to step
in the future by inviting also those colleagues who are able
to present anthropological topics interesting for primatolo-
gists and thus to encourage an interdisciplinary discussion
among primatologists and anthropologists. The themes will
be: Primate genetics, Primate ethology and socio-biology,
Primate evolution and paleoathropology and Primate ecol-
ogy and conservation among others. For more information
and registration go to: efp_prague2007.pdf>, or visit the web page: cuni.cz/kbio/efp>.

The 25th Annual Symposium for nonhuman primate
models for AIDS. September 10 to 13, California National
Primate Research Center, University of California, Davis.
The main objective is to serve as a scientific forum for the
dissemination and exchange of new research findings, ideas,
and to utilize the knowledge gained from these crucial
nonhuman primate studies to better understand how HIV
and SIV cause disease, and to facilitate the development of
new methods for the treatment, control and prevention of
AIDS in human populations. The symposium will focus on
the biology of primate lentivirus infection and the use of
nonhuman primate models for the study of viral pathogen-
esis, vaccines, and therapeutic approaches against primate
lentivirus infection and disease; primate genomics; viral
agents associated with simian acquired immunodeficiency
syndrome; and the mechanisms of natural resistance in sev-
eral primate species to endemic primate lentiviral infection.
All scientists interested in HIV/AIDS and related research
topics are invited to participate. For more information go
to: .

III Congreso Mexicano de Primatologia. Octubre 24 al
27. La Asociaci6n Mexicana de Primatologia convoca al
Tercer Congreso Mexicano de Primatologia, el cual se lle-
vard a cabo en el auditorio del Instituto de Investigaciones
Antropol6gicas de la Universidad Nacional Aut6noma de
Mexico, Ciudad Universitaria, M6xico DF Las temiticas
que se abordarin seran: Antropologia Biol6gica / Psicolo-
gia Evolutiva; Ecologia Conductual / Etologia; Conserva-
ci6n; Primatologia Medica. Para mayor informaci6n visit:
,Dudas: yahoo.com.mx>.

XI Congress de la Sociedad Mesoamericana para la Biolo-
gia de la Conservaci6n. Noviembre 26 al 30. La Sociedad
Mesoamericana para la Biologia y la Conservaci6n invita a

Neotropical Primates 14(2), August 2007

su XI Congreso: "Uniendo esfuerzos para enfrentar los retos del
cambio climdtico en Mexico y Mesoamirica" el cual se llevardi
a cabo en Cocoyoc, Morelos Mexico. Durante este magno
event y con la participaci6n de la comunidad mesoameri-
cana, se presentarin y discutirin temas relevantes y actuales
sobre biodiversidad, cambio climitico y conservaci6n que
son importantes para cientificos, pueblos indigenas, toma-
dores de decisions y representantes de comunidades. El
objetivo general del congress es aportar e integrar cono-
cimientos y herramientas para el desarrollo sostenible, la
formaci6n de capacidades y la gesti6n de los recursos natu-
rales de la region Mesoamericana, especialmente en el con-
texto del cambio climitico. Para mayores informes visitar:

VIII Curso Latino-americano de Biologia da Conservaiao
e Manejo da Vida Silvestre. Novembro 06 a 12 de Dez-
embro. Na 61tima d6cada o Curso Latino Americano de
Biologia da Conservagao e Manejo da Vida Silvestre vem
contribuindo para a formacao de profissionais comprome-
tidos com a conservagao da diversidade socioambiental da
America Latina. Durante as cinco semanas de capacitagao
os participants compartilham experiencias com renoma-
dos profissionais que atuam nas diferentes esferas da Biolo-
gia da Conservagao. 0 conteddo programitico deste curso
trata primeiramente de uma abordagem conceitual te6rica,
seguido de metodologias e ferramentas dteis em estudos de
campo e por 61timo, estudos de caso nos quais os alunos
vivenciarao a realidade de um program de conservagao na
regiao do Pontal do Paranapanema. As aulas serao minis-
tradas na sede do IPE, em Nazar6 Paulista SP e no mu-
nicipio de Teodoro Sampaio SP. Para mais informadao
visit o web page: asp?mes=Novembro&id= 104>.

Animal Training and Behavior Through Positive Rein-
forcement Further Challenging and Advanced Issues.
December 7 to 9, Munich, Germany. The seminar will be
imparted by Ken Ramirez, vice president for animal collec-
tions and animal training; he develops and supervises ani-
mal-care programs, staff training and development as well
as public presentation programs for the animal collection at
Shedd Aquarium. The Seminar will include themes related
to operant conditioning, training situations and problem
solving with positive reinforcement. For more information
visit: .

6.- Gbttinger Freilandtage. Primate Behavior and Human
Universals. December 11 to 14, Gbttingen, Germany. This
conference aims to bring together primatologists, evolu-
tionary anthropologists and psychologists to summarise
our current state of knowledge concerning behavioral
variation and its determinants within the order Primates,
including humans. Specifically, it will focus on three as-
pects: (1) comparative studies of behavioral adaptations
across (human and non-human) primates that examine
evolutionary principles, (2) the ability and failures of evo-
lutionary theory to explain human behavioral traits that

affect survival and reproduction, and (3) to identify and
explain human behavioral universals. For additional de-
tails contact Prof. Dr. Peter Kappeler or
visit the web page: de/welcome.html>.


III Confer&ncia Nacional do Meio Ambiente. Em maio de
2008 seri realizada a "III Conferencia Nacional do Meio
Ambiente", cujo objetivo seri construir um espago de con-
vergencia social para a formulagao de uma agenda national
do meio ambiente, por intermedio da mobilizagao, edu-
cacao e ampliacao da participagao popular, com vistas ao
estabelecimento de uma political de desenvolvimento sus-
tentivel para o Pais. Afinal, a definigao de political pdblicas
para um Brasil sustentavel depend de mudangas na forma
de atuacao das esferas governamentais, do setor produtivo,
das organizagoes da sociedade, chegando ao cotidiano de
cada cidadao. Em suas edikoes, a conferencia coloca para a
sociedade temas estratngicos para o Pais, que visam a con-
servagao da biodiversidade, da agua, do clima e dos recursos
energ6ticos, com vistas ao desenvolvimento sustentivel, le-
vando em consideracao que 6 possivel sim crescer sem de-
gradar a natureza. Quem quiser enviar textos para compor a
Comissao Nacional, pode envii-los at6 dia 15 de dezembro
para . Informacges sobre a
Conferencia Nacional em nacional> ou pelo e-mail .

9th Student Conference on Conservation Science. March
25 to 27, 2008. Department of Zoology, University of
Cambridge, United Kingdom. The SCCS is aimed at
young researchers working in universities departments,
conservation organizations, or resource management agen-
cies. Approximately 170 postgraduate students attend the
conference each year, from a broad range of disciplines
in conservation, including ecology, geography, sociology
and economics. The most important part of the three-day
programme will be poster sessions and 33 fifteen minute
talks by students on any aspect of conservation science.
There will also be workshops, presentations by conserva-
tion NGOs and agencies and social events designed to give
participants the opportunity to make new contacts in their
own and related disciplines. Prizes are awarded to posters
and talks of outstanding quality and relevance to conserva-
tion. For more information and applications go to: sccs-cam.org>.

22nd Annual Meeting of the Society of Conservation Bi-
ology. The 22nd annual meeting of the SCB will be held at
the Chattanooga Convention Center, Chattanooga, Ten-
nessee, USA, from July 13 to 17, 2008. The chair of the
meeting will be Dr. David A. Aborn, from the Department
of Biological and Environmental Sciences, University of
Tennessee at Chattanooga. As evidenced by several previ-
ous meeting themes, there are connections among many

Neotropical Primates 14(2), August 2007

aspects of the environment and its conservation, and recog-
nizing those connections is critical for achieving the goals of
conservation biology. To that end, the theme for the 2008
SCB annual meeting "T'ow the mountains to the sea" will
examine several major ecosystems, both as separate compo-
nents and as a connected entity. For more information and
submission dates, go to: .

XXIInd IPS Congress. August 3 to 8, 2008, Edinburgh
International Conference Center, Edinburgh, Scotland.
Sponsored by the Primate Society of Great Britain. Ab-
stracts for oral and poster presentations must be submitted
by 30th November 2007. For information consult the web
site: .

XXth International Congress of Zoology. August 26
to 29, Jussieu Grand Campus, Paris, France. Held
every four years, the International Congress of Zool-
ogy gives zoologists and professionals from all fields re-
lated to zoology the chance to come together to discuss
the current status of zoology and to share ideas about
future development of all its disciplines. The Internation-
al Society of Zoological Sciences (ISZS) and the Societe
Zoologique de France invite you to the XX International
Congress of Zoology. It will be co-hosted by the Universi-
ties Pierre et Marie Curie (PVI), Paris-Sud (PXI) and the
Museum National d'Histoire Naturelle. Registration will
open on 1st December 2007. For information about the
Congress, please visit: or

Monkeys of the Atlantic Forest of Eastern Brazil Pocket Identification Guide
Mail and Fax Order Form

Monkeys of the Atlantic Forest of Eastern Brazil Pocket Identification Guide by Russell A. Mittermeier,
Adelmar F. Coimbra-Filho, Maria Cecilia Martins Kierulff, Anthony B. Rylands, Sergio Lucena Mendes,
Alcides Pissinatti and Livia Meneghel de Almeida.
ISBN: 978-1-934151-12-9. First Edition.

Price: $7.95 (includes UPS Ground shipping within the continental United States)
For orders requiring faster service than UPS Ground, you will be responsible to pay all shipping costs.

Please call the phone number listed below for: overnight deliveries, wholesale orders, and international

Please complete the following form, print it out and mail or fax to:

Jill Lucena
Conservation International
2011 Crystal Drive, Suite 500
Arlington, VA 22202 USA
Tel: +1 (703) 341-2536
Fax: +1 (703) 553-4817

First Name Last Name

Company Name

Mailing Address City State Zip/Postal Code

Telephone Fax

E-Mail Address

Order Form should include credit card information or be sent along with check or money order, in
U.S. dollars, made payable to Conservation International. Please allow 2-3 weeks for delivery.

Quantity x $7.95 each Total:
Payment Enclosed (check or money order in US $ only)

Charge my credit card:
Name (as it appears on card)
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of the Atlantic Forest
of Eastern Brazil
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R- A. Mlrudn& Ah*.f Rfl $A
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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 publications, primatologi-
cal society information and suchlike.

Please send all English and Spanish contributions to: Erwin Palacios,
Conservaci6n Internacional-Colombia, Carrera 13 # 71-41 Bogoti
D.C., Colombia, Tel: (571) 345-2852/54, Fax: (571) 345-2852/54,
e-mail: , and all Portuguese contribu-
tions to: Julio Cisar Bicca-Marques, Departamento de Biodiversidade
e Ecologia, Pontificia Universidade Cat6lica do Rio Grande do Sul,
Av. Ipiranga, 6681 Pr6dio 12A, Porto Alegre, RS 90619-900, Brasil,
Tel: (55) (51) 3320-3545 ext. 4742, Fax: (55) (51) 3320-3612,
e-mail: .

Manuscripts may be in English, Spanish or Portuguese, and should
be double-spaced and accompanied by the text on CD for PC com-
patible text-editors (MS-Word, WordPerfect, Excel, and Access),
and/or e-mailed to (English, Spanish)
or (Portuguese). 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 their English manuscripts 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, Systemat-
ics, Genetics (when relevant for systematics and conservation),
Biogeography, Ecology and Conservation. Text for full articles
should be typewritten, double-spaced with no less than 12 cpi font
(preferably Times New Roman) and 3-cm margins throughout,
and should not exceed 25 pages in length (including references).
Please include an abstract in the same language as the rest of the
text (English, Spanish or Portuguese) and (optional) one in Portu-
guese or Spanish (if the text is written in English) or English (if the
text is written in Spanish or Portuguese). Tables and illustrations
should be limited to six, except in cases where they are fundamen-
tal for the text (as in species descriptions, for example). Full articles
will be sent out for peer-review. For articles that include protein
or nucleic acid sequences, authors must deposit data in a publicly
available database such as GenBank/EMBL/DNA Data Bank
of Japan, Brookhaven, or Swiss-Prot, and provide an accession
number for inclusion in the published paper.
Short articles. These manuscripts are usually reviewed only
by the editors. A broader range of topics is encouraged, includ-
ing such as behavioral research, in the interests of informing on
general research activities that contribute to our understanding
of platyrrhines. We encourage reports on projects and conserva-
tion and research programs (who, what, where, when, why, etc.)
and most particularly information on geographical distributions,
locality records, and protected areas and the primates that occur in
them. Text should be typewritten, double-spaced with no less than
12 cpi (preferably Times New Roman) font and 3-cm margins
throughout, and should not exceed 12 pages in length (including

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 that are publishable.
Tables. Tables should be double-spaced, using font size 10, and
prepared with MS Word. Each table should have a brief title.
News items. Please send us information on projects, field sites,
courses, Thesis or Dissertations recently defended, recent publica-
tions, awards, events, activities of Primate Societies, etc.
References. Examples of house style may be found throughout
this journal. In-text citations should be first ordered chronologi-
cally and then in alphabetical order. For example, "... (Fritz, 1970;
Albert, 1980, 2004; Oates, 1981; Roberts, 2000; Smith, 2000;
Albert etal., 2001)..."
In the list of references, the title of the article, name of the journal,
and editorial should be written in the same language as they were
published. All conjunctions and prepositions (i.e., "and", "In") should
be written in the same language as rest of the manuscript (i.e., "y"
or "e", "En" or "Em"). This also applies for other text in references
(such as "PhD thesis", "accessed" -see below). Please refer to these
examples when listing references:
Journal article
Stallings, J. D. and Mittermeier, R. A. 1983. The black-tailed mar-
moset (Callithrix argentata melanura) recorded from Paraguay. Am.
J. Primatol. 4: 159-163.
Chapter in book
Brockelman, W Y. and Ali, 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
(Natural 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.
UNESCO. 2005. UNESCO Man and the Biosphere Programme.
United Nations Educational, Scientific, and Cultural Organisation
(UNESCO), Paris. Website: http://www.unesco.org/mab/index.
htm. Accessed 25 April 2005. ("Acessada em 25 de abril de 2005"
and "Consultado el 25 de abril de 2005" for articles in Portuguese
and Spanish respectively).
For references in Portuguese and Spanish:
"and" changes to "e" and "y" for articles in Portuguese and Spanish
"In" changes to "Em" and "En" for articles in Portuguese and Span-
ish respectively.
"Doctoral thesis" changes to "Tese de Doutoramento" and "Tesis de
Doctorado" for articles in Portuguese and Spanish respectively.
"MSc Thesis" changes to "Dissertaiao de Mestrado" and "Tesis de
Maestria" for articles in Portuguese and Spanish respectively.
"Unpublished report" changes to "Relat6rio T&cnico" and "Reporte
no publicado" for articles in Portuguese and Spanish respectively.

Neotropical Primates
A Journal and Newsletter of the IUCN/SSC Primate Specialist Group
Vol. 14(2), August 2007



Further Notes on the Distribution of Endemic Bolivian Titi Monkeys, Callicebus modestus and Callicebus olallae
Jesus M artinez and Robert B W allace .............................................. ........................................................................ ........................ 47
Resource Use and Seed Dispersal by Red Howler Monkeys (Alouatta seniculus) in a Colombian Andean Forest
Paula Giraldo, Carolina Gdmez-Posada, Jesus Martinez and Gustavo Kattan................................................................................55
Cebusparaguayanus in Zoos: The Spontaneous Expression of Species-Specific Behaviors
A ldo M G iudice and Rom ina Pav .................................................................................................................................................. 65

Short Articles

Registro de Alimentaoio Noturna em Macaco-Prego (Cebus apella)
C laudia R Silva .............................................................................................................................................................................. 72
A Case of Spontaneous Tool-Making by a Captive Capuchin Monkey
Tiago Soares Bortolini andjulio Cisar Bicca-Marques .................................................................................. .............................. 74
Power lines and howler monkey conservation in Porto Alegre, Rio Grande do Sul, Brazil
Luisa Xavier Lokschin, Rodrigo Cambard Printes, Juliane Nunes Hallal Cabral and Gerson Buss............................................. 76
Preliminary Survey on the Current Distribution of Primates in Belize
Sidn S. W waters and O scar U lloa ............................................ ............... ......... ..... ................ ............................ .................................. 80
Sympatric Occurrence of Alouatta caraya and Alouatta sara at the Rio Yacuma in the Beni Department,
Northern Bolivia
Anna B. S. Biintge and Lennart W Pyritz................................ ......................................................................................... 82
Registro de Ocorrencia de Cebus kaapori (Cebidae: Primates) na APA Lago de Tucuruf
Fernanda Almeida Cunha, Maria Aparecida Lopes, Sidnei de Melo Dantas, Nivia Aparecida,
Silva do Carmo e Suleima do Socorro Bastos da Silva................................. .......................................................................84
Consumption of Carapa guianensis Bark by Cebus capucinus in La Reserva Biol6gica Indo-Mafz, Nicaragua
Thor H anson ................................................................................................................................................................................... 85

N ew s ............................................................................................................................................................................................ 87

P rim ate S societies ........................................................................................................................................................................ 90

R ecen t P u blication s ...................................................................................................................................................................90

M eetin g s ..................................................................................................................................................................................... 99

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