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Title: Neotropical primates
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Permanent Link: http://ufdc.ufl.edu/UF00098814/00052
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Title: Neotropical primates a newsletter of the Neotropical Section of the IUCNSSC Primate Specialist Group
Abbreviated Title: Neotrop. primates
Physical Description: v. : ill. ; 27 cm.
Language: English
Creator: IUCN/SSC Primate Specialist Group -- Neotropical Section
IUCN/SSC Primate Specialist Group -- Neotropical Section
Conservation International
Center for Applied Biodiversity Science
Publisher: Conservation International
Place of Publication: Belo Horizonte Minas Gerais Brazil
Belo Horizonte Minas Gerais Brazil
Publication Date: January 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: VID00052
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
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    Back Matter
        Back Matter 1
        Back Matter 2
        Back Matter 3
    Back Cover
        Back Cover
Full Text

ISSN 1413-4703



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

Volume 14
Number 1
January 2007

Erwin Palacios
Lia rii Cort6s-Ortiz
J6lio C6sar Bicca-Marques
Eckhard Heymann
Jessica Lynch Alfaro
News and Books 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, Conservation International -Colombia
Liliana Cort6s-Ortiz, Museum of Zoology, University of Michigan, Ann Arbor, MI, USA
J6iio Cisar Bicca-Marques, Pontificia Universidad Cat6lica do Rio Grande do Sul, Porto Alegre, Brasil
Eckhard Heymann, Deutsches Primatenzentrum, Gittingen, Germany
Jessica Lynch Alfaro, Washington State University, Pullman, WA, USA
News and Books Reviews
Brenda Sol6rzano, Instituto de Neuroetologfa, 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 Rodrfguez-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 F Ferrari, Universidade Federal do Sergipe, Aracaj6, Brazil
Russell A. Mittermeier, Conservation International, Arlington, VA, USA
Marta D. Mudry, Universidad de Buenos Aires, Argentina
Horicio Schneider, Universidade Federal do Pari, Campus Universitirio de Braganga, 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, Section on Great Apes, Liz Williamson
Regional Coordinators Neotropics
Mesoamerica, Ernesto Rodrfguez-Luna
Andean Countries, Erwin Palacios and Eckhard W. Heymann
Brazil and the Guianas, M. Cecilia M. Kierulff
Regional Coordinators Africa
West Africa, John F. Oates
East Africa, David Mbora
Regional Coordinator Madagascar
Jarg 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(1), January 2007 1

During the last 12 years Neotropical Primates has achieved the remarkable task of providing a network and forum of commu-
nication for all primatologists and university students involved in New World primate conservation, ecology and behavior,
in both captive and field-based research settings. This has been possible mainly to the tireless work of its founding editors,
Dr. Anthony Rylands and Dr. Ernesto Rodriguez-Luna, a dedicated team of assistant editors and designers, and the gener-
ous support of the Margot Marsh Biodiversity Foundation, Los Angeles Zoo, Earthkind, the Houston Zoological Society
Conservation Program, the Columbus Zoo, Wildlife Preservation Trust International, the Jersey Wildlife Preservation Trust,
Penscynor Wildlife Park, the Detroit Zoological Institute, the Brazilian National Biodiversity Working Group (GTB), the
Primate Society of Great Britain (PSGB), and Conservation International. As a result 51 issues have been published to date,
including about 370 articles and 60 thesis abstracts. Starting with volume 14, Neotropical Primates now has a new editorial
team, with the challenge of continuing to make Neotropical Primates a frequently cited and well-respected journal, provid-
ing information on activities related to the study and conservation of primates, and the conservation of their forest habitats
throughout Central and South America, and encouraging non-governmental and governmental institutions to keep these
efforts going. We are committed to this endeavour and as such want to renew an invitation to contributors to send us their
manuscripts, and to thank in advance all the institutions which make it possible for NeotropicalPrimates to continue being
a widespread and easily accessible journal of primatology.

Erwin Palacios
Liliana Cortds-Ortiz
Jflio CUsar Bicca-Marques
Eckhard Heymann
Jessica Lynch Alfaro

2 Neotropical Primates 14(1), January 2007


Carolina G6mez-Posada1, Jesus Martinez1'2'3, Paula Giraldo1'2'4 and Gustavo H. Kattan1

1 Fundacidn .....A- : \i : .-. Conservation Society Colombia Program, Cali, Colombia. AA. 25527,
e-mail: ,
2 Departamento de Biologia, Universidad del Valle, Cali, Colombia
3 Present address: Universidad de Los Lagos, Osorno, Chile, e-mail:
4Present address: Vrije University, Amsterdam, The Netherlands, e-mail: < : '."


We studied habitat use and ranging patterns of five red howler monkey troops for six months in a lower montane forest in
the Colombian Andes. The study area is a mosaic of mature and secondary forest and non-commercial ash, pine and oak
plantations that were established as part of a reforestation program. Home ranges (10 3 ha) and daily distances traveled
(554 248 m) were similar to those reported for lowland habitats. Home range use differed between groups using mature
forest versus secondary forest and plantations, reflecting structural differences between habitat types. We estimated a density
of 0.72 individuals and 0.1 groups per hectare. This high density is related to the protected status of the study area and the
howler monkeys' behavioral plasticity, which has allowed them to colonize new environments such as plantations.

Key Words: Alouatta seniculus, cloud forest, Colombia, density, home range, habitat use, mature and secondary forest, tree


Estudiamos los patrons de uso del espacio de cinco grupos de mono aullador rojo durante seis meses en un bosque montano
bajo en los Andes de Colombia. El area de studio es un mosaico de bosques maduros y de regeneraci6n, y plantaciones no
comerciales de urapin, pino y roble que fueron establecidas en un program de reforestaci6n. Las areas de actividad (10
3 ha) y las distancias recorridas diariamente (554 248 m) fueron similares a las reportadas para poblaciones de tierras bajas.
Los patrons de uso del Area de actividad difirieron entire los grupos que usaban bosque maduro y los que usaban combina-
ci6n de bosque secundario y plantaciones, lo cual refleja las diferencias estructurales entire los tipos de habitat. Estimamos
una densidad de 0.72 individuos y 0.1 grupos por ha. Esta alta densidad esti relacionada con el estado de protecci6n del area
y con la plasticidad conductual de estos monos, que les ha permitido colonizar nuevos ambientes como las plantaciones.

Palabras Clave: Alouatta seniculus, bosque nublado, Colombia, densidad, irea de dominio vital, uso de habitat, bosque
maduro y secundario, plantaciones de arboles


Home ranges and population densities of animals vary in
relation to factors such as body size, diet, habitat type, social
system, and human disturbance (Milton and May, 1976;
Crockett and Eisenberg, 1987; Peres, 1997; Chapman and
Balcomb, 1998). In general, for energetic reasons, animals
of large body size require larger home ranges and have lower
population densities than smaller animals within the same
trophic level (Fleagle, 1999; MacNab, 2002). Population
density is also modulated by habitat productivity and posi-
tion in the trophic pyramid for a particular species (Eisen-

berg, 1980; Fleagle, 1999; MacNab, 2002). In addition,
for animals like primates that live in groups, population
density depends on home range, troop size and the degree
to which ranges of different troops overlap (Eisenberg,
1980; Crockett and Eisenberg, 1987).

The red howler monkey (Alouatta seniculus) is one of the
largest Neotropical primates, with females reaching up to
6.3 kg and males 7.5 kg (Defler, 2003). Home ranges of red
howlers vary between 7 and 25 ha (Crockett and Eisenberg,
1987), but may reach up to 182 ha (Palacios and Rodri-
guez, 2001). Small home ranges have been associated with

Neotropical Primates 14(1), January 2007

the highly folivorous diet and sedentary habits of howler
monkeys, but may also vary on the basis of interspecific in-
teractions (Milton, 1980; Gaulin and Gaulin, 1982; Braza
et al., 1983; Crockett and Eisenberg, 1987). Depending on
habitat and group composition, among other factors, pop-
ulation densities of red howlers vary widely, with a mean of
34-55 ind/km2, but densities as high as 150 ind/km2 have
been reported in some habitats (Crockett and Eisenberg,
1987; Chapman and Balcomb, 1998).

Throughout its broad distribution in northwestern South
America, the red howler is found from lowland rain forest,
gallery forest, and dry woodlands to montane forest. In
the Colombian Andes it ranges up to 2400 m and occa-
sionally up to 3200 m (Hernindez-Camacho and Cooper,
1976; Defler, 2003). Montane populations of red howlers
in the Colombian Andes are threatened by habitat destruc-
tion and fragmentation. Few tracts of continuous forest
still exist in the Central and Western Cordilleras (Kattan
and Alvarez-L6pez, 1996), and many howler populations
are isolated in small fragments, sometimes as small as 10
ha (G6mez-Posada et al., 2005). Protecting and managing
these populations requires an understanding of patterns of
spatial and habitat use and population densities.

Humid montane forest differs from lowland rain forest in
having lower productivity (correlated with a decrease in
temperature) and lower plant diversity, especially when
above 1500 m (Gentry, 1992; Cavelier, 2001). Thus, pop-
ulation densities of howlers may be expected to be lower
in montane forest than in the lowlands. Relatively low
densities have been reported for two sites in the Colom-
bian Andes (Gaulin and Gaulin, 1982 = ca. 15 ind/km2;
Morales-Jimenez, 2002 = 31.3 ind/km2), but patterns of
habitat and space use and population densities have not
been rigorously documented. In this study, we present data
obtained over six months on population density and space
use of five red howler troops in a cloud forest in the Cen-
tral Cordillera of the Colombian Andes. Our study site is a
mosaic of habitat types, including old-growth and second-
ary forest, and monodominant patches of both exotic and
native trees that were established in a reforestation program.
Our study area is within an extensive, continuous forest
(several thousand hectares) and our data provide baseline
information for a more extensive study documenting re-
sponses of red howlers to fragmentation (G6mez-Posada,
unpublished data).

Study Area

The study was conducted at Otin Quimbaya Flora and
Fauna Sanctuary (Otin Quimbaya), a 489 ha protected
area located on the western slope of the Central Cordillera
of the Colombian Andes, east of the city of Pereira. The
study area ranges between 1800 and 2100 m. Otin Qui-
mbaya is contiguous with Ucumari Regional Park, encom-
passing 3980 ha. Both areas protect the Otin River drain-
age between elevations of 1750 and 2600 m. The study area

lies in the very humid lower montane forest life zone of the
Holdridge classification system (Londofio, 1994). Mean
annual rainfall is 2712 mm (El Cedral weather station, Ce-
nicaf6, 1995-2001), with a bimodal pattern. Rainy periods
occur in April-June and September-November. There is a
mild dry season in December-February, and a stronger one
in July-August (Aguilar and Rangel, 1994). In 2001 total
precipitation was 2117.8 mm. Mean annual temperature
is 15C.

Native forest in the Otin River drainage was largely cleared
during the first half of the 20th century, for the establish-
ment of cattle pastures. Some old-growth forest fragments
remained, mainly in deep canyons, although hardwoods
were extracted from most of the region. In the 1960s a re-
forestation program was initiated by local authorities with
the objective of stabilizing soils and stopping erosion in the
watershed. Some patches were planted with exotic Chinese
ash (Fraxinus chinensis) and cypress (Cupressus lusitanica),
and with native Andean oak (Quercus humboldtii). Most of
the area currently included in Otin Quimbaya, however,
was abandoned to natural regeneration, with seeds provid-
ed by native forest remnants.

Currently, the Otin drainage is 80% forested, with a
mosaic of old-growth forest, secondary forest of different
ages, and interspersed patches of monospecific tree planta-
tions. As these plantations were established for reforesta-
tion purposes, they were not managed and are presently
invaded by native vegetation, particularly in the understory
and edges (Durin and Kattan, 2005). The canopy remains
monodominant, though. Presently the Otin Quimbaya
area is covered by a mosaic of old-growth and secondary
forest on the mountainsides, and strips of ash plantations
on the valley floor, along the river. Oak and cypress planta-
tions occur in small patches near the river, neighboring ash
stands. A narrow dirt road cuts across the park and the ash


Between July and December 2001 we identified all red
howler monkey troops in 113 ha of Otin Quimbaya,
which included native forest of different ages, and ash,
cypress and oak patches. We selected five groups (labeled
C-G) for intensive observation. Each of these troops was
followed for a maximum of three days per month. Daily
travel routes involved going from a sleeping tree in the
morning at 0630-0800, through a series of feeding trees
throughout the day, to a different sleeping tree in the after-
noon at 1600-1700. We identified individuals by age and
sex following Defler (1981) and Soini (1992).

During observation periods, we recorded data on activ-
ity patterns and diet (Martinez, 2003; Giraldo et al., sub-
mitted). To evaluate habitat use, we followed each troop,
taking note of its location and habitat type every half hour.
We superimposed a 14 ha grid over a map of the study area,

Neotropical Primates 14(1), January 2007

and plotted all troop locations to obtain frequencies of use
of each 14 ha quadrant within their home ranges (NRC,
1981). To quantify patterns of habitat use, we added all
records in quadrants in each habitat type for each group,
and used a X2 test to compare habitat use among groups.
To establish whether quadrant use frequency distributions
differed from random (Poisson), we used a X2 test. This dis-
tribution is zero-truncated because in theory some cells will
remain unused (Robinson, 1986; Di Bitetti, 2001). We
used a Spearman rank correlation coefficient to correlate
the number of feeding and sleeping trees in each quadrant
and the quadrant's frequency of use. We pooled troops in
two categories according to the main habitat type they used
(two troops in old-growth forest versus three troops in sec-
ondary forest/plantations), and used a Mann Whitney U
test to compare the number of quadrants used per day, and
the mean number of records per quadrant between the two
habitat types. To test whether groups used habitat types in
proportion to their availability, we compared the frequency
of use of each habitat with its area within the home range
with a x2 test.

We estimated the home range size of each troop as the
number of 14 ha quadrants used at least once (NRC, 1981).
Home range overlap was calculated using the formula 0 =
HR*D/GS, where 0 is overlap, HR is mean home range,
D is population density and GS is mean group size. This
index reveals the number of troops that can overlap at any
point within the study area (Terborgh, 1983; DiBitetti,
2001). We correlated home range size and group size with a
Spearman rank correlation coefficient. Daily distance trav-
eled was defined as the distance covered by a group from
one sleeping tree to the next (NRC, 1981). We joined all
half-hour location points of each group with straight lines
to obtain the distance traveled in a day. Daily distances
traveled by different groups were compared with a Krus-

kal-Wallis test. The number of trees visited by groups in
old-growth forest versus secondary forest/plantation was
compared with a Mann-Whitney U test.


Eleven red howler monkey troops with a total of 82 in-
dividuals inhabited the 113 ha study area (Table 1, Fig.
1), for a density of 72.6 ind/km2 and 9.73 groups/km2.
We observed other groups outside the core study area, and
solitary adult males throughout the study area. Red howler
troops were stable throughout the study period (Table 1).
Mean group size was 7.3 individuals (DS=2.5, range 3-10)
and increased to 7.5 after an infant was born and a sub-
adult male joined group G. Groups were composed of one
adult male, one to three adult females, one or two sub-
adults and one to four juveniles and infants. Sex ratio was
biased toward females (1:0.6) and the ratio of adult females
to immature individuals (juveniles and infants) was 1:1.2.

Habitat use. The five more intensely studied troops used dif-
ferent habitat types in different proportions (2 = 1351.6,
df= 8, p < 0.01; Table 2). Cypress plantations were used as
corridors between forest patches and as sleeping trees, but
were used infrequently during the day. Sometimes when
howlers were foraging in secondary forest, where few large
trees were available, they moved to cypress patches for di-
urnal resting periods. Oak stands also were only used as
routes between ash stands and forest patches. Ash stands
had some dispersed Cecropia and Ficus trees that were used
as sleeping and feeding trees. Howlers also fed on immature
ash fruits and used big ash trees as sleeping trees (Giraldo
et al., submitted).

Home range use was not random, as howlers used some
quadrants more than expected (Fig. 2). The more inten-

Table 1. Size and composition of 11 red howler monkey troops at Otim Quimbaya Flora and Fauna Sanctuary, Central Andes of
Adult Subadult Juvenile
Group M F M F Infant Unknown Total
M* F* M F M F
A 1 1 2 1 1 6
B 1 3 1 2 3 10
C 1 2 1 1 5
D 1 3 1 1 1 1 2 10
E 1 2 1 1 1 2 8
F 2 2 1 1 2 2 10
G 1 2 1 1 1 2 8
H 10 10
I 7 7
J 1 2 1 1 5
K 1 1 1 3
Total 10 18 7 5 6 4 11 21 82
Mean 1.1 2.0 1.2 1.0 1.2 1.0 1.8 7.5
* M: males; F: females

Neotropical Primates 14(1), January 2007

sively used quadrants, corresponding to dormitories, la-
trines, and feeding trees (r = 0.63, p < 0.01), did not form
a core area but were dispersed throughout the home range.
The number of sleeping and feeding trees per quadrant was
lower for the three groups that mainly used plantations
and secondary forest than for the two troops using mature
forest (Z = 2.2, n = 143 and 63, p = 0.02) (Table 3). Home
range use was different between groups using mature forest
and groups using secondary forest and plantations. The

latter usually used one or two quadrants intensively for sev-
eral days, feeding and sleeping in the same area. After crops
were exhausted in these feeding trees, monkeys moved
throughout their home range in a haphazard way until
finding a new feeding tree. In contrast, troops in mature
forest used several quadrants each day, moving between
feeding trees, sleeping trees, and latrines, traversing their
home ranges in three or four days. The mean number of
quadrants visited per day was similar for mature forest and

Table 2. Home range composition by habitat type, and proportion of observations in each habitat type, for five red howler monkey troops
in Central Andes of Colombia.
Habitat Type
Group Mature Forest Secondary Forest Plantation N
% home range % obs. % home range % obs. % home range % obs.
C 90.9 97.2 9.1 2.8* 246
D 58.9 63.3 41.1 36.7 297
E 31.4 17.3* 68.6 82.7* 272
F 17.3* 24.0 57.7 52.7 25.0 23.3 146
G 100 100 267
Significant difference between observed and expected frequencies (p < 0.05, Z2 test)



S500 0 600 1000 Meters

Home Range
M Troop C
rT Troop D
STroop E
Troop F
S Troop G
...... Other troops.
Habitat type
Secundary forest.
I Mature forest

Figure 1. Map of Otim Quimbaya Flora and Fauna Sanctuary, Central Cordillera, Colombian Andes, showing habitat types and home
ranges of red howler groups.

Neotropical Primates 14(1), January 2007

for secondary forest/plantation groups (U = 88, n = 17
and 12, p = 0.5; Table 3), but coefficients of variation were
larger for the latter. The percent use of each quadrant was
larger for mature forest troops (Z = 1.9, n = 143 and 63,
p = 0.05) (Table 3).

Home range and daily distance traveled
Home ranges of the five troops varied between 7.5 and
14 ha, with a mean + SD of 10.2 3.03 ha (Table 4, Fig.
2). Bigger groups showed a tendency to have larger home
ranges (r, = 0.87, n = 5, p = 0.05). Groups C and G had
small, compact home ranges 7.8 0.4, with an area/pe-
rimeter ratio of 48:1, while the other groups had larger
(11.9 2.8) and elongated home ranges, with an area/pe-
rimeter ratio of 35:1 (Table 4, Fig. 2). Home ranges of the
five troops were contiguous and had an overlap index of

Figure 2. Intensity of use of quadrants in home ranges of five
groups of howler monkey at Otin Quimbaya. Frequency distri-
bution of quadrant use was significantly different from Poisson in
all cases (group C: X2 = 1389.7, df= 31, p < 0.01; group D: 1' =
590.9, df= 55, p < 0.01; group E: X = 1977.1, df= 34, p < 0.01;
group F: 2 = 98, df= 51, p < 0.01; group G: X = 3963, df= 29,
p < 0.01).

0.98 (i. e., any point within the study area was used by
0.98 groups). Unused areas between home ranges were
early second growth forest, which lacks the structure and
resources required by howlers (Fig. 1). On average, each
group shared 20.9% 12.9% of its home range with other
groups. When groups met at feeding trees, they engaged in
vocal displays and usually the smaller group retreated.

Daily distance traveled varied between 317.5 m (group D)
and 1321.2 (group E), with a mean of 553.9 247.9 (Table
4). Mean daily distance was similar among groups (H =
4.38, df= 4, p = 0.3). However, the coefficient of variation
for groups using mature forest was much smaller (28%)
than for groups using secondary forest/plantation (53%).
This reflected different patterns of habitat use. When trees
in plantations produced fruit, howlers exploited them in-
tensively, remaining near this tree for one or more days.
After the crop was exhausted, they traveled throughout the
home range in search of fruiting trees. In mature forest,
in contrast, howlers fed from several trees each day and
traveled through their home range in three or four days.
The number of feeding trees visited per day was lower for
secondary forest/plantation groups (5.7 1.6) than for
mature forest groups (9.1 4.4) (U = 54, n = 17 and 12,
p = 0.05).


Group sizes of red howler monkeys observed at Ot6n Qui-
mbaya are within the range of 2-16 individuals (mean =
6-9) usually reported for this species (Neville, 1972; Izawa,
1988, 1997; Soini, 1992; Chapman and Balcomb, 1998;
Defler, 2003). Group composition is also typical, with a
slightly higher proportion of adult females to adult males
(Defler, 1981), reflecting the red howler's social organiza-
tion: one dominant male, one to four adult females and
their offspring, and zero to three subadults (Izawa, 1988,
1997; Soini, 1992; Crockett, 1996). The ratio of adult
females to immature (juveniles and infants) may give an
idea of population health (Heltne etal., 1976). A high ratio
may indicate a declining population, and a low ratio may
indicate an expanding population. At Ot6n Quimbaya we
found 1.2 immature individuals for each adult female, sug-
gesting a growing population (Defler, 1981).

Red howler monkeys usually have densities of 34-55 ind/
km2 (Defler, 1981; Freese et al., 1982; Braza et al., 1983;
Terborgh, 1983; Soini, 1992; Chapman and Balcomb,
1998), but may vary from 4 to 150 (Neville, 1972; Klein
and Klein, 1976; Rudran, 1979; Freese etal., 1982; Crock-

Table 3. Number of feeding and sleeping trees per quadrant, number of quadrants visited per day and percent use of each quadrant (mean
+ SD, CV), for five red howler troops using two habitat types in the Central Andes of Colombia.
No. Trees No. Quadrants % Use
Mature forest (groups C and G) 2.0 + 2.5 9.6 + 3.5, 36.3 3.2 + 5.0, 157.8
0.9+ 1.3 11.7+6.9,60 2.1 +3.7, 178.5
Secondary forest and plantation (groups D, E and F) 0. 1.3 11.7 6.9,60 2.1 + 3.7, 178.5
p 0.02 p > 0.05 p 0.05

Neotropical Primates 14(1), January 2007

Table 4. Home range and daily distance traveled (x SD, n) for five red howler monkey troops at Otim Quimbaya Flora and Fauna
Sanctuary, Central Andes of Colombia.
Group Home range (ha) Distance (m) No. hours of observation No. days of observation
C 8.0 526.7 + 150.4, 6 119.4 18
D 14.0 412.2 + 120.2, 6 136.4 19
E 8.7 661.3 + 341.1, 9 132.0 15
F 13.0 660.0 +481.9, 2 52.1 9
G 7.5 528.3 + 154.6, 6 117.4 15
Mean 10.2 553.9 + 247.9, 29

ett and Eisenberg, 1987; Rylands and Keuroghlian, 1988;
Palacios and Rodriguez, 2001). Population densities of red
howler monkeys vary depending on factors such as habitat
characteristics (e. g., plant diversity and abundance, forest
productivity and structure; Freese et al., 1982; Crockett,
1985) and habitat heterogeneity and seasonality (Peres,
1997). Competition with other frugivorous species or with
other primates may keep densities low (Klein and Klein,
1976; Defler, 1981; Palacios and Rodriguez, 2001). Much
variability in red howler populations is related to their
recent history, such as human disturbance (habitat altera-
tion and fragmentation, hunting), fruit crop failure, and
disease (Freese et al., 1982; Crockett, 1985; Rylands and
Keuroghlian, 1988; Peres, 1990; Sussman and Phillips-
Conroy, 1995; Chapman and Balcomb, 1998).

The density of 72.6 ind/km2 at Otin Quimbaya corre-
sponded to the upper part of the range. The red howler is
the largest frugivore in our study area and is in sympatry
with only one other primate species, the night monkey
(Aotus lemurinus). Howler monkey populations at our site
were likely greatly reduced when this forest was exploited
in the early- to mid-20th century (Londofio, 1994). Forest
protection and restoration have presumably allowed howler
populations to recover in the last 40 years. Flexibility in
habitat use has allowed howlers to exploit new habitats
such as ash plantations, and they are not limited to mature
forest (Estrada and Coates-Estrada, 1996; Fedigan et al.,
1998; Fedigan and Jack, 2001). At Hato Masaguaral in
Venezuela, for example, densities over 70 ind/km2 have
been reported (Neville, 1972; Rudran, 1979; Crockett and
Eisenberg, 1987). Population size at this site has increased
in part in response to forest recovery (Crockett and Eisen-
berg, 1987; Crockett, 1996). Fedigan and Jack (2001)
found that in 28 years since the creation of Santa Rosa Na-
tional Park in Costa Rica, the population of black howler
monkey (Alouattapalliata) has increased seven-fold, due to
protection and increase in forest cover.

Habitat use
Howler monkeys at Otin Quimbaya used the different
habitat types in proportion to their availability, as has also
been found in the lowlands, where howlers are reported as
habitat generalists (Neville, 1972; Soini, 1982; Stevenson et
al., 1991; Palacios and Rodriguez, 2001). In our six-month
study we observed low intra- and interspecific synchrony in

fruit and new leaf production, and no absolute fruit scar-
city for howlers (Giraldo et al., submitted). Tropical mon-
tane forests do not present drastic periods of fruit scarcity,
in contrast to the lowlands (Giraldo, 1990; Ataroff, 2001;
Cavelier et al., 2001). Quadrant use by monkeys at our site
was dictated by the presence of feeding trees. Probably for
this reason, there was no core area in the home range and
quadrant use was not random.

Differences in structure and composition among habitat
types at Otin Quimbaya generated differences in habitat
use by howlers. In mature forest, the canopy is heteroge-
neous and densities of feeding and sleeping trees are high.
Howlers used feeding trees for a short time and moved
among them, traversing their entire home range in a few
days, as occurs in lowland forest (Stevenson et al., 1991;
Izawa, 1997). Plantations, in contrast, are more homo-
geneous (monodominant canopy) and resource trees are
more dispersed. Thus, howlers spent several days at one
or two fruiting trees until exhausting the fruit crop, and
then moved to another tree, which could be located in a
far quadrant of their home range. This resulted in some
quadrants being used intensively, whereas others were used
only as movement routes. Groups living in plantations also
had a less diverse diet than mature forest groups (Giraldo et
al., submitted). It is unlikely that monkeys could survive in
plantations without neighboring tracts of native forest, and
without having dispersed Ficus and Cecropia trees within
the plantation (Giraldo et al., submitted). The Moraceae
are very important for howler survival in isolated forest
patches, and in disturbed and second-growth forest (Ry-
lands and Keuroghlian, 1988; Schwartzkopf and Rylands,
1989; Estrada and Coates-Estrada, 1996; Fedigan et al.,

Home range and daily distance traveled
Red howler home ranges vary widely, but tend to be small
(6-30 ha). Home ranges at our site were similar to those
reported for lowland forest (Neville, 1972; Defler, 1981;
Crockett and Eisenberg, 1987; Soini, 1992; Izawa, 1997).
Small home ranges in howler monkeys reflect their high use
of leaves, a low-quality nutritional resource that is abundant
and widely distributed (Milton, 1980; Gaulin and Gaulin,
1982; Braza et al., 1983). A study in the Central range of
the Colombian Andes at 2300 m of elevation (Gaulin and
Gaulin, 1982), about 300 km south of our study area, re-

Neotropical Primates 14(1), January 2007

ported a home range of 22 ha and a density ca. 15 ind/km2
for a red howler group in a mature forest. At this site howl-
ers are at the limit of their elevational range, and are sym-
patric with Cebus apella, a very active frugivore-insectivore
that could represent strong competition for howlers.

A previous study at OtGn Quimbaya (Morales-Jimenez,
2003) reported a home range of 14.5 ha for a troop in
mature forest and 21.2 ha for a troop in ash plantation.
This author suggested that the larger home range of the
plantation troop was due to lower resource availability, as
found in our study (Giraldo et al., submitted). The plan-
tation troop studied by Morales-Jimenez (2003) coincides
with our troops D and E, which could suggest that a new
group formed in this area. Fedigan and Jack (2001) found
that population increase of mantled howler monkey at
Santa Rosa National Park in Costa Rica was due to new
group formation as forest recovered. Mantled howlers rap-
idly colonized secondary forest as trees reached a sufficient
diameter at breast height to support their weight.

Home ranges of howler monkeys may decrease when
population densities increase (Crockett and Eisen-
berg, 1987). For example, at La Macarena, Colombia
(67 ha, 17-30 ind./km2; Stevenson, et al., 1991, 2000)
and Capard, Colombia (182 ha, 4 ind./km2; Palacios and
Rodriguez, 2001), densities are low and home ranges
large. In contrast the opposite is observed at Hato Masa-
guaral in Venezuela (7-10 ha, 83-118 ind/km2; Crockett
and Eisenberg, 1987) and OtGn Quimbaya (10.2 ha, 72.6
ind./km2; this study). Home ranges of troops using mature
forest at OtGn Quimbaya were compact and small, whereas
they were elongated and larger for plantation troops. This
reflects structural and compositional differences between
habitat types, as well as patch shape. In plantations the
forest canopy is homogeneous and more discontinuous,
and feeding trees and sleeping trees are more dispersed. In
addition, plantations are in strips along the river and the
road. Groups D and E could cross the road only at cer-
tain points where tree canopies provided a bridge. Thus,
movement routes for plantation troops sometimes were
long and linear. In mature forest, in contrast, movement
routes were more tortuous and uniform, covering similar
distances each day.

Daily travel routes ofhowlers are usually around 500-600 m
in lowland habitats (Neville, 1972; Rudran, 1979; Steven-
son et al., 1991). Howler troops tend to be stable and use
the same routes repeatedly. For example, in ten years of
following a troop at La Macarena, Colombia, Izawa (1997)
observed little changes in travel routes. Mean daily move-
ments at OtGn Quimbaya (317-1321 m/day) were similar,
with variations related to habitat structure.

Home ranges and space requirements of montane red
howler monkeys observed in this study are similar to those
reported for lowland forest populations. The relatively high
population density estimated for OtGn Quimbaya is relat-

ed to the recent history of protection and forest recovery
in the area. Plasticity in habitat use has helped the howler
population to recover, as they have been able to use tree
plantations that offer some resources. Patterns of move-
ment and home range use, however, differ between habitat
types (mature forest vs. secondary forest and plantations).
These differences are related to differences in resource dis-
tribution and availability in the different habitats.


We thank the National Parks Unit and the staff at Otin
Quimbaya for logistical support and for permits. Thanks to
Carolina Murcia and fellow researchers at WCS Colombia/
EcoAndina for continued support and discussion of ideas,
and to Leonor Valenzuela for help with the map. Finan-
cial support was provided by a grant from the John D. and
Catherine T. MacArthur Foundation.


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Neotropical Primates 14(1), January 2007 11


Amrei Baumgarten1 and G. Bruce Williamson1

1Department of Biological Sciences, Louisiana State University, 107 Life Science Building, Baton Rouge, Louisiana 70803, USA


We studied the geographic distribution of the endangered black howler monkey, Alouattapigra, in its southeastern range in
eastern Guatemala along its putative contact zone with the mantled howler monkey, A. palliata. A distributional survey of
both species was conducted through direct observations and interviews, and the data analyzed with GIS to detect potentially
limiting geographic and ecological factors. Alouatta pigra was not associated with any specific vegetation type or altitudinal
range, as it occurred in many forest types from the lowlands up to elevations of 2705 m a.s.l., in contrast to prior reports of
it being restricted to elevations below 400 m a.s.l. and having a strong affinity for riverine forests. Alouattapalliata did not
occur at the higher elevations. Distribution of both howler monkey species was unaffected by the presence of Ateles, by pro-
tection status of the survey site, or by hunting activities. Deforestation appears to have restricted both species in some areas.
Throughout the southeastern range of A. pigra, we found no evidence for current sympatry with A. palliata, as their distribu-
tions are separated by a river barrier and unsuitable dry shrubland. Ecological preferences also seem to reinforce allopatry,
with only A. pigra occurring in wet and cold montane forests of the Sierra de las Minas. Our data indicate that highland
habitats in Mexico and Guatemala, previously considered unsuitable for A. pigra, may need to be evaluated for distribution
assessments and management plans. Populations of A. pigra in the diverse protected ecosystems in eastern Guatemala may
be an important component for conservation of the species.

Key Words: Alouatta pigra, A. palliata, geographic distribution, allopatry, eastern Guatemala


Se estudi6 la distribuci6n geografica del mono aullador negro, Alouatta pigra, y el mono aullador de manto, A. palliata, en
el este de Guatemala, en donde reports previous proponen traslape de rangos. Esta region constitute el limited sureste del
rango de A. pigra, especie en peligro de extinci6n y end6mica del sur de M6xico, Belice y Guatemala. La distribuci6n de
ambas species se determine mediante observaciones directs y encuestas. Los datos fueron analizados con SIG para detec-
tar posibles factors limitantes, tanto ecol6gicos como geogrificos. La presencia de A. pigra no estuvo asociada con ningun
tipo de vegetaci6n particular o rango altitudinal, encontrindose la especie en various tipos de bosque desde los 0 hasta los
2705 msnm. Estos datos no coinciden con studios previous que described a A. pigra como una especie restringida a eleva-
ciones por debajo de los 400 msnm y con alta afinidad por bosques riberefios. Alouattapalliata no se registry en altitudes
elevadas. La distribuci6n de ambas species de monos no se vio afectada por la presencia de Ateles, por el estado de protecci6n
del sitio de muestreo, ni por actividades de caceria. En algunas areas las species se ven restringidas por la deforestaci6n. No
se encontr6 evidencia de simpatria ni traslape de rangos. Los rangos de ambas species se separan por un rio y por habitat
inadecuado de bosque seco. Ademis, preferencias ecol6gicas parecen reforzar la alopatria, puesto que dnicamente A. pigra
ocurre en los bosques humedos y frios de la Sierra de las Minas. Nuestros datos sugieren que habitats montanos en M6xico
y Guatemala previamente categorizados no aptos para A. pigra, necesitarian considerarse en evaluaciones de distribuci6n y
planes de manejo. Las poblaciones de A. pigra en los diversos ecosistemas protegidos en el este de Guatemala podrian ser
importantes components en la conservaci6n de la especie.

Palabras Clave: Alouatta pigra, A. palliata, distribuci6n geogrifica, alopatria, oriented de Guatemala

Introduction assessments and action plans for the species mandate up-
dating distribution maps and describing the status of wild
The black howler monkey, Alouatta pigra, is endemic to populations (Rodriguez-Luna et al., 1996; Matamoros et
Guatemala, Belize and southern Mexico. Recently, it al., 1997).
was categorized as endangered in the IUCN Red List of
Threatened Species due to population size reduction based Limited knowledge of the geographic distribution of
on habitat decline (Cuar6n et al., 2003). Conservation A. pigra is based on museum specimens (Smith, 1970;

Neotropical Primates 14(1), January 2007

Hall, 1981) and broad field studies done in Belize and
Mexico (Horwich and Johnson, 1986; Watts et al., 1986)
and Guatemala (Curdts, 1993). Recent fieldwork in the
Yucatan peninsula detailed the species' distribution and
habitat preferences (Navarro et al., 2003; Serio-Silva et
al., 2006), but other areas of Mexico and Guatemala need
similar detailed studies. Of particular importance are po-
tential contact zones with the mantled howler monkey, A.
palliata, at the northwest and southeast range limits, where
sympatry has been reported in Tabasco, Mexico (Smith,
1970; Cortes-Ortiz et al., 2003) and suspected in eastern
Guatemala (Horwich and Johnson, 1986; Curdts, 1993).
Contact zones are important for the preservation of pri-
mate biodiversity (Jones and Bicca-Marques, 2004) and es-
sential for the study of genetics and ecology of speciation
(Jiggins et al., 1996).

The studies in southern Belize and eastern Guatemala
have resulted in ambiguous and inconclusive definition
of ranges for the two species. For example, Horwich and
Johnson (1986) identified sympatry at the Belize-Guate-
mala border around the Sarstdn River, based on sugges-
tions of palliata-like individuals on the Guatemala side
of the river, where only A. pigra was expected. Curdts
(1993) proposed large areas of sympatry in the moun-
tains of Baja and Alta Verapaz and Sierra de las Minas,
Guatemala, but provided no empirical evidence. The ob-
jective of the present study was to define the geographic
distribution of A. pigra and A. palliata in their contact
zone in eastern Guatemala at the southeastern range
limit of A. pigra. In addition we hoped to identify eco-
logical factors and geographic barriers that might affect
both species' distributions.


Study area
The contact zone and putative area of sympatry of
A. pigra and A. palliata lies within 880-90 W longitude
and 15-16 N latitude, encompassing the eastern part
of Guatemala, bounded by Belize, Honduras and the
Atlantic Ocean (Fig. 1). Five Guatemalan departments
are partly or totally within the area: Alta Verapaz, Baja
Verapaz, Izabal, Zacapa and El Progreso. The area is to-
pographically and ecologically heterogeneous with eleva-
tions ranging from sea level to 3,000 m a.s.l. (CONAP,
2005) and ecosystems varying from flooded coastal forest
to montane cloud forest. The region is transversed by
several mountain ranges and large rivers (Fig. 1). Con-
tinuous tracts of tropical forests are restricted primarily
to protected areas. Forest conversion by traditional slash-
and-burn maize cultivation is combined with intensive
socioeconomic activities: coffee and cardamom planta-
tions in the highlands, and cattle ranching, banana, Af-
rican palm and rubber plantations in the lowlands. The
Department of Izabal, which occupies a large part of the
study area, has 20% of its total area in cattle pastures and
crop monocultures (INAB, 2001).

Following Brockelman and Ali (1987), we conducted a
distributional field survey during the dry season, February
to April 2005. Cartographic maps (1:25000), vegetation
maps, and a LandsatTM satellite image (2003) were used to
identify suitable monkey habitat, access routes, and survey
sites. Survey sites were selected to test potential geographic
barriers and to cover the altitudinal ranges and vegetation
types of the region. Additionally, site selection was depen-
dent on the presence of suitable forest cover and existence
of key persons who would facilitate entrance to villages,
contact appropriate interviewees, and serve as translators
in K'ekchi villages. Survey sites were accessed using vehicle,
boat, mule or by foot. At each survey site geographic loca-
tion and altitude were recorded with a Garmin GPS 72.

Howler monkey occurrences were detected by two meth-
ods: interviews (Pinto and Rylands, 1997; Iwanaga and
Ferrari, 2002) and broad forest surveys along trails and
rivers (Brockelman and Ali, 1987). Interviews consisted
of non-leading questioning about the primates in the area,
as well as other questions related to the interviewee's occu-
pation, residence time in the area, and hunting activities.
Additionally, interviewees were asked to recognize photo-
graphs of possible local primate species (Alouatta, Ateles,
Cebus) and South American Alouatta species, as well as
playbacks of loud calls of A. pigra and A. palliata. People
interviewed were mainly subsistence hunters, town elders
with knowledge of the forest, park rangers, landowners,
and field biologists. Forest surveys were made during
3-day visits to key sites in early mornings and late after-
noons during peak hours of howler monkey vocal activity,
in order to locate individuals and make visual and/or audi-
tory identification.

Species presence/absence records obtained from valid in-
terviews and/or direct observations at each site were incor-
porated into a Geographic Information System (ArcView
version 3.3). Digital maps of the vegetation types of Gua-
temala (INAB, 2001; CCAD-WB, 2003), protected areas,
and elevation curves were overlaid with the geographical
coordinates of sites to create joined attribute tables and
allow analyses of the distribution of howler monkeys in
relation to physical and ecological features. Information
collected through interviews on hunting pressure and on
the occurrence of the spider monkey, Ateles ..- ..i was
also related to the presence/absence records of the howler
monkeys. A search of collection databases of 18 natural his-
tory museums in North America and Europe and a review
of literature were made to obtain any historic records of the
occurrence of A. pigra and A. palliata in the region.

Statistical analyses
We used tests for homogeneity of proportions with a lo-
gistic analysis approach to detect habitat-specific distribu-
tional patterns. Specifically, we used a nominal model to
test if vegetation type had an effect on the probability of
occurrence of each species and an ordinal model to test for

Neotropical Primates 14(1), January 2007

a trend in probability of occurrence along an altitudinal
gradient. Also, tests of independence were used to assess
howler monkey association with (a) the presence of Ateles
and (b) the protection status of the site. All statistical tests
were performed with SAS version 9.0.


A total of 58 sites were visited (Fig. 1); interviews were
conducted at 47 sites and direct observations at 23 sites.
We conducted 97 interviews, 59% with ladino and 41%
with K'ekchi interviewees. Five interviews were classified
as invalid due to inconsistencies in descriptions or identi-
fications of photographs. Playbacks proved useful only to
identify A. palliata, as interviewees clearly distinguished the
calls. In contrast, at A. pigra sites, interviewees could not
distinguish between the two species' calls. Overall, Alouatta
pigra was found at 26 sites: seven through both interviews
and direct observations, seven through observations only,
and 12 sites through interviews only (Table 1). Alouatta
palliata was found at 12 sites: five through interviews and
direct observation, three through observations only, and
four through interviews only (Table 2).

Alouatta pigra was verified along both banks of the Rio
Sarstdn, the Atlantic coast, the north shore of Lake Izabal-
Rio Dulce, both banks of the Rio Polochic and inland in
the Purulha mountains, Sierra Yalijux, Sierra Santa Cruz,
and Sierra de las Minas (Fig. 1, Table 1). Alouatta palliata
was verified along the Atlantic coast south of the Rio Dulce
and inland along the Rio Motagua valley and in Sierra
Caral (Fig. 1, Table 2).

Of five potential vegetation types, A. pigra was found in
four and A. palliata in three (Table 3). Neither species oc-
curred in "seasonal evergreen shrubland with mixed forest".
Only A. pigra occurred in "tropical evergreen and semiever-
green mixed forest" typical of cloud forest. Alouattapalliata
was most often in "agroproductive systems with significant
portions of broad-leaved forest", although the associa-
tion was not statistically significant (test of homogeneity
of proportions 2 = 7.6, df= 4, p = 0.10). In contrast, A.
pigra, was less frequent in this disturbed vegetation type
and more commonly associated with flooded forests, mixed
forests and broad-leaved forest (test of homogeneity of pro-
portions 2 = 14.41, df = 4, p < 0.01). Survey sites were
distributed across elevations from sea level to more than

Table 1. Localities where Alouatta pigra occurs (see Fig. 1), protection status, and detection method.
Site No. Locality Protection status Method
1 Chelema private reserve sighting, interview
2 Lowland south of Chichipate- 11,. None sighting, interview
4 Selich .- 11 ,. None interview
6 5.. .. ...II.c. None interview
7 Boquer6n None sighting
8 Sakitzul finca proposed private reserve interview
9 Guitarra- .11 '. None interview
14 Calaja 11 -. None interview
15 Rio Sarstun, Belize side Sarstoon Temash Reserve sighting
16 Rio Sarstun, Guatemala side Rio Sarstun Multiple Use Area sighting
17 Rio Sarstun, Belize side Sarstoon Temash Reserve sighting
18 Sarstn i- I i -. Rio Sarstun Multiple Use Area sighting, interview
19 Calix Black creek finca None interview
20 Choc6n-Machacas scientific station Choc6n Machacas Biotope interview
21 Mario Dary Biotope Mario Dary Biotope interview
22 Las Cabanas scientific station core area SM BR' sighting, interview
24 trail to Volcan Las Palomas core area SM BR1 sighting
25 Alejandria finca multiple use zone SM BR' sighting, interview
26 Los Angeles d I,. buffer zone SM BR1 interview
27 San Vicente II .11 buffer zone SM BR' interview
28 Manguitos II -Ill I-. lowlands None interview
30 Semuy II ,ill ,. lowlands Bocas Polochic Wildlife Reserve interview
32 Selempin biological station Bocas Polochic Wildlife Reserve sighting, interview
33 Rio Oscuro Bocas Polochic Wildlife Reserve sighting
34 Lake Izabal shore Bocas Polochic Wildlife Reserve sighting
35 Naranjal Yaxte 11 ,. None sighting, interview
' SM BR: Sierra de las Minas Biosphere Reserve.

Neotropical Primates 14(1), January 2007

Figure 1. The study area in eastern Guatemala, showing the location of the survey sites, mountains, major rivers, and department limits.
Asterisk shows the historic location of A. pigra in Quirigui, Izabal.

Table 2. Localities where Alouatta palliata occurs (see Fig. 1), protection status, and detection method.
Site No. Locality Protection Status Method
41 Juan Vicente Creek Rio Dulce National Park none sighting, interview
42 Casa Guatemala Rio Dulce National Park none sighting, interview
43 Rio Frio lowland None interview
45 Cayo Piedra finca Rio Dulce National Park none sighting
50 Laguna Santa Isabel shore Punta Manabique Wildlife Reserve sighting
51 Estero Lagarto i ,. Punta Manabique Wildlife Reserve sighting, interview
52 Cabo Tres Puntas .11 ,. Punta Manabique Wildlife Reserve sighting, interview
53 San Vicente Paul 11 ,. None interview
54 San Vicente Paul highlands proposed Sierra Caral reserve none interview
56 Animas .11 ,. highlands proposed Sierra Caral reserve none interview
57 Champas finca lowlands None sighting, interview
58 Cacaol .11 'c. None sighting

Table 3. Vegetation types (UNESCO classifications following INAB, 2001) of the survey sites and number of sites with presence/absence
records for each species.
No. of No. of No. of No. of sites No. of sites
Vegetation type No. of sites with sites with with both without howler
A. pigra A. palliata spp. monkeys
1. Tropical evergreen broad- leaved periodically flooded forest 13 10 3 0 0
2. Tropical evergreen and semievergreen broad-leaved forest 21 9 3 0 9
3. Tropical evergreen and semievergreen mixed forest' 6 4 0 0 2
4. Seasonal evergreen shrubland with mixed forest1 3 0 0 0 3
5. Agroproductive systems with significant portions of 15 3 6 0 6
Broad-leaved forest
Total 58 26 12 0 20

'Mixed forest = broad-leaved and needle-leaved forest

Neotropical Primates 14(1), January 2007

Table 4. Altitudinal ranges (following INAB, 2001) of the survey sites and number of sites with presence/absence records for each

Altitudinal Number of sites No. of sites with No. of sites with No. of sites with wiou.t oier
descriptor Elevation (m) Number of sites A. pigra A. palliata both spp. monkeys
Lowland 0-499 39 18 10 0 11
Sub-montane 500-999 8 1 2 0 5
Lower montane 1000-1499 5 2 0 0 3
Upper montane 1500-2000 3 2 0 0 1
Altimontane >2000 3 3 0 0 0
Totals 58 26 12 0 20

2500 m a.s.l. (Table 4). Alouattapigra was found in all alti-
tudinal ranges (Table 4) and no ordinal association was de-
tected (test of homogeneity of proportions, ordinal model
x2 = 1.61, df = 1, p = 0.20). The altimontane survey sites
(> 2000 m a.s.l.) were all occupied by A. pigra, the highest
being 2705 m a.s.l. in Sierra de las Minas. In contrast, A.
palliata occurred significantly more frequently in lowland
sites and was not recorded at elevations higher than 1000
m a.s.l. (test of homogeneity of proportions, ordinal model
2 = 4.07, df= 1, p < 0.05).

Of the 10 major protected natural areas surveyed, 80%
sustained howler monkey populations: Alouatta pigra was
found in six and A. palliata in two (Tables 1 and 2). Of the
total survey sites, 32 sites (55%) were in protected areas
with enforcement and 26 sites (45%) were located outside
protected areas or in protected areas without vigilance. Oc-
currence of howler monkeys was not associated with the
protection status of the survey site (test of independence
G2 = 0.33, df = 1, p = 0.59).

Of 97 interviewees, 76% denied hunting or ever hearing
of anyone hunting howler monkeys. Most (19 of 22) of
the remaining 24% responded that they had only heard of
other people killing howler monkeys and did not consider
them a valuable bush meat; two respondents used howler
monkeys as bait for river shrimp, and one hunted these
primates for meat and medicine (howler monkey broth
was used as treatment for respiratory illness). Howler mon-
keys kept as pets were rare. The reports on hunting activity
were distributed across the survey sites, unrelated to ethnic
group of interviewees or geographic region. The only other
primate species in the area reported in the interviews was
the spider monkey Ateles I. .. The white-faced capu-
chin monkey, Cebus capucinus, was not known to locals
and probably does not extend into Guatemala. Records for
Ateles ..O .. presence were slightly lower than those for
Alouatta species (34% and 40% of interview sites, respec-
tively), and its range overlapped with both A. palliata and
A. pigra. Ateles occurred in sites with and without howler
monkeys and there was no evidence of association or dis-
sociation with Alouatta (test of independence G2= 0.11, df
= 1, p = 0.76).

No evidence was found for a current zone of sympatry be-
tween Alouatta pigra and A. palliata. Only one interviewee
claimed to have seen both species in mixed troops, but this
was disputed by other interviewees at the same site (site
19) and it conflicted with records from surrounding sites as
well. The identified range limits for A. pigra are: Lake Izabal
and Rio Dulce in the east, unsuitable habitat of pine forest
west of Purulhi in Baja Verapaz in the southwest and pine
forest and dry shrubland in foothills of Sierra de las Minas
in the south (Fig. 1). The southeastern range limit is not as-
sociated with any barrier. We found the most southeasterly
populations of A. pigra in the tip of Sierra de las Minas
(site 35), and historical records place the species farther
east in Quiriguk in the middle Motagua valley (Salvin and
Goodman, 1879; museum specimen in the Smithsonian
National Museum of Natural History, USNM 238704)
(Fig. 1). The western range limit of A. palliata is dry de-
ciduous shrubland. The heavily disturbed area to the south
of Lake Izabal in the lower Motagua valley is the northwest
limit and the Rio Dulce the north limit.


This study reports the distribution of the endangered black
howler monkey, Alouatta pigra, in its southern geographic
range in topographically and ecologically heterogeneous
eastern Guatemala, providing needed information on de-
tailed occurrence localities (Rodriguez-Luna et al., 1996;
Matamoros et al., 1997) and new reports of its ecological
tolerances. It also contributes to the limited knowledge of
the status and distribution of the mantled howler monkey,
A. palliata, in Guatemala. Alouatta pigra was widespread
and occupied several different habitat types across altitudi-
nal ranges from sea level up to 2705 m. Our results show
no evidence of significant association with a specific veg-
etation type or altitude. In his preliminary study in Gua-
temala, Curdts (1993) also found A. pigra at high eleva-
tions above 2000 m a.s.l. These results contrast with the
reports that A. pigra is restricted to elevations below 400
m a.s.l., with a strong affinity for riverine forest (Horwich
and Johnson, 1986; Watts et al., 1986; Ostro et al., 2000).
Models of the species' current range based on preference for
lowland forests (Luecke, 2004) need to consider potential

Neotropical Primates 14(1), January 2007

premontane and montane habitats to avoid underestima-
tion of actual range.

Forest disturbance by humans influences the distribution
patterns of A. palliata and A. pigra in the region mainly
by eliminating suitable monkey habitat. This is especially
the case along the Rio Motagua Valley and south shores
of Lake Izabal and Rio Dulce. Here extensive monocul-
tures and cattle pastures have replaced the original tropi-
cal forest (INAB, 2001), which may account for the high
frequencies of A. palliata in disturbed vegetation types.
The ability of A. palliata to survive in fragmented habi-
tats is widely recognized by other authors (e.g., Estrada and
Coates-Estrada, 1984; Silva L6pez et al., 1988; Clarke etal.,
2002). Deforestation in the range of A. pigra occurs mainly
in the upper and middle Rio Polochic valley, where the spe-
cies was once very common (Salvin and Godman, 1879),
as well as north of Lake Izabal. The species was present in
some disturbed sites, in accordance with other reports of A.
pigra thriving in forest fragments (Horwich and Johnson,
1984; Baumgarten, 2000; Estrada et al., 2002).

The absence of A. palliata from the Cerro San Gil area (sites
44, 47, 48, 49) was unexpected, since the area has protected
status and extensive forest cover with other wildlife, includ-
ing the spider monkey. The premontane and montane el-
evations at these sites may act as a limiting factor for the
species. Other studies have also found that A. palliata is
absent from elevations above 700 m a.s.l. in mountainous
ranges in Mexico (Silva L6pez et al., 1988; Estrada and
Coates-Estrada, 1996). The species is present at elevations
up to 1500 m a.s.l., but only at lower latitudes in Central
America (Timm et al., 1989). The red howler monkey, Al-
ouatta seniculus, occurs at high elevations in the Colombian
Andes (Hernindez-Camacho and Cooper, 1976; Gaulin
and Gaulin, 1982).

We found little evidence for hunting pressure on howler
monkeys and no association between hunting and the dis-
tribution of the two species. Hunting of howler monkeys
is also limited in Belize (Horwich and Johnson, 1984) but
more common in Mexico (Estrada and Coates-Estrada,
1984; Horwich and Johnson, 1984; Silva L6pez et al.,
1988). In contrast, subsistence hunting has accounted
for local extinctions of Alouatta in localities in the Brazil-
ian Amazon (Peres, 1990) and the Peruvian and Bolivian
Amazon (Freese et al., 1982). Alouatta occurrence may be
influenced by competition with other primates in some
habitats (Eisenberg, 1979), although not always (Peres,
1997; Iwanaga and Ferrari, 2002). We found no evidence
for competitive exclusion between Ateles and Alouatta spe-
cies. In Mexico the spider monkey Ateles is more wide-
spread than Alouatta and sympatric with it in only a few
places (Watts et al., 1986; Rodriguez-Luna et al., 1987;
Silva L6pez et al., 1988).

The study region has a high potential for the conservation
of the endangered black howler monkey, A. pigra, since six

of its protected areas harbor the species (Table 1), the largest
being the Sierra de las Minas Biosphere Reserve (246,803
ha) (CONAP, 2005). Furthermore, the low hunting pres-
sure and the occurrence of the species outside protected
areas provides the opportunity for community-based con-
servation projects and sustainable resource programs, which
have proven effective for howler monkey conservation in
Belize and Mexico (Horwich, 1998). The occupied habi-
tats are ecologically heterogeneous in comparison with the
species' range in the Yucatin Peninsula, which is predomi-
nantly lowland rainforest (Horwich and Johnson, 1986;
Watts et al., 1986; Navarro et al., 2003). The populations
of A. pigra living in cloud forests at high elevations are an
important component in conserving the species variability
and its gene pool. Previously considered to be unsuitable
habitats, highlands will have to be considered in distribu-
tion assessments and management plans for the species.
The severe deforestation south of Lake Izabal-Rio Dulce
puts the populations of the mantled howler monkey A. pal-
liata at risk in Guatemala. These are the most northerly
populations of the subspecies A. palliata palliata. Impor-
tant conservation actions here include conservation in the
protected areas without enforcement, the legal declaration
of the proposed Sierra Caral protected area, and the estab-
lishment of the Mesoamerican Biological Corridor project
that connects Punta de Manabique Wildlife Reserve with
protected areas in Honduras.

The results of this study are relevant to clarifying the dis-
tribution of Alouatta pigra and A. palliata in their contact
zone in eastern Guatemala. We found no evidence for cur-
rent range overlap or sympatry as well as no support for
the previous proposed sympatric areas around Rio Sarstun
(Horwich and Johnson, 1986) or in the highlands of Baja
Verapaz, Alta Verapaz, and Sierra de las Minas (Curdts,
1993). In the northwestern part of the putative contact
zone, the Rio Dulce acts as a physical barrier separating
both species. Curdts (1993) also reported A. pigra on the
north bank and A. palliata on the south bank of this river.
River boundaries are often limiting factors for the distri-
bution of primates (Ayres and Clutton-Brock, 1992; Wal-
lace et al., 1996). In the eastern part of the contact zone,
A. pigra occurs on the southwest side of Lake Izabal and
tip of Sierra de las Minas. South of Lake Izabal suitable
monkey habitat has been removed and A. pigra is known
only from historic records. Further east in the lower Mo-
tagua Valley we registered exclusively A. palliata. It is
possible that a narrow sympatry existed south of the lake
before forest loss. Further range overlap is not evident,
probably explained by ecological differences. Our results
suggest that A. palliata is associated with forests found at
low elevations, precluding its expansion into the wet and
cold habitats of Sierra de las Minas where A. pigra is found.
Habitat preferences seem to act as a barrier between howler
monkey species in other cases of near sympatry (Crockett,
1998) as reported for A. fusca and A. caraya in northern
Argentina (Di Bitetti et al., 1994) and A. seniculus and
A. caraya in southwestern Amazonia (Iwanaga and Fer-

Neotropical Primates 14(1), January 2007

rari, 2002). Further, A. pigra's lack of wide expansion into
A. palli,,m.' range in the lowlands may perhaps involve
assortative mating or hybrid inviability.

In the extreme south of the potential contact zone, the
ranges of both species are interrupted by dry deciduous
shrubland south of the Sierra de las Minas. The semidesert
characteristics of this vegetation clearly constitute an effec-
tive ecological barrier between the species. In contrast to the
scenario in Guatemala, A. pigra and A. palliata in Mexico
have a broad sympatric area that extends over the lowlands
of the states of Tabasco and Campeche. Nevertheless, spe-
cies introgression seems to be limited as Smith (1970) found
no evidence for hybridization based on museum specimens
from the same localities, although occasional interbreeding
may occur in mixed troops of howler monkeys in the area
(Cortes-Ortiz et al., 2003).


This study was supported by two Louisiana Governor's
Office of Environmental Education Research Grants. We
are grateful to FUNDAECO, Fundaci6n Defensores de
la Naturaleza, FUNDARY, CECON-USAC, Parroquia
El Estor, Finca Tijax, Finca Cilix and Finca Champas for
technical support and research permission. Special thanks
to all interviewees and enthusiastic guides and to Daniel
Tenes, Carlos Avendafo, Haroldo Garcia and Jos6 Moreira
for assistance in the field. We thank Kellen Gilbert for com-
ments on this manuscript. This study was carried out with
the required permits from the Consejo Nacional de Areas
Protegidas-Guatemala. The senior author was supported by
a grant from the Fulbright Faculty Development Program
Central America.


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in disturbed forest areas of Sierra Santa Marta, Mexico.
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their distribution and status. Primate Conservation (7):

Neotropical Primates 14(1), January 2007 19

Swen C. Renner1, Regula Rieser2 and Robert H. Horwich3

1 Conservation & Research Center, National Zoological Park, Smithsonian Institution, 1500 Remount Road, Front Royal, VA
22630, USA, e-mail:
2 Universitdt Ziirich, Winterthurerstrasse 190, CH-8057 Ziirich, Switzerland
3 Community Conservation, 50542 One Quiet Lane, Gays Mills, WI 54631, USA


Groups of the Guatemalan black howler monkey (Alouatta pigra) in a fragmented northern Guatemalan cloud forest in
the Sierra Yalijux were analyzed for group composition and size using an ornithological census technique. Audio and visual
records were combined to estimate each group's home range size. The 361 observations in 2002 estimated 50 individuals in
seven groups. Two home range classes were observed with 1.87 ha and > 2.87 ha. Compared to other recent studies, the
density of A. pigra in the Sierra Yalijux is considerably higher than in other areas (83.3/km2 vs. 15.1/km2 in Quintana Roo
and 23.0/km2 in Palenque) but about half the density found in Belize (178/km2). We discuss the relevance of the .r ir-.. 1..
assessment method for howler monkeys.

Key words: Alouatta pigra, Neotropics, Guatemala, cloud forest, Sierra Yalijux, ecology, GIS, home range, minimal convex
polygon, saraguate


Utilizando una tecnica ornitol6gica de censos se analizaron la composici6n y tamafio de grupos del mono aullador negro
Guatemalteco (Alouatta pigra) en un bosque nublado fragmentado al norte de Guatemala en la Sierra Yalijux. A partir de
361 observaciones hechas en 2002 se estimaron 50 individuos en siete grupos. Se encontraron dos classes de tamafio de area
de uso vital 1.87 ha y > 2.87 ha. Comparada con aquellas de otros studios recientes, la densidad de A. pigra en la Sierra
Yalijux es considerablemente mris alta que en otras areas (83.3/km2 vs. 15.1/km2 en Quintana Roo y 23.0/km2 en Palenque)
pero cerca de la mitad de la densidad encontrada en Belice (178/km2). Discutimos la relevancia del m6todo de evaluaci6n
ornitol6gico utilizado para los monos aulladores.

Palabras Clave: Alouatta pigra, Neotr6picos, Guatemala, bosque nublado, Sierra Yalijux, ecologia, GIS, area de dominio
vital, minimo poligono convexo, saraguate


The black howler monkey, Alouatta pigra, endemic to Me-
soamerica, is threatened due to widespread habitat destruc-
tion causing severe population declines. The IUCN 2004
Red List of Threatened Species classifies A. pigra as Endan-
gered (A4c) (see Rodriguez-Luna et al., 1996a, 1996b;
Crockett, 1998; Cuar6n et al., 2003), and it is listed on
Appendix I of CITES the Convention on International
Trade in Endangered Species of Wild Flora and Fauna. A.
pigra occurs in the tropical and semi-deciduous forests of
Yucatan (Mexico), Belize, western Honduras and northern
parts of Guatemala (Emmons and Feer, 1997; Reid, 1997).
A. pigra is highly territorial, with each group (generally of
about five to ten individuals) occupying relatively exclusive
ranges (Reid, 1997).

Currently, there are considerable regional differences in
the conservation status of the black howler, with popula-
tions in areas with high deforestation being more threat-
ened than others. Populations of A. pigra in Belize are
comparatively well-studied (e.g., Horwich and Johnson,
1984; Horwich etal., 2001a, 2001b; Estrada et al., 2002a,
2002b, 2004). Although A. pigra is thought to occur at
lower elevations (Horwich and Johnson, 1986), the status
of A. pigra in the central mountains of Guatemala remains
relatively unknown. We here add data on a highland pop-
ulation of A. pigra and discuss the use of an ornithologi-
cal census method applied to assess howler monkeys. We
discuss density and territoriality of the A. pigra highland

Neotropical Primates 14(1), January 2007


Study site
We studied a population of A. pigra in a tropical cloud
forest on the northern slopes of the central mountain
ridge of northern Guatemala (Fig. 1), in the southern-
most part of the species' distribution (Emmons and Feer,
1997). The study site is near the community of Chelemhi,
10.8 km north of Tucurd, Alta Verapaz (central co-ordi-
nates: 90004'W, 1523'N; 1,980-2,550 m a.s.1.). The site is
part of the Sierra Yalijux which merges in the west with the
Sierra Caquipec (9011 'W, 1523'N, 2,000-2,200 m a.s.1.)
12 km west of Chelemhi. To the north are the lowlands of
El Peten.

Human disturbance in the primary forest of the study area
is limited to subsistence hunting and occasional logging.
The forests of the highlands of the Sierra Yalijux, while
isolated from other forests, are relatively intact, with only
3.08% being lost between 1986 and 2000. This is a rela-
tively low mean annual deforestation rate of 0.2% (Voigt,
2004; Markussen and Renner, 2005; Renner et al., 2006).
Neighboring the forests of the Sierra Yalijux are two further
large forest fragments, totaling 5,500 ha (Sierra Yalijux:
2,200 ha; Sierra Caquipec: 3,200 ha; unnamed fragment
in between: 100 ha) of mature pine-oak cloud forests
(Markussen, 2004; Renner et al., 2006). All three forest
fragments are separated from each other (Markussen and
Renner, 2005; Renner et al., in press), and are 52 km east
and 15 km north, respectively, from the nearest mature
highland cloud forests of Sierra de Chamaa and Sierra de
las Minas. The nearest lowland rainforest of El Peten is
25 km north of the forest fragments. A. pigra is not report-
ed there and is most unlikely to migrate such a distance,
even though they may cross open areas on the ground.
The three fragments are not fragmented themselves and
each consists of closed forest cover. Further forest patches
of small size (< 20ha) are found in the surrounding areas
of these three major fragments. All forest is surrounded
by different land use, including secondary vegetation and
taller secondary forest. This tall secondary forest is ap-
proximately 15 years old and 20 m tall and has already
established two vegetation strata (Renner et al., 2006b).

The deforestation rate in the Sierra Yalijux was around
0.2% for 1986 to 2000 (Markussen, 2004; Voigt, 2004;
Markussen and Renner, 2005; Renner et al., in press) and
low for Guatemala (country mean 1.7%). For further de-
tails on the region refer to Markussen and Renner (2005),
Renner (2003, 2005) and Renner et al. (2006a, 2006b).

Census protocol
We applied a survey method in Chelemhi that is com-
monly used in bird surveys. Triangulation and assessment
of vocalizing male individuals has been established in orni-
thology for a long time. The method is commonly used to
assess individual bird territories or relative abundance for
birds in temperate forests and in the tropics either applying

Figure 1. The study site (white circle) in Guatemala and altitudes
>1,000 m (gray shaded area) and >2,000 m (black shaded area).

point counts or transect-assessments. An observer records
all songs and sights along a transect or point count grid
and notes species, time, date and estimated or measured
distance from the census grid. Here we used the established
ornithological transect grid (Renner, 2003; Renner et al.,
2006) to assess the howler monkey population. The observ-
ers (SCR, RR) noted time, direction and estimated distance
of howling individuals. One observer (RR) then located the
howling groups and visually observed the groups in greater

We carried out a survey of A. pigra along a 2,000 m tran-
sect line near the community of Chelemhi between March
and October in 2001 and 2002. We surveyed all transects
on a monthly basis for five days per month, from 08:00 to
14:30. From July 22 to September 26, 2002, surveys were
carried out daily. Each survey day we proceeded slowly
along transects. We recorded all instances of howling,
determining the direction by compass bearing and esti-
mating the distance in three categories (close: < 100 m,
medium: 100-300 m, and far: > 300 m). To determine
the exact observer's location we established fixed points
every 25m along transects using global positioning sys-
tems (GPS). When two observers were carrying out the
survey the troops were located by triangulation. Once we
heard or saw a howler group we followed it as long as pos-
sible to determine their troop composition as to age and
sex as described by Reid (1997) and Emmons and Feer

Groups were considered to be discrete units when audio
observations indicated spatial distances of more than
100 m. Groups A to D and G (exceptions: E and F;
Table 1) were recorded by simultaneous vocalizations four


Neotropical Primates 14(1), January 2007

times during the study period with an indicated in-between
difference of more than 100 m. Often adjacent groups will
roar at each other in close proximity at territorial borders.
Sometimes it is impossible to tell whether they are different
troops until they move away from each other and the terri-
tory border (Horwich, pers. obs.). Thus, in addition to the
audible locating procedures, all groups (A to G; Table 1)
were observed visually several times. Groups that could not
be seen were not included.

We mapped all locations of A. pigra troops in the study
region with ArcGIS 9.1 and classified the land cover by
analyzing a Landsat ETM+ scene (path 020, row 049 of
January 23, 2000) with ground-truthing data (Renner and
Markussen, in press). For analyses of the home range we
used the ArcView 3.3 Animal Movement extension to cal-
culate the minimal convex polygon (MCP) and the kernel
home range (KHR). For the KHR we used the 25, 50, 75
and 95% confidence intervals and plotted the results. For
the total population estimate we added the maximum ob-
served troop sizes and assumed this cumulative number
represents the total number of individuals in the study

We also established a transect of 2,000 m in the Sierra
Caquipec close to the small village of Chicacnab, and for
14 days surveyed the A. pigra population. The Sierra Ca-
quipec is the western extension of the Sierra Yalijux but
the forests, with similar pine-oak cloud forest vegetation,
are isolated by a 500 m gap. This second site was included
so that we could have some idea as to whether the popula-
tion density in Chelemhi is high or low compared to other
forest fragments in the region.


We observed seven groups ofA. pigra with 50 individuals in
the 60.05 ha study site near Chelemhi (Fig. 2, Table 1). All
groups were separable by territorial vocalizations and visual
observations. Home range size varied from 1.01 to 5.53 ha
(mean 3.03 ha 1.69 sd.) as measured with the minimal
convex polygons (Table 1). The seven groups ranged in size
from six to ten individuals with 50 individuals in total. The
mean number of males per group was 1.50 0.55, and the

number of females was 1.33 0.55 (Table 1). The popu-
lation density of the forest fragment was 83.3 individuals
per km2.

Further groups were registered by their vocalizations out-
side our study plot (Fig. 2). One group occupied a prima-
ry forest patch (isolated from the Sierra Yalijux; 17.80 ha)
approximately 700 m to the south of group D. Two other
groups were located 500 m north and 900 m west of
group A, and another two troops approximately 1.1 km
and 2.2 km, east-north-east of group G; the latter four
all within the Sierra Yalijux. However, the home range
size estimate for the distant groups is very rough, because
they were far away and never seen. The distance increased
the error for area-estimation. We also observed howlers in
the vicinity of the study area along a path of 4,500 m;
that path was used on a monthly basis during the study

A. pigra in Chelemhi exhibited more frequent territorial
howling than in Chicacnab (12 km west of Chelemhi).
In Chicacnab we only heard one group howling 500 m
away but never could observe them. The surveys close to
Chicacnab (Sierra Caquipec) revealed only one howling
individual in the far distance of the settlement in primary
forests. Density estimates could not be determined here. In
the Sierra Yalijux, A. pigra used mainly primary forest or
old secondary growth at least 15 m in height with at least
two strata (see Renner et al., 2006, for further description
of the vegetation). Thus, 90% (= 1,800 m) of the survey
transects were in primary cloud forest and 10% in old sec-
ondary forest.


Remote assessment using vocalization and triangulation
We applied a survey method in Chelemhi howlers that
is commonly used in bird surveys. While we could de-
termine the howling units of A. pigra by triangulation
and remote assessment, visual contact for group census
and composition is essential since usually only one or
two adult monkeys participate in howling while the re-
mainder of the group stays silent. Thus a combination of
both methods is needed for a detailed census of howlers.
However, the method could be useful to estimate gross

Table 1. Groups of Alouattapigra observed in the Sierra Yalijux near Chelemhi in 2002.
Sub-Adult Sub-Adult
Group Total Adult Male Adult Female Male Female Sub-Adult Infant Area (ha)
Male Female
A 9 1 1 1 1.05
B 8 2 2 3 4.87
C 9 2 1 1 1 4.01
D 5.53
E 8 1 2 1 1.02
F 6 1 1 1.87
G 10 2 2 3 1 2.87

Neotropical Primates 14(1), January 2007

Figure 2. Records of the black howler monkey (Alouattapigra) in the Sierra Yalijux, near the settlement of Chelemhi in 2002. For observa-
tions on the groups (A to G) see Table 1. Probabilities are derived from the Kernel Home Range; Minimum Convex Polygons are derived
from the outermost border of observations per group. Groups were separated according to simultaneously observed howling of groups.

densities of howlers in unstudied areas in shorter time
periods by vocalization records alone. The distinction
between A. pigra groups in the Sierra Yalijux was gener-
ally reliable and clearly separable due to the observations
and territorial howling of the group members, however
we cannot exclude the possibility that in some instances
solitary males were howling.

Howler density and territorial overlap
Howling has been observed to have a territorial function
in A. seniculus and A. pigra (Sekulic, 1982; Estrada et al.,
2004) although this may not always be the case (Estrada et
al., 2002b, 2004). Some authors have argued that howling
is more correctly a mechanism only for intergroupp spac-
ing" (Kitchen et al., 2004), and Sekulic (1982b), studying
Alouatta seniculus, reported on "floating territories".

There is evidence that A. pigra troops occupy their own
exclusive range (Reid, 1997) and that they defend this
range (Horwich 1983a). However, for A. pigra (Ostro
et al., 2001; Pavelka et al., 2003), A. seniculus (Crockett
and Janson, 2000), and other mammals (Pen and Weiss-
ing, 2000), home range overlap has been reported. If this
were true in Chelemha by this survey method, territory
size might decrease and the separation of groups might not
be apparent. A. pigra populations in Belize always showed
some range overlap (Horwich 1983a, b) especially as den-
sities increased (Horwich, pers. obs.). Horwich (1983a)
reported a female crossing into another territory to breed
with the male away from her troop, who later threatened
the same male in an inter-troop territorial dispute. Thus,
there appear to be clear boundaries where adjacent troops
howl at each other across the territory line and distinctions
are possible only by a combination of audio and visual ob-

I ] Mimimum convex polygon E Primary forest
': Milpa, settlements, sec. vegetation M Old secondary forest

Neotropical Primates 14(1), January 2007

servations. Audible and remote detection as well as distin-
guishing the troops by howling alone, as suggested here as
an alternative, might be more difficult, since in some cases
two troops are howling at one location and are not recog-
nizable as two distinguished troops without a visual double
check. Our data for groups E and F (Fig. 2) illustrate this
since no simultaneous or interactive howling was observed.
Thus, they could be merged into one group. However,
since troops over 10 individuals are rare in A. pigra, these
are probably distinct troops.

Home ranges in this study are larger than from other study
sites (see Estrada et al., 2004), and the population density
in this study is intermediate compared to other sites. For
example, lower densities are reported for A. pigra in vari-
ous sites in Mexico (Gonzales-Kirchner, 1998; Estrada et
al., 2002a; Estrada et al., 2004), including Yaxchilan (12.8/
km2), Quintana Roo (15.1/km2), Calakmul (15.2/km2),
and Palenque (23.0/km2). Black howlers in Tikal (El Peten,
Guatemala) also show low population density (17.8/km2).
However, higher density estimates as in Chelemhi have
been reported in Belize, with up to 178 individuals per km2
(Horwich et al. 2001). The population was considered to
be crowded due to fragmentation (Silver et al., 1998; Ostro
etal. 1999, 2000; Horwich et al., 2001). The Sierra Yalijux
has a clearly higher population density than the Mexican
and northern Guatemalan sides but is still below numbers
from more fragmented landscapes of Belize.

Crowding populations?
As observed by Ostro et al. (2001), low density popula-
tions consist of one male with two females and high density
populations of multi-males with > 2 females. The groups
we observed in Chelemhi consist of more than one male
and several females (with some exceptions, Table 1), and
therefore are most likely high-density groups. The com-
paratively high population density ofA. pigra in Chelemhi
implies crowding in the remaining cloud forest. However,
distribution in Chelemhi is patchy. Several groups in ad-
dition to the seven groups as shown in Figure 2 are present
in the area. While the seven groups in Chelemhi seem to
clump together, there is no indication of other individuals
between the widely spaced groups. Therefore we conclude
that the populations have a patchy distribution, probably
due to patchy distribution of nutrition in the Chelemhi
mature forests. However it remains open whether or not A.
pigra crowds in response to the decreased habitat (Markus-
sen and Renner, 2005). Other research teams report crowd-
ed populations from fragmented forest patches in Belize
(Silver et al., 1998; Chapman and Balcom, 1998; Ostro et
al. 1999, 2000; Horwich et al., 2001) with approximately
twice as high population densities than we found in the
Sierra Yalijux (compare above).

Fragmentation and deforestation in the Sierra Yalijux
Fragmentation and deforestation influences behavior and
distribution of A. pigra (e.g., Silver et al., 1998; Estrada et
al., 2002b). Forests are the major habitat for A. pigra as for

most of the other howler species (Estrada et al., 2002b),
even when disturbed (Lyon and Horwich, 1996). No indi-
vidual of any group in the Sierra Yalijux has been observed
in any vegetation used by humans, except for tall secondary
forest (Fig. 2). Only once during our study an individual
of A. pigra was reported in a corn field and shot by a local
farmer (A. Schumacher, pers. comm.). This individual was
likely to have been hunted in mature forest, where the
hunter feigned a case of food competition, as only rarely are
the howlers consumed by locals following poor harvesting
seasons (D. Unger, pers. comm.). Normally, consumption
of howler meat is despised by the local community. Secre-
tive behavior in Chicacnab might be one howler response
to hunting (K. Eisermann, pers. comm.). However, A. pigra
can utilize almost any kind of habitat and will come to the
ground to cross narrow forest gaps and will feed in scrub
areas or in areas low to the ground (Horwich, pers. obs.).


The study was performed in accordance with the current
laws of Guatemala and CONAP (Comisi6n Nacional de
Areas Protegidas) authorized the study (No. 139-2001). We
would like to thank Danielle Shanahan, PiaTerranova, and
Anthony Rylands for help on English and valuable com-
ments on earlier versions of the manuscript. In addition,
our special thanks are going to Vera and Markus Reinhard,
Armin Schumacher, and all local people of the Comuni-
dad de Chelemhi for their great support during the study
period in the field and accommodation, as well as to David
Unger, Knut Eisermann, and Claudia Avendano.


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Alta Verapaz, Guatemala. Ecotropica 12: 43 -49.
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and Rylands, A. B. 1996a. Plan de Accidn para los Pri-
mates Mesoamericanos. IUCN/SSC Primate Specialist
Group, Xalapa, Veracruz, Mexico.
Rodriguez-Luna, E., Cortes-Ortiz, L., Mittermeier, R. A.,
Rylands, A. B., Wong Reyes, G., Carrillo, E., Matam-
oros, Y., Nufez, E and Motta Gill, J. 1996b. Hacia un
plan de acci6n para los primates Mesoamericanos. Neo-
trop. Primates 4(Suppl.): 9-23.
Sekulic, R. 1982a. The function of howling in red howler
monkeys (Alouatta seniculus). Behavior 81: 38-54.
Sekulic, R. 1982b. Daily and seasonal patterns of roaring
and spacing in four red howler Alouatta seniculus troops.
Folia Primatol. 39: 22-48.
Silver, S. C., Ostro, L. E. T., Yeager, C. P. and Horwich, R.
H. 1998. Feeding ecology of the black howler monkey
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Universitdit Gattingen, Gattingen.

Neotropical Primates 14(1), January 2007


Briseida Dego de Resende
Dilmar A. G. Oliveira
Eduardo D. Ramos da Silva
Eduardo B. Ottoni


Primates emit different kinds of vocalizations in different
contexts (Struhsaker, 1967; Snowdon and Pola, 1978; Sey-
farth et al., 1980; Robinson, 1982; Boinski et al., 1999;
Oliveira and Ades, 1998; Maccowan et al., 2001; Di Bitet-
ti, 2001, 2003). For example, long calls can serve as local-
ization cues for conspecifics and are often produced in the
context of territorial encounters, mate attraction, and isola-
tion/group cohesion (Waser, 1982; Miller and Ghanzanfar,
2002). Vocalizations used in close-range social interactions
may be given in many different situations, such as rest-
ing, grooming, foraging or playing (Seyfarth, 1988). Some
primates have different alarm calls for different predators
(Struhsaker, 1967; Fichtel and Hammerschmidt, 2002;
Fichtel and Kappeler, 2002). An often-cited example is the
alarm repertoire of vervet monkeys. In this system, receiv-
ers respond differently to different calls: for example, they
look up and move down or into cover just after an eagle
alarm call, and they run into the trees just after a leopard
call (Struhsaker, 1967; Seyfarth etal., 1980). The appropri-
ate response contingent upon danger increases chances for
survival and reproduction, thus, improvement in fitness.
Vervet alarm call specificity improves with age: at first, in-
fants do not show much discrimination among predator
classes (they may give eagle calls for non-raptors, for ex-
ample); then, they give alarm calls for raptors that are not
their predators; and finally they learn to vocalize only for
the correct raptor predator (Seyfarth and Cheney, 1986;
Seyfarth, 1988). Nevertheless, it is difficult to separate the
role of genetics and environment affecting development
(Seyfarth and Cheney, 1986; Seyfarth, 1988).

Tufted capuchins (Cebus apella) were studied by Robinson
(1982) and, more recently, by Boinski et al. (1999) and
Di Bitetti (2001, 2003). Di Bitetti (pers. comm.), studying
wild Cebus nigritus from Iguazu, Argentina, recorded what
he called the "wah wah wah": a vocalization produced by
only the adult or subadult males, usually in response to
a sudden loud and low frequency sound. The "wah wah
wah" vocalization has a duration of 2-3 sec and consists of
a repetitive series of 16-22 broad band sounds which con-
tain some tonal components. Di Bitetti reported that males
would usually stop doing the activity they were engaged in,
and look alert and attentive while vocalizing in response to

a loud, explosive sound. The two most common sounds Di
Bitetti reported as eliciting this vocalization were thunder
and falling branches or falling trees.

Preliminary observations
In our studies of both captive and wild capuchins, we also
noted the "wah wah" or "rumble call" vocalization. In May
and June 1998, BDR worked with a group of four cap-
tive capuchins (two adult males, two adult females) from
Quinzinho de Barros Zoo, Sorocaba, Sao Paulo, Brazil, and
she noticed that they emitted a particular vocalization just
after explosions coming from a nearby quarry, and that the
monkeys approached their cage mates after the explosions.
Later that year, in August and September, she also noticed
that four captive monkeys from Catanduva's Grove, Cat-
anduva, Sao Paulo, Brazil, emitted what seemed to be the
same vocalization just after firecracker explosions.

After we observed these vocalizations in various popula-
tions, we decided to study them in more detail, including
the acoustic properties, contexts of emission and associated
behaviors, to test our hypothesis that loud and sudden ex-
plosive noises are the proximate causes of this vocalization.
In this paper, we report all the occurrences of the "rumble
call" given by semifree-ranging capuchins of Tiete Ecologi-
cal Park (TEP), and wild capuchins from Carlos Botelho
State Park (CBSP) and Jaragua State Park (JSP), all in the
state of Sao Paulo, Brazil. For a description of the parks and
the capuchin groups, see: TEP-Ottoni and Mannu, (2001);
CBSP-Izar (2004); and JSP-Izar et al. (in prep.). We also
describe a field experiment with a group of approximately
50 free-ranging monkeys in a 7 ha forest fragment in Flo-
rinea, Sao Paulo, to test the ability to elicit the "rumble
call" vocalization by producing loud sudden noises. This
fragment is surrounded by a sugar cane plantation and has
a road running through it. We performed the experiment
at this field site because staff from the other parks did not
permit shooting fireworks, and because observations of the
monkeys were easier in the forest fragment. We did not
recognize individuals in this group.


Field observations
Monkeys from TEP were followed from March 2000 to April
2004 by BDR and Michele Verderane, for a total of 3500
hours of observation. Monkeys from CBSP were followed
from November 2001 to December 2002 by Patricia Izar, for
a total of 1032 hours of observation, and monkeys from JSP
were followed from January 2004 to July 2004 by EDRS, for
a total of 485 hours of observation. All occurrences of these
calls were collected in each site, and, whenever possible, we
registered the precipitating sound (e.g.: thunder, firecracker,
explosion), which individual emitted the call, and other be-
haviors that coincided with the vocalization. One limitation
to the study was that there was only one researcher working
at each site at a given time, and a single individual is unable
to monitor all the group members simultaneously.

Neotropical Primates 14(1), January 2007

Field experiment
In order to generate more controlled observations of the
behaviors and contexts involved in the rumble calls, we
performed the following experiment with a group of free-
ranging monkeys in Florinea. A total of eight firecrackers
were shot into the air, with a minimum interval of 30 min
between each shot. Two shots were executed on 20 June
2004, at 17:00 and 17:30; and six on 21 June 2004, from
07:00 to 11:00. To record the vocalizations, we used a
Sennheiser ME-67TM microphone and a Sony TCD-D8TM
DAT recorder. Recordings started 5 min before each shot
and ceased 5 min after it. Whenever possible, the number
of monkeys visible during the shots was registered, as well
as their age class and the behaviors they displayed imme-
diately after the explosions. However, it was not possible
to register how many monkeys vocalized each time, or
where all of them looked, or the behavior of every visible

We converted the audio recordings into wave files in an
AMD Athlon XPTM computer with a Philips PSC-706TM
soundcard and analyzed them with the Avisoft SASLab
ProTM software. We generated sonograms with a 22 kHz
sampling rate and measured time and frequency param-
eters of calls.


Field observations
Table 1 summarizes the frequency of rumble call events in
capuchin monkey groups from three parks in Sao Paulo.
In most observations, multiple individuals gave the rumble
call and some group members were observed looking at each
other, or running towards each other, giving the impression
that they were looking for social contact. Capuchins never
emitted this vocalization when there was no precipitating
loud sudden noise. However, on some occasions when we
were traveling with the monkeys, we heard thunder or
other explosions but the monkeys did not respond with
rumble calls. In these cases, the explosions tended to be
quieter and more distant.

Tablel. Frequencies of contexts of capuchin monkey rumble call
events in three parks in Sao Paulo, Brazil. TEP = Tiet6 Ecological
Park; JSP = Jaragui State Park; CBSP = Carlos Botelho State Park.
*Possible causes for rumble calls classified as unidentified in JSP
include trucks passing by and objects falling to the ground. Each
event that resulted in rumble calls is counted as one observation.
Each observation may include vocalizations by several monkeys.
Thunder 12 3 7
Firecrackers 23 3 1
Quarry Explosions 0 9 0
Low-Flying Plane 0 0 1
Unidentified 3 19* 0
Total 38 34 9

Field experiment
In all of the eight cases in which we experimentally fired
shots, the capuchin monkeys emitted rumble calls im-
mediately after the shots. Capuchins in the sugar cane
plantation or in the road ran towards the forest imme-
diately after the shots and vocalized only after they had
entered the forest. Both adults (males and females) and
juveniles emitted the vocalization. It was not possible to
determine if infants vocalized. Rumble calls (Fig. 1) occur
as series of harsh pulses, with each pulse lasting about
100 to 120 ms. They reach 7-10 kHz, but the energy is
concentrated between 0,9- 3 kHz. They form quick trails
of pulses (up to 15/second), with emissions lasting from
10 to 20 seconds after each firecracker was shot. The most
intense calls took place immediately after the firecrackers
exploded, with the vocalizations dampened by the noise
of the firecracker. This fact, and the occurrence of simul-
taneous emissions by several individuals, complicates
sound analysis. On 21 June 2004, while we were waiting
30 minutes between a shot and the next one, a vehicle
with a damaged exhaust pipe passed along the road and
emitted explosive noises that also elicited the monkeys'
rumble calls.


The rumble call is contingent on thunder, skyrockets,
explosions or other explosive noises. Both juveniles and
adults make the rumble vocalization. At the moment, we
cannot determine the role of learning in the development
of the rumble call, but it probably has a strong innate
component, as the same vocalization was heard in differ-
ent and distant populations, always contingent upon the
same type of external stimulus (Argentina: Di Bitetti; Su-
riname: Boinski; Northeast and Southeast of Brazil: Izar,
Resende, Verderane and Ramos da Silva). For example,
Patricia Izar reported that similar rumble calls were emit-
ted on two occasions immediately after thunder by mem-
bers of a group of Cebus libidinosus from Gilbu5s, Piaui,
Brazil in the course of 42 hours of contact time (Patricia
Izar, pers. comm.).

Apparently, the sound is a stimulus that elicits the mon-
key's vocalizations. The signaler and the receivers fled
from unprotected sites. If it is an adaptive behavior, with
an innate component, we wonder what kind of fitness
benefit it could provide. As the calls were mainly emitted
in response to explosive noises, and as we have registered
that monkeys ran to the trees just after listening them,
we could hypothesize this vocalization is a sort of alarm
call, impelling the monkeys to protect themselves, pos-
sibly from a thunderstorm, or from a tree falling. How-
ever, according to Di Bitetti (pers. comm.), the acoustic
structure of the rumble call is not ideal for long-distance
communication, and it does not seem to have any acous-
tic similarity to other spacing calls, which seem to be
related to each other. As Seyfarth (1988) states, there is
a direct relation between the function of a call and its

Neotropical Primates 14(1), January 2007

Figure 1. Sonograms depicting the noise stimuli and elicited vocalizations. A) An experimental session with skyrocket rumbles visible as
most intense (darker) and longer sounds at approximately 0.3, 2.5 and 3.1 s. Rumble calls are more clearly visible after rumbles, as trails
of pulses. B) An amplified section of the first sonogram, showing six calling pulses.

acoustic properties: a call that cannot be heard by subjects
far from the signaler cannot be considered an alarm call.
For Di Bitetti, the rumble call is a vocalization, produced
mostly by adult or sub-adult males, that serves to mediate
social relationships among them, and has no relationship
with the cohesion-spacing vocal system or the alarm call
system. In 16% of his records, this vocalization was pro-
duced in social contexts, mostly during reunion displays,
without any previous explosive sound. That is why he be-
lieves it may function as an appeasement call; this could
explain its occurrence during tense situations. The fact
that we also scored female vocalizations indicates that,
even if it is related to social mediation, this behavior is
not exclusively male.

In conclusion, we know that loud explosive noises elicit
this vocalization, and that it is similar across populations
from different and distant parts of Sao Paulo State. Ju-
veniles, males and females give rumble calls. As far as we
know, non-tufted Cebus do not exhibit any rumble calls in
response to explosive noises. The biological function of the
rumble call and its ontogeny are interesting puzzles that
demand more elaborate experimental designs. These exper-
iments should focus on individual observations of subjects
from different age and sex classes, scoring their vocal be-
havior, and the response exhibited just after explosions.


This paper benefited greatly from discussions with and
comments from Dr. Mario Di Bitetti and Dr. Patricia Izar,
who also kindly shared their data with us. We also thank
Michele Verderane for sharing her data from PET, Dr.
Carlos Alberts for making Florinea's experiment possible,
and the staff from Tiete Ecological Park and Jaragua Park
for their support. This work was funded by a grant from the
FAPESP to BDR (#99/11573-2) and CNPq grants to DO
(#305.372/2002-5), ERS (#119939/2004-5), and EBO

Briseida D6go de Resende, Universidade de Sao Paulo
(USP), e-mail: , Dilmar A. G. Oliveira,
CEPESBI / Universidade Regional de Blumenau (FURB),
Eduardo D. Ramos da Silva and Eduardo B. Ottoni, Uni-
versidade de Sao Paulo (USP).


Boinski, S., Gross, T. S. and Davis, J. K. 1999. Terrestrial
predator alarm vocalizations are a valid monitor of stress
in captive brown capuchins (Cebus apella). Zoo Biology
18: 295-312.

Neotropical Primates 14(1), January 2007

Di Bitetti, M. 2001. Food-associated calls in tufted capu-
chin monkeys (Cebus apella). Doctoral thesis, State
University of New York at Stony Brook, New York.
Di Bitetti, M. 2003. Food-associated calls of tufted capu-
chin monkeys (Cebus apella nigritus) are functionally ref-
erential signals. Behaviour 140: 565-592.
Fichtel, C. and Hammerschmidt, K. 2002. Responses of
red-fronted lemurs to experimentally modified alarm
calls: Evidence for urgency-based changes in call struc-
ture. Ethology 108: 763-777.
Fichtel, C. and Kappeler, P. M. 2002. Anti-predator behav-
ior of group-living Malagasy primates: Mixed evidence
for a referential alarm call system. Behav. Ecol. Sociobiol.
51: 262-275.
Izar, P. 2004. Female social relationships of Cebus apella
nigritus in a southeastern Atlantic forest: An analysis
through ecological models of primate social evolution.
Behaviour 141: 71-99.
Maccowan, B., Franceschini, N. V. and Vicino, G. 2001.
Age differences and developmental trends in alarm peep
responses by squirrel monkeys (Saimiri sciureus). Am. J.
Primatol. 53: 19-31.
Miller, C. T. and Ghazanfar, A. A. 2002. Meaningful
acoustic units in nonhuman primate vocal behavior. In:
The Cognitive Animal, C. Allen, M. Bekoff and G. M.
Burghard (eds.), pp.265-273. The MIT Press, Cam-
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grooming interactions as indicators of the social organi-
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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.
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tion to language. In: Primate Societies, B. B. Smuts, D.
L. Cheney, R. M. Seyfarth, R. W. Wrangham and T. T.
Struhsaker (eds.), pp. 440-451. The University of Chi-
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T. Snowdon, C. H. Brown and M. R. Petersen (eds.),
pp.117-143. Cambridge University Press, New York.


Marcos de Souza Fialho
Eleonore Z. E Setz


Like most other howler monkeys, brown howlers (Alouatta
guariba) form one-male groups with up to 10 individu-
als. Even if there is more than one adult male, the alpha
male howler monkey usually monopolizes all reproductive
females and sires all young (Pope, 1990). However, extra-
group copulations (EGCs) have been observed in Alouatta
pigra (Horwich, 1983) andA. seniculus (Agoramoorthy and
Hsu, 2000). Here we report the first EGCs observed in A.
guariba clamitans.


We studied brown howler troops in hillside forest in Porto
Alegre (3012'S, 51o04'W), Brazil, during the summer
(Nov 1998 Jan 1999, 483 obs. hours) and winter (Jun
-Aug 1999, 386 obs. hours; Fialho and Setz, 2000). Study
group GA was comprised of three adult males, three adult
females, and four immatures. An adult male had emigrated
from this group in October 1999 (MMA Jardim, pers.
comm.). A neighboring group (GB) had five individuals.
The GB alpha male was larger and had a more intense red-
dish coloration than any GA adult male.


Daily inter-group encounters between the study groups
were accompanied by extended vocalizations, but they were
usually peaceful. However, an aggressive encounter between
GA and GB occurred on June 12. During this encounter,
the GA group chased and bit individuals from GB, and one
GB individual fled to the ground. Only the GB alpha male
was not attacked. Shortly after this aggressive encounter,
the GB alpha male copulated with a GA female, just a few
meters away from other GA group members. The observing
males of GA group did not react. On June 13, the same two
individuals performed two more EGCs. In the morning,
the male inspected the female's genitalia twice and copulat-
ed with her; an hour and a half later, the large GB male was
feeding in a Ficus tree where GA group was resting. The GB
male approached their group more closely, and GA group
members became agitated. The GB alpha male vocalized
within a few meters of the group, and the female left her
group and followed him for about 50 meters. The female
produced nasal sounds ("Hummm, hummm"), while flick-

Neotropical Primates 14(1), January 2007

ing her tongue rhythmically in and out of her mouth. The
male approached and mounted her.

On the days that the EGCs were observed (June 12 and
13) no within-group copulations were observed for the GA
group. We did not follow the group on June 14 and 15.
Early on June 16, the GB alpha male visited GA's home
range again; he approached, vocalized and left, followed by
the same female as above. Simultaneously, another female
disappeared from GA, but minutes later the two females
reappeared in the group. Later on the same day we saw
three copulations involving a resident GA male and the
female that had copulated with the GB male. This was the
first within-group sexual activity we had observed in GA.
A group GA female carrying an infant observed the copula-
tions but did not react. In summary, all EGC copulations
occurred in the mornings, and each one lasted a minute
or less. About five minutes after each EGC, the GB male
chased the female for several meters. All EGCs involved the
same pair and occurred at the periphery of GA's area, where
home ranges overlapped. The GB male did not follow the
GA group when it moved away from the edge of its range
after these encounters.


Extragroup copulations have been described both in Old
(Smuts, 1987) and New World monkeys (Digby, 1999),
and in monogamous (Mason, 1966; Palombit, 1994;
Reichard, 1995) as well as polygynous species, including
Alouatta spp. (Horwich, 1983; Agoramoorthy and Hsu,
2000). The behavioral repertoire of A. pigra during EGCs
(Horwich, 1983) is more diverse than in A. guariba. How-
ever, the male vocalization in our study has not been de-
scribed for either A. pigra or A. seniculus. In general, EGCs
are similar across Alouatta species. For example, after male
solicitation, the female moves towards the male (Horwich,
1983), and rhythmic tongue flicks precede copulations
(Horwich, 1983; Mendes, 1989; Agoramoorthy and Hsu,
2000). Genital inspection was observed in both A. senicu-
lus (Agoramoorthy and Hsu, 2000) and in A. guariba (this
study). Extra-group copulations last about one minute
across Alouatta species (Horwich, 1983; Agoramoorthy and
Hsu, 2000; this study). After copulation the male chases
the female (Horwich, 1983) and no agonistic behaviors are
directed at the female by her group mates (Horwich, 1983;
Agoramoorthy and Hsu, 2000). In 44% of observed EGCs
in A. seniculus, one or more resident males had visual con-
tact with the mating pair but did not react (Agoramoorthy
and Hsu, 2000); group members also appeared indifferent
to EGCs in this study.

EGCs may be rare in Alouatta compared to other primates.
In the common marmoset, C .-( jacchus, for example,
EGCs occur frequently during group encounters, and males
act aggressively and chase females after copulations (Digby,
1999). Among primates, females are responsible for most
copulation solicitations (Smuts, 1987). In all A. seniculus

EGCs, it was the female who took the lead (Agoramoor-
thy and Hsu, 2000). In A. guariba females also initiated
EGCs. The few data available suggest that females are more
prone to EGCs in multi-male groups (Horwich, 1983; Ag-
oramoorthy and Hsu, 2000; Kowalewski et al., 2006).

Observations of EGCs in Alouatta are consistent with two
hypotheses proposed by Smuts (1987) to explain female
mate choice: (a) the search for genetically superior males
and (b) the preference for non-familiar males (see also
Agoramoorthy and Rudran, 1993). The first hypothesis
is supported by observations on A. guariba (this study)
and A. seniculus (Agoramoorthy and Hsu, 2000) in which
males involved in EGCs were noticeably larger than those
belonging to the female's group. Agoramoorthy and Hsu
(2000) suggested that by copulating with neighboring
males a female could reduce the likelihood of infanticide if
her group was taken over by a new male. Six out of seven A.
seniculus females involved in EGCs had previously lost in-
fants through infanticide (Agoramoorthy and Hsu, 2000).
EGCs could also be a prelude to female dispersal to the
neighboring group. However, in our group, this had not oc-
curred by January 2000, when both GA females gave birth
in GA (MMA Jardim, pers. comm.). The small number of
EGCs observed in howler monkeys does not yet allow us to
falsify any of these competing hypotheses.


We thank the Urban Monkey Project (Universidade Feder-
al do Rio Grande do Sul and Secretaria Municipal do Meio
Ambiente de Porto Alegre), FAPESP grant 98/03018-6,
and Jdlio C. Bicca-Marques and Woody Benson for com-
ments on the manuscript. Denise Brutto permitted us to
do research in her area.

Marcos de S. Fialho, Universidade Estadual de Campinas,
Institute de Biologia, Departamento de Zoologia, Bairro
Barao Geraldo, Campinas, Sao Paulo, Brazil, 13083-970;
e-mail: , and Eleonore Z. E
Setz, Department of Zoology, Institute of Biology, Univer-
sidade Estadual de Campinas, Campinas, Sao Paulo, Brazil,
e-mail: .


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Carlos Henrique de Freitas Burity'
Leandro Duarte da Cruz'2
Vera Lfcia Rocha+
Nelson Barroso da Conceifao2
Daniel Eduardo da Luz'
Durval da Silva Santos'
Devylson da Costa Campos'
Alcides Pissinatti3


The golden lion tamarin, Leontopithecus rosalia (Linnaeus,
1766), is an endangered species (IUCN, 2004) accord-
ing to the World Conservation Union, Species Survival
Commission. The first geographical study of this species,
by Wied-Neuwied (1826), described L. rosalia as distrib-
uted along the coast of the state of Rio de Janeiro between
22and 23S, from the Sao Tomd Cape to the municipal-
ity of Mangaratiba. In 1969, Coimbra-Filho hypothesized
that the historical distribution of this lion tamarin species
extended across the length of the coast of the state of Rio de
Janeiro in lowland forests and at low altitudes usually not
exceeding 300 m a.s.1. (Coimbra-Filho, 1969; Kleiman and
Rylands, 2002). According to Coimbra-Filho, the historical
distribution of L. rosalia comprised several municipalities
of the Fluminense lowlands, including Duque de Caxias.
Based on population counts performed between 1962 and
1969, Coimbra-Filho reported that L. rosalia was extinct in

17 municipalities, including Duque de Caxias (Coimbra-
Filho, 1969; Kleiman and Rylands, 2002).

In the 1990s, censuses across the range of L. rosalia by Ki-
erulff (1993) and later, Kierulff and Proc6pio de Oliveira
(1996) found L. rosalia in only four of the municipalities
described by Coimbra-Filho: Silva Jardim, Casimiro de
Abreu, Cabo Frio, and Saquarema. More recently, L. ro-
salia was found in Araruama in some mountainous areas
of Maca6 de Cima (Rylands et al., 1993), but this recent
expansion of their range clearly resulted from human in-
terference and is not indicative of past distribution. In
the most recent census of L. rosalia distribution (Kierulff
and Rylands, 2003), the authors reported a population of
562 individuals distributed in groups of three to six and
restricted to the aforementioned four municipalities. They
also reported reintroduced populations throughout the
length of the BR-101 road between the municipalities of
Rio Bonito and Casimiro de Abreu in Rio de Janeiro state.
Here, we report the occurrence of golden lion tamarins in
the Taquara Municipal Natural Park, a conservation unit of
the municipality of Duque de Caxias (RJ), where the spe-
cies was considered extinct during the most recent census
(Kierulff and Rylands, 2003).

Materials and Methods

In August 2006, golden lion tamarins were observed in the
Taquara Municipal Natural Park (22035' S, 43o14' W, ap-
proximately 76 m a.s.1.), municipality of Duque de Caixas,
Rio de Janeiro. The 190 km2 park was created according to
Law 1157 (November 11, 1992), and its northern limit
is the Taquara River, near the Ndcleo Colonial of Duque
de Caxias District Three (Fig. 1). The lion tamarins were
observed by the authors during visits to the park guided
by biologists.

Results and Discussion

Based on reports of the presence of golden lion tamarins
near the conservation unit, we interviewed local inhabitants
and showed them pictures to identify the species they had
observed. When golden lion tamarins were confirmed as
the species sighted, we began periodic morning surveys by
walking existing trails close to the areas where the animals
had been spotted. In the first encounter with lion tama-
rins, we observed a non-habituated group of approximately
12 animals that fled towards the Environmental Protec-
tion Area of Petr6polis (APA Petr6polis), a conservation
unit contiguous with the Taquara Municipal Natural Park.
Subsequent sightings of the same group were recorded at
an altitude of approximately 76 m a.s.l. Occasionally, the
group was observed foraging in sympatry with groups of
C .-.' jacchus, C .-'. penicillata and, possibly, hy-
brids of these two introduced marmoset species.

Increased control of access into Taquara Municipal Nat-
ural Park will allow L. rosalia to safely utilize the forest

Neotropical Primates 14(1), January 2007

Figure 1. (A) State of Rio de Janeiro (RJ), with the original distribution of Leontopithecus rosalia (lines) and fragmented current distribu-
tion (< reintroduced population and =town, according to Rylands et al., 2002). (B) Map indicating the location of the L. rosalia group
seen in the Taquara Municipal Natural Park (star) in the municipality of Duque de Caxias, Rio de Janeiro (RJ), the new southernmost
location for the species.

here, as well as in the higher altitude protected area of
the APA Petr6polis. Research on home range use, activity
budget, and interspecific interactions with other monkey
species by this lion tamarin group (possibly with the aid
of radiotelemetry) at the park, as well as an assessment of
the size of this golden lion tamarin population, its genetic
structure, distribution, and possible threats to its surviv-
al, are urgently needed to evaluate its long-term viabil-
ity. These data will also serve as baseline information for
future programs of population supplementation or species

Carlos Henrique de Freitas Burity, email: unigranrio.com.br>, Leandro Duarte da Cruz, Daniel
Eduardo da Luz, Durval da Silva Santos, Devylson da
Costa Campos, Institute of Biosciences, Biological Scienc-
es Unigranrio, Duque de Caxias, RJ, Vera Ltcia Rocha,
Nelson Barroso da Conceigio, Municipal Environment
Secretariat (SEMMA) Duque de Caxias, RJ, and Alcides
Pissinatti, Rio de Janeiro Primate Center (CPRJ-FEEMA)
- Guapimirim, RJ.


Coimbra-Filho, A. F 1969. Mico-Leao, Leontideus rosa-
lia (Linnaeus, 1766): Situacao atual da esp6cie no Brasil
(Callitrichidae, Primates). Anais Acad. Brasil. Ciencias 41
(Suppl.): 29-52.

IUCN. 2004. Red List of Threatened Species. redlist.org>.
Kierulff, M. C. M. 1993. Status and distribution of the
golden lion tamarin in Rio de Janeiro. Neotrop. Primates
1(4): 23-24.
Kierulff, M. C. M. and Proc6pio de Oliveira, P. 1996. Re-
assessing the status and conservation of the golden lion
tamarin Leontopithecus rosalia in the wild. Dodo, J. '
Preserve. Trust 32: 98-115.
Kierulff, M. C. M. and Rylands, A. B. 2003. Census and
distribution of the golden lion tamarin (Leontopithecus
rosalia). Am. J. Primatol. 59(1): 29-44.
Kleiman, D. G. and Rylands, A. B. 2002. Lion Tamarins:
Biology and Conservation. Smithsonian Institution Press,
Washington, DC.
Rylands, A. B., Coimbra-Filho, A. F and Mittermeier, R.
A. 1993. Systematics, geographic distribution, and some
notes on the conservation status of the Callitrichidae. In:
Marmosets and Tamarins: Systematics, Behaviour and Ecol-
ogy, A. B. Rylands (ed.), pp.11-77. Oxford University
Press, Oxford.
Rylands, A. B., Kierulff, M. C. M. and Pinto, L. P. S. 2002.
Distribution and status of lion tamarins. In: Lion Tama-
rins: Biology and Conservation, D. G. Kleiman and A. B.
Rylands (eds.), pp.42-70. Smithsonian Institution Press,
Washington, DC.
Wied-Neuwied, Prinz Maximilian zu, 1826. Beitrdge zur
Naturgeschichte von Brasilien, vol.2.

Neotropical Primates 14(1), January 2007


Tadeu Artur de Melo Jfnior
Fernando Jose Zara


Army ants have different intra- and extranidal symbiotic
associations with invertebrates and vertebrates (Gottwald,
1995). Vertebrates that feed on arthropods disturbed by
the army ants include the anuran Bufo marinus; lizards such
as Ameiva, Kentropyx, Anolisfrenatus, and Tupinambis meri-
anae; a large number of birds; and callitrichids (Willis and
Oniki, 1978, 1992; Rylands et al., 1989; Martins, 2000;
Melo Jdnior and Zara, pers. observ.). The best studied associ-
ations between vertebrates and army ants in the Neotropical
region are those between birds and ants. More than 50 spe-
cies from the families Cuculidae, Cracidae, Dendrocolapti-
dae, Formicariidae, Thamnophilidae, Rallidae, Tinamidae
and Thraupidae have been observed to follow army ants
and to pick off arthropods flying or sitting on forest litter
that was disturbed by the army ants Eciton burchelli and
Labiduspraedator (Willis and Oniki, 1978, 1992). Accord-
ing to Gottwald (1995), swarm-following birds have to
compete with marmosets for this food resource stirred up
by progressing army ants. However, marmosets only op-
portunistically exploit the arthropods flushed by swarms of
E. bu:ch//c. and L. praedator (Rylands etal., 1989; Martins,
2000). These two army ant species present a similar broad
swarm-raiding pattern (Teles da Silva, 1982; Rylands et al.,
1989; Gottwald, 1995) that may cause higher prey distur-
bance in the forest litter than the columnar raiding pattern
of Eciton hamatum (Teles da Silva, 1982).

In the Neotropics, five species from the genus Calli-
thrix- C. humeralifer, C. kuhli, C. flaviceps, C. ... 0..,
and C. aurita-have been reported to take arthropods
over swarms of army ants (Rylands et al., 1989; Martins,
2000). Marmosets seem to associate more frequently with
E. bu:/he// than with L. praedator (Rylands et al., 1989),
and C aurita was only observed in association with La-
bidus sp. during the dry season (Martins, 2000). Associa-
tion with army ants has not been recorded for C. penicil-
lata and C. jacchus so far, possibly due to a relative scarcity
or absence of army ants in the drier regions of cerrado
(Brazilian savanna) and semi-deciduous scrub and wood-
lands of Brazilian caatinga (Rylands et al., 1989). How-
ever, L. praedator has a wide geographic distribution and
ranges from central Mexico (San Luis Potosi) to northern
Argentina (Borgmeier, 1955; Rettenmeier, 1963); it has
been collected in some regions of the Cerrado (Kempf,
1972; Watkins, 1976). Here we report the association be-
tween C .-.'. penicillata and the army ant L. praedator

on three different occasions at two localities in the Cer-
rado and the Atlantic Forest of Brazil.


One observation was made during a field study in the
Parque Estadual Fernao Dias (4404'W, 1956'S), Minas
Gerais, located in a transitional area between the Atlan-
tic Forest and the Cerrado. This protected area has about
2000 ha and varies in altitude between 850 and 950 m
a.s.1. (Melo Jdnior, 2000). The other two observations were
made at the Parque Estadual da Ilha Anchieta (45001'W,
2332'S), Ubatuba, Sao Paulo. This protected 828-ha
island is located 600 m from the continent. Both observa-
tions on the island were made at different points of the
Saco Grande trail, which crosses a fragment of rainforest.
In March 1983, five C. penicillata were introduced to Ilha
Anchieta by the Fundagao Parque Zool6gico de Sao Paulo
(Guillaumon et al., 1989), and now these marmosets are
the most abundant primate on this island (Galetti, pers.
comm.). Army ant samples were collected and identified
as L. praedator according to Borgmeier (1955, p. 84: major
worker) and Watkins (1976: worker key 8).

Results and Discussion

The first association was observed on 11 October 1999,
in the late dry season. A group of seven black-tufted mar-
mosets, C. penicillata, was seen following the army ant
L. praedator. Total observation time lasted 42 minutes,
but the total time that this group followed the army ants
was longer, because the marmosets were already near the
ground when observations began. On eight different op-
portunities, individual marmosets were seen to take in-
sects on the ground, an uncommon behavior for marmo-
sets. Two other observations of C penicillata following the
army ant L. praedator were made on 23 January 2000 and
7 February 2003, both during the rainy season on Ilha
Anchieta. The first observation lasted for 12 min and oc-
curred at 08:45. Two individuals from a group of six indi-
viduals were seen close to the swarm front, at about 0.5 m
from the ground. The other members sat on branches at
around 3-5 m above ground and emitted alarm calls to-
wards the observers. To avoid interference, we retreated
to a distance of 5 m and made further observations using
binoculars. Marmosets were observed taking arthropods
from the ground. The army ants showed a small swarm
front (0.5 m wide) near the marmosets. Other swarm
fronts (not exploited by marmosets) were observed, but
most of the ants remained in columns. The raiding pat-
tern seemed more columnar, similar to the description
by Fowler (1979). The marmosets stopped foraging over
the army ants due to disturbance from arriving tourists.
The second record started at 09:15 when a group of seven
marmosets was observed capturing flushed arthropods
over the swarm front during 38 min. This time the army
ants were foraging in a swarm raiding pattern, similar
to descriptions by Borgmeier (1955) and Rettenmeier

Neotropical Primates 14(1), January 2007

(1963). This swarm raid was more vigorous than the
first and more than 2 m wide. The marmosets were po-
sitioned at the front of the swarm and over the fan area
on branches at 0.3-1 m above ground. Three different
individuals were observed going down to the ground and
capturing arthropods ahead of the swarm on seven differ-
ent occasions.

Our observations indicate that C penicillata may follow
the army ant L. praedator and use disturbed arthropods
as a food source, similar to what has been described for
C humeralifer, C kuhli, C flaviceps, C. f... ..1, (Rylands
et al., 1989), and C aurita (Martins, 2000). Flushed ar-
thropods captured by the marmosets included crickets,
grasshoppers, cockroaches, and spiders, in line with what
has been described for other C .-'. species (Rylands et
al., 1989; Martins, 2000). Moths, true bugs (Hemiptera),
and beetles that are usually preyed upon by L. praedator
(Borgmeier, 1955; Rettenmeier, 1963; Gottwald, 1995)
were also preyed upon by the marmosets during our ob-
servations. We did not observe the marmosets taking prey
that had been captured by the army ants, as reported for C
humeralifer by Rylands et al. (1989).

The C penicillata individuals that captured insects were
located on branches below 1 m, similar to other C .-'.
species during ant following (Rylands et al., 1989). They
were always in the center of the swarm front or fan area.
This positioning resembles that described for dominant
birds during ant following. According to Willis and Oniki
(1978, 1992), large dominant birds occupy the central and
probably best zone of high prey disturbance in the swarm
front, while medium-sized birds were chased off to more
peripheral zones with fewer flushed arthropods. Accord-
ing to Martins (2000), for C aurita following army ants is
more attractive during the dry months, when availability of
arthropods and other resources is low. Paradoxically, during
the dry and cold months, L. predator performs fewer and
less intense raids (Fowler, 1979). For C penicillata, fol-
lowing behavior seems to be more opportunistic and re-
lated to the presence or absence of the army ants in swarm
raid pattern rather than to season. Rylands et al. (1989)
suggested that this association has not been observed for
C penicillata and C jacchus possibly due to the relative
scarcity or absence of this ant species in their ranges. This
is probably true for C jacchus in the Caatinga, where there
are no records for L. praedator. In the Cerrado, however,
L. praedator is one of the most common army ants col-
lected (Zara, unpubl. data), and original records of these
army ants in the Cerrado were presented in Kempf (1972)
and Watkins (1976).


We are grateful to the directors of Parque Estadual da Ilha
Anchieta (Instituto Florestal SP) and Parque Estadual
Fernao Dias for permits to visit these areas. TAMJ received
grants from FAPESP (97/04642-2) and CNPq. FJZ ex-

tends thanks to FAPESP (proc. 05/04707-5). Our grati-
tude to Cynthia P. A. Prado for smoothing out the English,
and to Simone Batalha Velten and Alexandre de Almeida
for field assistance.

Tadeu Artur de Melo J6nior, UNIFRAN, Universidade de
Franca, Avenida Armando Salles de Oliveira s/n, Franca,
SP, Brazil, CEP 14404-600, e-mail: ,
and Fernando Jose Zara, UNESP Campus do Litoral
Paulista, Unidade de Sao Vicente, Praga Infante Don Hen-
rique s/n Sao Vicente, SP, Brazil, CEP 11330-900, e-mail:


Auricchio, P. 1995. Primatas do Brasil. Terra Brasilis, Sao
Borgmeier, T. 1955. Die Wanderameisen der Neotropisch-
en Region (Hym. Formicidae). Studia Ent. 1: 1-716
Eisenberg, J. F. and Redford, K. H. 1999. Mammals of the
Neotropics, Vol 3: The Central Neotropics: Ecuador, Peru,
Bolivia, Brazil. The Chicago University Press, Chicago.
Fowler, H. G. 1979. Notes on Labidus praedator (Fr. Smith)
in Paraguay (Hymenoptera: Formicidae: Dorylinae: Eci-
tonini). J Nat. Hist. 13: 3-10.
Gottwald Jr., W. H. 1995. The Army Ants: The Biology of
Social Predation. Cornell University Press, London.
Guillaumon, J. R. et al. 1989. Plano de manejo do Parque
Estadual da Ilha Anchieta. Institute Florestal Srie Regis-
tro, Sao Paulo.
Kempf, W. W. 1972. Catilogo abreviado das formigas da
Regiao Neotropical. Studia Ent. 15: 3-344.
Martins, M. M. 2000. Foraging over army ants by C
aurita (Primates: Callitrichidae): Seasonal occurrence?
Rev. Biol. Trop 48: 261-262.
Melo J6nior, T. A. 2000. Morfometria, comportamento
alimentar e social do cuitelao, Jacamaralcyon tridactyla
(Piciformes: Galbulidae) em duas reserves no Estado de
Minas Gerais. Dissertagao de Mestrado, UNESP, Rio
Claro, SP
Rettenmeyer, C. W. 1963. Behavioral studies of army ants.
Univ. Kans. Sci. Bull. 44: 281- 465.
Rylands, A. B., Monteiro da Cruz, M. A. 0. and Ferrari,
S. E 1989. An association between marmosets and army
ants in Brazil.J. Trop. Ecol. 5: 113-116.
Teles da Silva, M. 1982. Behaviour of army ant Eciton
burchelli and E. hamatum (Hymenoptera: Formicidae)
in the Belen region. III. Raid activity. Ins. Soc. 29: 243-
Watkins, J. E 1976. The Identification and Distribution
of New Word Army Ants (Dorylinae: Formicidae). Baylor
University Press, Waco.
Willis, E. 0. and Oniki, Y. 1978. Birds and army ants.
Ann. Rev. Ecol. Syst. 9: 342-263.
Willis, E. 0. and Oniki, Y. 1992. Aves e as formigas de cor-
reigao. Bol. Mus. Paraense E. Goeldi 8: 123-150.

Neotropical Primates 14(1), January 2007


Marcelo Marcelino de Oliveira
Marcos de Souza Fialho

Na Estagao Experimental em Ouro Preto do Oeste, no
estado de Rond6nia, Brasil, terd inicio a implementagao
de models experimentais de cercas para evitar a predagao
de frutos de cacau (Theobroma cacao) por macacos-prego
(Cebus apella). Esta agao 6 resultado de um trabalho de ne-
gociacao do Centro de Protegao de Primatas Brasileiros,
6rgao especializado em conservagao e manejo de primatas
do Institute Brasileiro do Meio Ambiente e dos Recursos
Naturais Renoviveis (CPB-IBAMA), para resolver uma si-
tuacao de conflito entire primatas e agricultores. A lavoura
de cacau 6 uma cultural em expansao na Amaz6nia, fomen-
tada pelo Ministerio da Agricultura atrav&s da Comissao
Executive do Plano da Lavoura Cacaueira (CEPLAC),
tendo como foco pequenos agricultores em areas de assen-
tamento rural. A predagao de frutos de cacau por Cebus
apella comegou a ser constatada pela CEPLAC em sua
Estagao Experimental ha mais de dois anos e registros de
predagao em propriedades rurais ji foram comunicadas a
Superintendencia do IBAMA em Rond6nia (Supes-RO).
0 aumento da predagao nas areas de experiment e de
produgao de cacau na estagao, levou a CEPLAC a solici-
tar a Gerencia Executiva do IBAMA em Ji-Parand (RO)
providencias para retirada dos animals, pedindo inclusive
apoio ao Ministerio Pdblico Federal visando pressionar o
IBAMA nesse sentido. Um estudo elaborado pela Univer-
sidade Federal de Rond6nia (UNIR) em setembro deste
ano, a pedido da CEPLAC, identificou oito esp6cies de
primatas na irea e registrou a ocorrencia de dois grupos
de Cebus apella, alkm de alguns individuos solitarios (Mes-
sias et al., 2006). 0 estudo recomendou a translocaiao
de um desses grupos, constituido aparentemente por at6
20 animals. Em reuniao realizada em Porto Velho (RO)
no dia 13 de novembro de 2006, com a participagao da
CEPLAC, da Supes-RO e da UNIR, os representantes do
CPB conseguiram o convencimento de que a melhor solu-
gao para o conflito era o cercamento das areas experimen-
tais de plantio de cacau, mantendo-se assim a convivencia
com os animals. Em visit a irea em Ouro Preto do Oeste
foi definida a implantagao de dois models de cerca, ao
long de aceiros ji existentes, num total de 4,5 Km de ex-
tensao por 4 metros de largura. 0 primeiro modelo, ji em
uso pela CEPLAC em um pequeno trecho do aceiro, 6 o
de uma cerca eletrificada de arame e tela, com amperagem
adequada para apenas repelir os animals sem causar danos
fisicos, com 1,0 m de altura. 0 segundo modelo, trata-se
de uma cerca de tela com 1,5 metro de altura, encimada
por uma chapa galvanizada de 1,0 m de largura disposta

em angulo de 45, corn a extremidade voltada para a mata.
A CEPLAC testari a aplicabilidade de ambos os models,
verificando o custo de sua instalagao e manutencao e sua
eficiencia na contencao dos macacos-prego, bem como,
de outros potenciais predadores dos frutos. 0 CPB espera
que o modelo com melhor resultado seja, a partir de entao,
incorporado no conjunto das tecnicas e m6todos que a
CEPLAC transfer aos agricultores, para implantagao da
cultural do cacau.

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


Messias, M. R., Ferronato, M. L. e Oliveira, M. A. 2006.
Inventario e estimativa populacional da mastofauna da
EXTEX da CEPLAC Ouro Preto do Oeste Subsidios
para o manejo das esp6cies potencialmente predadoras dos
frutos de cacau. UNIR e CEPLAC. Relat6rio T&enico.


On 13 April 2007, during a preliminary field survey of the
current distribution and conservation status of the yellow-
tailed woolly monkey (Oreonaxflavicauda) in the depart-
ments of San Martin and Amazonas, Peru, we encountered
a group of eight 0. flavicauda consisting of five adults and
three juveniles. The encounter took place in a privately
owned, unprotected forest at an elevation of 1900 m a.s.l.,
1 km NW of the village of Santa Rosa (0540'13.5"S,
7755'08.0"W). This area is highly disturbed primary
montane forest interspersed with pasture. The group was
followed for approximately one hour. During the entire
encounter we observed the presence of a female white-
bellied spider monkey (Ateles belzebuth) freely associating
with all members of the 0. flavicauda group. The female
spider monkey continued to travel with the group as they
entered the next valley, but we were unable to follow. Both
species showed complete tolerance to each other and an
equal intolerance of the presence of humans, with loud
vocalizations and branch shaking. Observation was made
easier by the aggressive approach of the group toward us.
We believe this is the first record of such an association
and highlights the need for further detailed study of both
species in the wild, in particular the critically endangered
yellow-tailed woolly monkey about which we know so


We would like to thank Fernando Guerra-Vasquez, Helene
and Carlos Palomino (IKAMA Peru), and the people of
Santa Rosa and La Esperanza for their help, and the Gobi-

Neotropical Primates 14(1), January 2007

erno Regional de San Martin. Also we would like to thank
Aap, Holland, IPPL-UK and The Monkey Sanctuary Trust,
UK for generously funding this project.

Noga Shanee and Sam Shanee, Neotropical Primate Con-
servation, 2 Even Gvirol, Kfar Saba, 44393, Israel, neoprimate.blogspot.com>, e-mail: com>.

Angela M. Maldonado, Fundaci6n Entropika, Cra
9 No. 14-17, Leticia, Amazonas, Colombia, e-mail:


Carolina Gdmez-Posada

At least seventy percent of the original montane ecosys-
tems have been lost from the Colombian Andes. Forest
remnants are mostly small, isolated, and on privately-held
lands. Animals persisting in this landscape have adapted to
human encroachment and some have managed to survive
in highly degraded habitats. However, this does not guar-
antee the long-term survival and health of populations.
Andean forests in Colombia urgently need management
and conservation programs. We have developed a project
aimed at evaluating how howler populations are respond-
ing to loss and fragmentation of their natural habitats in
the Colombian Andes, with the expectation that this could
lead to measures to prevent further losses. We are studying
howler populations in montane forest in three provinces of
Colombia in the coffee growing region and Cauca Valley
(900 to 2200 m a.s.1.), evaluating the demographic and be-
havioral responses of howlers to habitat fragmentation and
diminished resources. This research has included eleven
theses of undergraduate and graduate students from nine
Colombian universities, addressing the following topics:

* Status of wild populations of red howler monkey in
forest fragments (natural forest, "guadua" bamboo
forest, forestry plantations);
* Ranging patterns, use of food resources and habi-
tat of red howler monkey in isolated Andean forest
* Genetic variability and endogamy of red howler
monkey in isolated Andean forest fragments in the
coffee region in Colombia;
* Use of anthropogenic habitats by monkeys and pro-
ductive systems as alternative tools for conservation in
private lands;
* Conservation strategy of the red howler monkey in the
coffee region in Colombia.

We have interacted with government agencies, large pri-
vate land owners, and small ranch farmers. We consulted
with farmers and the largest forestry plantation com-
pany in Colombia to understand their points of view,
in order to explore different management options that
would allow them to use their land without destroying
the resources needed for monkeys and other wildlife to
survive. These results were used to develop the conser-
vation strategy for this species in the region, and some
of the proposed conservation actions now have been
implemented by land owners and local environmental

This project is funded by national and international in-
stitutions: Corporaci6n Aut6noma Regional del Valle
del Cauca CVC, Fundaci6n para la Promoci6n de la
Investigaci6n y la Tecnologia del Banco de la Repdbli-
ca, Instituto de Investigaciones Biol6gicas Alexander
von Humboldt (Colombia), John D. and Catherine T.
MacArthur Fundation, U. S. Fish and Wildlife Service,
and Idea Wild.

Carolina G6mez-Posada, Wildlife Conservation Society
Colombia Program / Fundaci6n EcoAndina, Cali, Co-
lombia. AA. 25527, e-mail: .


A ministry do Meio Ambiente, Marina Silva, receberi o
maior premio das Nagoes Unidas na drea ambiental, o
"Champions of the Earth" (Campe6es da Terra) de 2007,
como reconhecimento ao seu trabalho em favor da pre-
servagao da floresta amaz6nica e da valorizagao das comu-
nidades locais e tradicionais da regiao. 0 andncio foi feito
nesta quinta-feira (01) pelo Programa das Na6oes Unidas
pelo Meio Ambiente (PNUMA). Marina Silva 6 uma das
sete personalidades que serao premiadas. 0 "Campe6es
da Terra" seri entregue numa cerim6nia prevista para o
dia 19 de abril, em Cingapura. 0 sub-secretirio da Or-
ganizagao das Nagoes Unidas (ONU) e diretor-executivo
do PNUMA, Achim Steiner, em carta enviada a ministry,
elogia a sua dedicacao na defesa das quest6es ambientais.
"Vossa Excelencia reconheceu que a construgao de uma
alianga de apoio para proteger o meio ambiente exige
adesao a um conjunto bisico de valores. Sua crenga de
que o sucesso da luta para salvar a vida sobre a Terra exige
que principios estejam a frente de nossos esforgos fez com
que fosse merecedora do premio Champions of the Earth
2007", escreve ele. 0 premio existe desde 2004. Seu ob-
jetivo 6 contemplar pessoas que tenham uma contribui-
gao significativa e reconhecida, global e regionalmente,
na protegao e gestao sustentivel do meio ambiente e dos
recursos naturais.

Fonte: InforMMA.

Neotropical Primates 14(1), January 2007


O passado 27 de maio, durante a terceira edigao do Viva
a Mata, event promovido pela Fundagao SOS Mata
Atlntica, o Instituto BioAtlntica (IBio) langou o livro
"Meu p6 de Mata Atlntica Experiencias de recompo-
sigao florestal em propriedades particulares no Corredor
Central". Resultado do Programa Conservagao em Terras
Privadas do IBio, o livro traz os primeiros dados do mo-
nitoramento iniciado no ano de 2004 em ireas de flores-
ta restauradas em propriedades localizadas no Corredor
Central da Mata Atlntica. 0 projeto de monitoramen-
to e a publicacao foram feitos com recursos do Fundo de
Parcerias para Ecossistemas Criticos (Critical Ecosystem
Partnership Fund CEPF) e da Agencia Americana para
o Desenvolvimento Internacional (USAID). "Nas viagens
a Bahia e Espirito Santo, no inicio das atividades do Pro-
grama de Conservagao da Biodiversidade em Terras Priva-
das, tivemos contato com proprietirios rurais do Corredor
Central da Mata Atlntica que vem desenvolvendo corn
recursos pr6prios agoes de recomposigao florestal. Em visi-
tas as dreas e nas conversas com esses proprietirios, consta-
tamos a ausencia de inventirios e monitoramento. Ainda
que por si s6 sejam dignas de reconhecimento, 6 preciso
avaliar o papel dessas agoes na redugao dos efeitos da frag-
mentagao do habitat natural," explica o engenheiro flores-
tal Beto Mesquita, coordenador do program Conservagao
em Terras Privadas do IBio e um dos autores do livro. "De
modo geral hi uma lacuna de conhecimento a respeito do
sucesso de projetos de recomposigao florestal. Em nosso
estudo, al6m de aspects floristicos e fision6micos consi-
deramos avifauna como grupo bioindicador para avaliar
o papel destas ireas como corredores de biodiversidade,"
comenta Ludmila Pugliese de Siqueira, gerente de projetos
do IBio e autora do livro.



Thiago Romero

O Museu Paraense Emilio Goeldi (Mpeg) e o Instituto Na-
cional de Pesquisas da Amaz6nia (Inpa) deram um passo
important para o monitoramento e a preservagao da bio-
diversidade na Amaz6nia. As duas instituigoes, vinculadas
ao Ministerio da Ciencia e Tecnologia (MCT), integraram
suas bases de dados cientificos sobre o bioma que ocupa
quase a metade do territ6rio national. 0 novo ambiente
on-line interliga os sistemas computacionais da Rede CT
Petro Amaz6nia e do Programa de Pesquisa em Biodiversi-
dade (PPBio-Amaz6nia), realizados em parceria pelas duas
instituigoes. Todo o material gerado pelos dois projetos esti
disponivel para consult pdblica pela internet. Novas in-
forma6oes poderao ser inseridas em tempo real a partir de

A Rede CT Petro Amaz6nia redne instituigoes de ensino
superior e de pesquisa da Amaz6nia e tem o objetivo de
desenvolver tecnologias voltadas para a recuperacao de areas
degradadas na floresta resultantes da exploracao de recursos
minerals, como o petr61leo e o gis natural. 0 PPBio-Amaz6-
nia desenvolve agoes de pesquisa voltadas para political de
conservagao e uso sustentivel da biodiversidade na regiao,
por meio da manutencao de acervos e colleges biol6gicas e
inventirio de esp6cies vegetais e animals. Mais informagaes:
ou br/ctpetro>.

Fonte: php?id=6557>.


The Chicago Zoological Society is soliciting new proposals
for the Chicago Board of Trade Endangered Species Fund
for the first grant cycle of 2007. The Committee is look-
ing for projects that will be conducted between June 2007
and February 2008. The Fund will support small projects,
usually up to $5,000 (smaller requests will fare better). For
more information contact: Daniel M. Brooks, Ph.D., Cu-
rator of Vertebrate Zoology, at .


Starting next March, Mona Foundation will present cours-
es on primate ethology. The main goal of these courses is
to study and comprehend the behaviour of non-human
primates, not only on a theoretical basis but also on a prac-
tical one. The estimated duration of each course will be of
15 hours (8.5 of theory and 6.5 of practice) along two days
(Fridays and Saturdays) and will take place on the third
weekend of each month. The schedule of the course will
be from 10.00h to 18.30h each day. The application fee
includes a file with a CD-ROM, a field notebook and a
diploma. Next courses taking place on the 1S semester of
2007 (Basic Level), 16th and 17th of March, 20th and 21" of
April, 18' and 19' of May, 15th and 16thof June. For more
information regarding the courses, please contact: Miquel
Llorente at , or visit www.fundacionmona.org/final/castellano/noticies_marco.


El Simposio de Recursos Gen&ticos paraAm6rica Latinay el
Caribe (Sirgealc) es el principal foro bianual americano, en
el cual diversos experts en el tema comparten las experien-
cias derivadas de la instrumentaci6n y el seguimiento de los

Neotropical Primates 14(1), January 2007

proyectos cientificos relacionados con los recursos gen&ticos,
provenientes de las plants, los animals y los microbios. En
este event se analizan ademas los avances en el tenor y se
proponen las estrategias y las tareas prioritarias para la region
latinoamericana. Esta sexta emisi6n del Sirgealc se llevard a
cabo en la Ciudad de Mexico, del 12 al 16 de Noviembre
de 2007 y los temas principles serin: Estrategias de con-
servaci6n de los recursos gen&ticos, Educaci6n en recursos
gen&ticos en todos niveles y Marco regulatorio de acceso a
los recursos gen&ticos. Para mayores informes visitar www.coyoacan-global.com/sirgealc/convocatoria.htm>.


Temos a satisfacao de anunciar o langamento da pigina ele-
tr6nica do Museu de Biologia Prof. Mello Leitao. La voce
poderia encontrar a informacao sobre a hist6ria, as ativida-
des e a noticia relacionadas com o museu. Enderego: melloleitao.iphan.gov.br>.


The Pitheciines include some of the most unusual and in-
triguing primates of the Neotropics. Several are threatened
with extinction, and for many we lack all but the most basic

The Pitheciine Action Group (PAG) exists to promote the
conservation of the genera Callicebus, Cacajao, Chiropotes and
Pithecia. PAG's main objectives are to coordinate and encour-
age research, act as a clearinghouse for information and re-
source sharing, and provide scientific and technical informa-
tion to guide conservation strategies and policy decisions.

Founded informally in 2005, PAG was launched officially
at the Pitheciins: Ecology and Conservation symposium of
the 2006 IPS Congress in Uganda, and is now part of the
IUCN/SSC Primate Specialist Group. PAG has an Executive
Committee of four, a 25-member Coordinating Committee
and Sub-Committees for each genus that includes pitheciine
experts from all range countries. There is also the PAG-Net-
work with an open-list membership; this is the information
exchange facet of the Group, to which institutions, research-
ers and interested individuals are invited to contribute.

PAG will soon launch a dedicated website and an electronic
newsletter. As well as information about current group ac-
tivities, website resources will include lists of priority taxa
and research topics, key literature, theses and dissertations,
photos and videos, recordings of vocalizations and distribu-
tion of museum specimens.

Current PAG activities include a 45-chapter, 82-author
book, Evolutionary Biology and Conservation of Titis, Sakis
and Uacaris, to be published by Cambridge University Press,

and participation, together with the Brazilian Government's
Environmental Protection Agency (IBAMA), in the formu-
lation of Action Plans for endangered Brazilian taxa.

For further information about PAG, please e-mail
or visit www.pitheciineactiongroup.org>.

Liza Veiga, Museu Paraense Emilio Goeldi, Belkm, Brazil,
e-mail: and Adrian
Barnett, University of Surrey Roehampton, UK, e-mail:

PAG Executive Committee Liza Veiga, Adrian Barnett,
Stephen Ferrari (Universidade Federal de Sergipe, Sao
Crist6vao, Brazil, e-mail: )
and Marilyn Norconk, Kent State University, USA, e-mail:

PAG Coordinating Committee Liza Veiga; Adrian Bar-
nett; Stephen Ferrari; Marilyn Norconk; Jdlio Cesar Bicca-
Marques (Pontificia Universidade Cat61lica do Rio Grande
do Sul, Porto Alegre, Brazil, e-mail: );
Jean-Philippe Boubli (The University of Auckland, New
Zealand, e-mail: ); Mark Bowler
(University of Kent, UK, e-mail: hotmail.com>); Thomas R. Defler (National University
of Colombia, Bogota D.C., Colombia, e-mail: asdefler@gmail.com>); Anthony Di Fiore (New York
University, USA, e-mail: );
Maria Cecilia Kierulff (Fundagao Parque Zool6gico de
Sao Paulo, Brazil, e-mail: ); Edu-
ardo Fernindez-Duque (University of Pennsylvania, USA,
e-mail: ); Eckhard Heymann
(Deutsches Primatenzentrum, Goettingen, Germany, e-
mail: ); Shawn Lehman (University
of Toronto, Canada, e-mail: ca>); Laura Marsh (Global Conservation Institute, Santa
Fe, USA, e-mail: );
Marcelo Oliveira (IBAMA Centro de Protegao de Pri-
matas Brasileiros, Joao Pessoa, Brazil, e-mail: Oliveira@ibama.gov.br>); Erwin Palacios (Conservation
International Colombia, Bogoti D.C., Colombia, e-mail:
); Liliam Pinto (Universidade
Estadual de Campinas, Sao Paulo, Brazil, e-mail: uol.com.br>); Leila M. Porter (Northern Illinois Univer-
sity, USA, e-mail: ); Helder
Queiroz (Mamiraud Escrit6rio Belkm, Universidade Fed-
eral do Pari, Brazil, e-mail: );
Jos6 Silva Jdnior (Museu Paraense Emilio Goeldi, Belkm,
Brazil, e-mail: ); Bernardo
Urbani (University of Illinois at Urbana-Champaign, USA,
e-mail: ); Rob Wallace (Wild-
life Conservation Society-Bolivia, La Paz, Bolivia, e-mail:
); Kirsten Pullen (Paignton Zoo En-
vironmental Park, Devon, U.K., e-mail: paigntonzoo.org.uk>); Rolando Aquino (Estaci6n Experi-
mental del IVITA, Iquitos, Peru, e-mails:

Neotropical Primates 14(1), January 2007

yahoo.es> and ) and Bruna Be-
zerra (University of Bristol, UK, e-mail: ac.uk>).


Primate Conservation, Incorporated (PCI) is a nonprofit
foundation founded to fund field research that supports
conservation programs for wild populations of primates.
Priority will be given to projects that study, in their natural
habitat, the least known and most endangered species. The
involvement of citizens from the country in which the pri-
mates are found will be a plus. The intent is to provide sup-
port for original research that can be used to formulate and
to implement conservation plans for the species studied.
PCI will grant seed monies or provide matching grants for
graduate students, qualified conservationists, and primatol-
ogists to study rare and endangered primates and their con-
servation in their natural habitat. All appropriate projects
will be considered, but the regions of current interest are
Asia and West Africa. For more information and grant ap-
plications, go to
or contact Ray Hamel at .


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

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

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

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

Application and award details: Individual awards tend to be
in the range of 250 to 500. Two closing dates apply: the
last day of February and the last day of August. Applica-
tions must be made on the Application Form, or following
the same format, and should be sent by post to reach the
Convener on or before the relevant closing date. Applica-
tions by e-mail or fax will not normally be accepted.

Applications forms are available on the PSGB website
or can be obtained directly from:
David A. Hill (Convenor of the CWP), School of Bio-
logical Sciences, University of Sussex, Falmer, Brighton
BN1 9QG, UK, Tel: +44 1273 606755 ext. 2755; Fax:
+441273678433, e-mail: .


Key Topics in Landscape Ecology: Key Issues in Theory, Meth-
odology, and Applications, edited by R. J. Hobbs and Jian-
guo Wu. 2007. Cambridge University Press. 400 pp. ISBN
0521850940. Landscape ecology is a relatively new area of
study, which aims to understand the pattern of interaction
of biological and cultural communities within a landscape.
This book brings together leading figures from the field to
provide an up-to-date survey of recent advances, identify
key research problems and suggest a future direction for de-
velopment and expansion ofknowledge. Providing in-depth
reviews of the principles and methods for understanding
landscape patterns and changes, the book illustrates con-
cepts with examples of innovative applications from differ-
ent parts of the world. Forming a current 'state-of-the-sci-
ence' for the science of landscape ecology, this book forms
an essential reference for graduate students, academics,
professionals and practitioners in ecology, environmental
science, natural resource management, and landscape plan-
ning and design. Contents: 1. Perspectives and prospects
on landscape ecology R. Hoobbs and Jianguo Wu; 2.
Adequate data of know accuracy are critical to advancing
the field of landscape L. R. Iverson; 3. Landscape pat-
tern analysis: key issues and challenges H- Li and J. Wu;
4. Spatial heterogeneity and ecosystem processes M. G.
Turner and J. A. Cardille; 5. Landscape heterogeneity and
metapopulation dynamics L. Fahrig; 6. Determining pat-

Neotropical Primates 14(1), January 2007

tern-process relationships in heterogeneous landscapes R.
H. Gardner, J. D. Forester and R. E. Plotkick; 7. Scale and
scaling: a cross-disciplinary perspective J. Wu; 8. Op-
timization of landscape pattern J. Hof and C. Flather;
9. Advances in detecting landscape changes at multiple
scales: examples of northern Australia J. A. Ludwig; 10.
The preoccupation of landscape research with land use and
land cover M. Antrop; 11. Applying landscape-ecologi-
cal principles to regional conservation: the wild Country
Project in Australia; 12. Using landscape ecology to make
sense of Australia's last frontier D. Bowman; 13. Transfer-
ring ecological knowledge to landscape planning: a design
method for robust corridors C. C. Vos, P. Opdam, E. G.
Steingrover and R. Reijnen; 14. Integrative landscape re-
search: facts and challenges G. Fry, B. Tress and G. Tress;
15. Landscape ecology: the state of the science J. Wu and
R. Hobbs.

Primate Anti-Predator Strategies, edited by Sharon L. Gursky
and K. A. I. Nekaris. 2007. Springer. 369pp. ISBN: 978-
0387348070. Part of the Series: Developments in Primatol-
ogy: Progress and Prospects. Since the 1960s, primatologists
have recognized the impact of predation on the evolution
of morphology, the social systems and cognitive behavior
of monkeys and apes, but few studies considered its impact
on the prosimians lemurs, lorises, galagos and tarsiers.
This comprehensive volume, written by experts in the field,
narrows this gap by highlighting the effect of predation on
the order Primates in general. Theoretical approaches to
understanding how primates perceive predation threat, as
well as proximate and ultimate causes to address threat and
attack, are considered across the primate order. Although
this volume concentrates on the least known group in this
theoretical area the prosimians contributions by re-
searchers on numerous primate taxa across four major geo-
graphical regions make this a novel and exciting contribu-
tion to students interested in primate evolution and ecology.
Contents: 1. Predation and Primate cognitive evolution K.
Zuberbiihler; 2. Predation on Primates: A biogeographical
Analysis D. Hart; 3. Primates and other prey in the sea-
sonally variable diet of Cryptoprocta ferox in the dry forest
of western Madagascar L. Dollar, J. U. Ganzhorn and
S. M. Goodman; 4. Predation on Lemurs in the rainfor-
est of Madagascar by multiple predator species: Observa-
tions and experiments S. M. Karpanty and P. C. Wright;
5. Predation, communication and cognition in Lemurs M.
Scheumann, A. Rabesandratana and E. Zimmermann; 6. A
consideration of leaping locomation as a means of preda-
tor avoidance in Prosimian Primates R. H. Crompton
and W. I. Sellers; 7. Anti-predator strategies of Cathemeral
Primates: Dealing with predators of the day and night I.
C. Colquhoun; 8. Moonlight and behavior in nocturnal
and cathemeral Primates, especially Lepilemur leucopus: Il-
luminating possible anti-predator efforts L. T. Nash; 9. A
comparison of calling patterns in two nocturnal primates,
Otolemur crassicaudatus and Galago moholi as a guide to pre-
dation risk S. K. Breader; 10. Predator defense by Slen-
der Lorises and Pottos K. A. I. Nekaris, E. R. Pimley and

K. M. Ablard; 11. The response of spectral tarsiers toward
avian and terrestrial predators S. L. Gursky; 12. Talk-
ing defensively, a dual use for brachial and exudate of slow
and pygmy Lorises L. R. Hagey, B. G. Fry and H. Fitch-
Snyder; 13. Anti-predator strategies in diurnal Prosimian,
the ring-tailed lemur (Lemur catta), at the Beza Mahafaly
Special Reserve, Madagascar L. Gould and M. L. Sauther;
14. Howler monkeys and harpy eagles: A communication
arms race- R. Gil-da-Costa; 15. Effects of habitat structure
on perceived risk of predation and anti-predator behavior of
vervet (Cerocpithecus aethiops) and patas (Erythrocebuspatas)
monkeys K. L. Enstam; 16. Predation risk and habitat use
in Chacma Baboons (Papio hamadryas ursinus) R. A. Hill
and T. Weingrill; 17. Reconstructing hominin interactions
with mammalian carnivores A. Treves and P. Palmqvist.

Ecological Census Techniques, edited by W J. Sutherland.
2006. Cambridge University Press. 448 pp. ISBN: 978-
0521606363. This is an updated version of the best selling
first edition, Ecological Census Techniques, with updat-
ing, some new chapters and authors. Almost all ecologi-
cal and conservation work involves carrying out a census
or survey. This practically focused book describes how to
plan a census, the practical details and shows with worked
examples how to analyse the results. The first three chapters
describe planning, sampling and the basic theory neces-
sary for carrying out a census. In the subsequent chapters
international experts describe the appropriate methods for
counting plants, insects, fish, amphibians, reptiles, mam-
mals and birds. As many censuses also relate the results to
environmental variability, there is a chapter explaining the
main methods. Finally, there is a list of the most common
mistakes encountered when carrying out a census. Contents:
1. Planning a research programme W. J. Sutherland; 2.
Principles of sampling J. J. D. Greenwood and R. A. Rob-
inson; 3. General census methods -J. J. D. Greenwood and
R.A. Robinson; 4. Plants -J. M. Bullock; 5. Invertebrates
- M. Ausden and M. Drake; 6. Fish I. M. C6td and M.
R. Perrow; 7. Amphibians T. R. Halliday; 8. Reptiles S.
Blomberg and R. Shine; 9. Birds D. W. Gibbons and R.
D. Gregory; 10. Mammals C. Krebs; 11. Environmental
variables J. C. Jones, J. D. Reynolds and D. Raffaelli; 12.
The twenty commonest censusing sins W J. Sutherland.

Primates in Perspective, edited by C. J. Campbell, A. Fuen-
tes, K. C. MacKinnon, M. Panger and S. K. Bearder, 2006.
Oxford University Press. 736 pp. ISBN: 9780195171334. It
is the first edited volume to offer a comprehensive overview
of primatology since 1987. Forty-four original essays-by
fifty-nine leading researchers in the field today-provide
wide-ranging and contemporary coverage of all of the major
areas of primatology. Arranged in six sections, the text
begins with an introduction to prim i ... 1. ._, and a review of
the natural history of the major taxonomic groups within
the order Primates. It goes on to cover methodologies and
research design for both field and captive settings; primate
reproduction; primate ecology and conservation and their
roles in the daily lives of primates; and such aspects of social

Neotropical Primates 14(1), January 2007

behavior and intelligence as communication, learning, and
cognition. The volume ends with a concluding chapter by
the editors that discusses the future of primatological re-
search. Ideal for introductory primatology courses, Primates
in Perspective can also be used in upper-division behavior
and conservation courses. Additionally, it is an invaluable
reference tool for primate researchers.

Mamiferos del Ecuador Gula de Campo, por Diego Tirira.
Ediciones Murcidlago Blanco. 576pp. ISBN: 9978-44-651-
6. La mas complete obra sobre la fauna de mamiferos del
Ecuador que se haya publicado hasta el present, con mas
de 380 species descritas. Para cada especie se incluye infor-
maci6n sobre: Identificaci6n (Principales medidas morfo-
m&tricas, Descripci6n de caracteristicas externas y patrons
de coloraci6n), Historia Natural (Ecologia, patrons de ac-
tividad, sociabilidad, hibitos alimenticios, refugios, areas de
vida, territories, locomoci6n, estrato utilizado y reproduc-
ci6n, Etologia), Sonidos, Distribuci6n y Habitat regionn,
rango altitudinal, tipos de bosques, mais de 350 mapas de
distribuci6n), Situaci6n Actual (Categorias de amenaza y
protecci6n segdn el Libro Rojo de los mamiferos del Ecua-
dor, UICN y CITES, principles amenazas y perspectives de
sobrevivencia de la especie, Areas protegidas). Ademis, claves
de identificaci6n, guia de huellas, comentarios taxon6mi-
cos, referencias '-..., *i, seleccionadas y anexos. Para
mayor informaci6n visitar: com/mamiferosdelecuador/publicaciones.htm#negro>.

ManagingProtectedAreas:A Global Guide. Edited by Michael
Lockwood, Graeme Worboys and Ashish Kothari, 2006.
Published with IUCN. 800PP. ISBN: 9781844073030.
Managing Protected Areas is an authoritative handbook,
produced by IUCN that spans the full terrain of protected
area management and is the international benchmark for
all professionals, students and academics worldwide. The
book employs dozens of detailed international cases stud-
ies, hundreds of concise topical snapshots, maps, tables,
illustrations and a colour plate section, as well as evalua-
tion tools, checklists and numerous appendices to cover all
aspects of park management from biodiversity to natural
heritage to financial management. he book establishes a
conceptual underpinning for protected area management,
presents guiding principles for the 21st century, reflects
recent work on international best practice and provides an
assessment of skills required by professionals. The publica-
tion is relevant to the full range of management systems
worldwide, balancing more traditional, developed country
approaches with developing country systems including
participatory, integrated, multi-sectoral and value-driven
approaches. Contents: Introduction, Part I: Setting the Con-
text, Natural Heritage, Social Context, Global Protected
Area Framework, Values and Benefits, Threats to Protected
Areas, Governance, Process of Management, Building Ca-
pacity, Part II: Principles and Practice, Establishing Pro-
tected Areas, Obtaining, Managing and Communicating
Information, Management Planning, Finance and Eco-
nomics, Administration, Sustainability Practice and Sus-

tainable Use, Operations Management, Natural Heritage
Management, Cultural Heritage Management, Managing
Threats, Incident Management, Tourism and Recreation,
Collaborative Management, Community Conserved Pro-
tected Areas, Linking the Landscape, Marine Protected
Areas, Evaluating Management Effectiveness, Building
Support, Futures and Visions.

The Conservation Project Manual. Edited by Colin Bibby
and Claire Alder, 2003. BP Conservation Programme,
Cambridge UK. ISBN: 1901930394. The basic aim of this
book is to help people to improve the process by which
they go about planning and managing small to medium-
sized conservation projects. By using a number of tried
and tested methods and illustrative examples, the manual
will enable users to work through their own concepts and
create a project plan. The manual is founded on the belief
that a good plan makes the fundraising and subsequent
implementation and management of a project easier and
success more likely. The book is divided in seven sections:
Section 1 Why Plan?, Section 2 Knowing where to fit in,
Section 3 Planning a Projetc, Section 4 Fundraising prin-
ciples, Section 5 Project Implementation, Section 6 Moni-
toring and Evaluation, and Section 7 Sources for Further
Information. This book is available in the web at conservation.bp.com/advice/project.asp>.


Aguiar, L. M., Ludwig, G., Svoboda, W. K., Teixeira, G. M.,
Hilst, C. L. S., Shiozawa, M. M., Malanski, L. S., Mello,
A. M., Navarro, I. T. and Passos, E C. 2007. Use of traps
to capture black and gold howlers (Alouatta caraya) on
the islands of the Upper Parand River, southern Brazil.
Am. J Primatol. 69(2): 241-247.
Anemone, R. L., Dirks, W., Moore, W. and Van Regen-
morter, J. 2007. Primate fossils, geological marker beds,
and anachronistic faunal assemblages from the early Pale-
ogene of southwestern Wyoming. Am.J. Phys. Anthropol.
Suppl. 44: 64-65.
Arroyo-Rodriguez, V., Serio-Silva, J. C., Alamo-Garcia, J.
and Ordano, M. 2007. Exploring immature-to-mother
social distances in Mexican mantled howler monkeys at
Los Tuxtlas, Mexico. Am. J Prim. 69(2): 173-181.
Barnett, A., Bezerra, B. and Andrade, E. 2007. Report of
two visits to Jad National Park, studying golden-backed
uacaris (Cacajao melanocephalus ouakary) and their habi-
tat, 13 Oct.-11 Nov. and 5-15 Dec., 2006. In: Manaus:
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Bloch, J. I., Silcox, M. T., Boyer, D. M. and Sargis, E. J.
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Clarke, M. R., Zucker, E. L. and Ford, R. T. 2007. Be-
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sex in monomorphic juvenile howlers (Alouatta palliata).
Am. J Primatol. 69(4): 477-484.

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Colquhoun, I. C. 2007. Anti-predator strategies of cathem-
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Couette, S. 2007. Morphologic and genetic differentia-
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Garber, P. A. 2007. Primate locomotor behavior and ecolo-
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Hart, D. 2007. Predation on primates: A biogeographi-
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multimodal communication applied to male mantled
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MacKinnon, K. C. 2007. Social beginnings: The tapestry
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havioral diversity in a radiation of primate seed predators.
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K. C. MacKinnon, M. Panger and S. K. Bearder (eds.),
pp.123-138. Oxford University Press, New York
Ruiz-Garcia, M., Escobar-Armel, P., Alvarez, D., Mudry,
M., Ascunce, M., Gutierrez-Espeleta, G. and Shostell,
J. M. 2007. Genetic variability in four Alouatta species
measured by means of nine DNA microsatellite markers:
Genetic structure and recent bottlenecks. Folia Primatol.
78(2): 73-87.
Sabbatini, G., Stammati, M., Tavares, M. C. H. and Vi-
salberghi, E. 2007. Response toward novel stimuli in a
group of tufted capuchins (Cebus libidinosus) in Brasilia
National Park, Brazil. Am. J Primatol. 69(4): 457-470.
Stone, A. I. 2007. Responses of squirrel monkeys to season-
al changes in food availability in an eastern Amazonian
forest. Am. J Primatol. 69(2): 142-157.
Stone, A. I. 2007. Age and seasonal effects on predator-
sensitive foraging in squirrel monkeys (Saimiri sciureus):
A field experiment. Am. J Primatol. 69(2): 127-141.
Sussman, R. W. 2007. A brief history of primate field stud-
ies. In: Primates in Perspective, C. J. Campbell, A. Fuentes,
K. C. MacKinnon, M. Panger and S. K. Bearder (eds.),
pp.6-10. Oxford University Press, New York.
Vogel E. R., Neitz, M. and Dominy, N. J. 2007. Effect
of color vision phenotype on the foraging of wild white-

Neotropical Primates 14(1), January 2007

faced capuchins, Cebus capucinus. Behav. Ecol. 18(2):
White, F J., Overdorff, D. J., Keith-Lucas, T., Rasmus-
sen, M. A., Kallam, W. E. and Forward, Z. 2007. Female
dominance and feeding priority in a prosimian primate:
experimental manipulation of feeding competition. Am. J.
Primatol. 69(3): 295-304.


Selected abstracts from the 76th Annual Meeting of the
American Association of Physical Anthropologists, Phil-
adelphia, PA, USA, 28-31 March, 2007, in: Am. J. Phys.
AnthropoL (Supp. 44): 60-256.

Alport, L. A. comparative analysis of lingual fungiform pa-
pillae and diet in primates, p.62.
Anderson IV, J. D., Owren, M. J. & Boinski, S. Impact-am-
plitude of controlled percussive strikes provides evidence
of site-selection for enhancement of acoustic display be-
havior in brown capuchins (Cebus apella) in Suriname,
Ascunce, M. S., Hasson, E., Mulligan, C. J. & Mudry, M.
D. Mitochondrial sequence diversity of the southernmost
extant New World monkey, Alouatta caraya, p.66.
Bales, K. L. Proximate mechanisms of paternal care in New
World primates, p.68.
Bezanson, M. Ontogenetic patterns of positional behav-
ior during play in Cebus capucinus and Alouatta palliata,
Boinski, S., Fragaszy, D. M., Ehmke, E. & Kauffman, L.
Wild brown capuchins in Suriname exploit combinatorial
manipulation for multiple functions, p.77.
Boore, J. L., DeGusta, D., Fourcade, H. M., Engel, D.,
Morgan, J. & Gignoux, C. R. Primate phylogeny and di-
vergence dates based on complete mitochondrial genomes,
Byron, C. D., Dhabliwala, J. & Bowles, K. Sagittal crests,
temporal fossae, and their putative relationships to mas-
ticatory function and encephalization in a diverse sample
of strepsirhine [sic] and platyrrhine primates, p.82.
Colburn, J. & Boinski, S. Effects of avian attendants on pre-
dation risk and foraging efficiency of squirrel monkey and
brown capuchin troops in Suriname, p.88.
Cole III, T. M. Stabilizing selection and adaptive optima in
the evolution of platyrrhine body size, p.88.
Coleman, M. N., Kay, R. E & Colbert, M. W Evaluating
hearing sensitivity in Homunculuspatagonicus, p.89.
Constantino, P. Evidence of convergent masticatory mor-
phology among durophagous anthropoids from four dif-
ferent clades, p.89.
Cooke, S. B., Halenar, L. B., Rosenberger, A. L., Tejedor,
M. F. & Hartwig, W. C. Protopithecus, Paralouatta, and
Alouatta: The making of a platyrrhine folivore, p.90.
Crofoot, M. C. Dominance and inter-group relationships:
Do group identity, group size and interaction location

affect the outcome of between-group contest competi-
tion in Cebus capucinus?, p.93.
DeLuycker, A. M. Activity pattern and habitat use of the
Rio Mayo titi monkey (Callicebus oenanthe) in a premon-
tane forest in the Alto Mayo, northern Peru, p.96.
Derby, A. M. Ecological influences on red howler monkey
density: A comparison of two Western Amazonian sites,
Di Fiore, A. & Fernandez-Duque, E. A comparison of
paternal care in three socially-monogamous neotropical
primates, pp.99-100.
Dietz, J. M. Why mating systems vary in cooperative-
breeding lion tamarins, p.100.
Digby, L. J. Determining home range "volumes" in pri-
mates: Why are we using two-dimensional measures for
species that live in a three-dimensional world?, p. 100.
Dobson, S. D. Social cohesion and the evolution of facial
expression in nonhuman anthropoids, pp. 100-101.
Eng, C. M., Ward, S. R., Winters, T. M., Kingsbury, T. D.,
Vinyard, C. J. & Taylor, A. B. Mechanics of the mastica-
tory apparatus favor muscle force production at wide jaw
gapes in tree-gouging marmosets, p.107.
Fernandez-Duque, E. Costs and benefits of paternal care in
free-ranging owl monkeys (Aotus azarai), p. 108.
Field, M. Y. If you give a monkey an onion: An introduc-
tion to fur rubbing in human-commensal white-fronted
capuchin monkeys (Cebus ilb],f ons), p.108.
Fleagle, J. G. & Gilbert, C. C. Comparing primate crania,
Ford, R. T. Lack of special relationships between male and
female mantled howling monkeys (Alouatta palliata),
Ford, S. M. & Boinski, S. Primate predation by harpy
eagles in the Central Suriname Nature Reserve, p.109.
Garber, P. A., Gomes, D. E & Bicca-Marques, J. C. Experi-
mental field study of handedness in wild tufted (Cebus
nigritus) and white-faced (Cebus capucinus) capuchins:
Evidence for individual and species differences, p. 112.
Guillot, D. M., Lawler, R. R. & MacLatchy, L. M. Patterns
of positional behavior among atelines: A comparative
analysis of Alouatta seniculus, Lagothrix lagotricha, and
Ateles belzebuth in Ecuador, p.121.
Hogg, R. T. Canine dimorphism, dental growth, and the
evolution of anthropoid mating systems: The platyrrhine
angle, p.130.
Jack, K. M., Lenz, B. B., Healan, E., Rudman, S., Schoof,
V. & Fedigan, L. M. The effects of observer presence on
the behavior of three groups of Cebus capucinus in Area
de Conservaci6n Guanacaste, Sector Santa Rosa, Costa
Rica, pp.134-135.
Janson, C. H. Predator detection and the evolution of pri-
mate sociality: Insights from experiments on a rain forest
primate, p. 136.
Kamilar, J. Predicting within-species variation in primate
behavior and ecology: A quantitative approach using
comparative methods, p.139.
Kitchen, D. M., Kenney, S. & Wilson, R. E. Do male black
howler monkeys (Alouatta pigra) assess the fighting abil-

Neotropical Primates 14(1), January 2007

ity of individual rivals during inter-group encounters?,
Knittel, J. M. Trabecular and cortical architecture of captive
versus wild specimens of Loris tardigradus and C
jacchus using micro-computed tomography, p. 145.
Kowalewski, M. & Garber, P. A. Mating promiscuity, en-
ergetics, and reproductive strategies in black and gold
howler monkeys (Alouatta caraya), p. 147.
Lawrence, J. M. The pair bond in brown titi monkeys
(Callicebus brunneus): Male and female reproductive in-
terests, pp. 151-152.
Luecke, L. G., Martinez-Mota, R. & Estrada, A. Census-
es of parapatric howler monkeys (Alouatta palliata and
Alouattapigra) in Pantinos de Centla Biosphere Reserve,
Tabasco, Mexico, p.160.
Lynch Alfaro, J. W., Gutikrrez-Espeleta, G., Olson, L. E.,
Ross, V., Neitzel, S., Sukraw, K., Drown, D. M. & Alfaro,
M. E. Patterns of mtDNA genetic diversity in capuchin
monkeys at regional and local scales, p. 161.
Matthews, L. J. & Schmitt, C. A. Courtship behaviors of
genus Cebus: A test case for inferences from phylogeny,
McKusick, G. L. & Teaford, M. E Premolar microwear of
three New World monkeys: Cebus apella, Pitheciapithecia,
and Ateles belzebuth, p. 169.
McLaughlin, M. V. & Fernindez-Duque, E. Spatial dis-
tribution of feeding and sleeping trees in territorial owl
monkeys (Aotus azarai), pp.169-170.
Mendoza, S. P. & Mason, W. A. What should we be look-
ing for in brain and physiology to understand paternal
care?, pp.170-171.
Morales-Jimenez, A. L. Body size and home range in non-
human primates: Do they scale?, p.174.
Mork, A. L., Horton, W. E. & Vinyard, C. J. Comparative
histology of the mandibular condylar cartilage in gouging
and non-gouging platyrrhines, p. 174.
Murphy, W. J. The use of rare genomic changes in mamma-
lian phylogeny estimation and the phylogenetic position
of Primates, p. 176.
O'Malley, R. C. & Fedigan, L. M. Hand preferences
during foraging in white-faced capuchins (Cebus capuci-
nus), p.180.
Organ, J. M. Evolutionary convergence of tail structure in
prehensile- and nonprehensile-tailed primates and pro-
cyonids, p. 181.
Peker, S., Pave, R., Zunino, G. E. & Kowalewski, M. M.
Observation of a birth in wild black and gold howler
monkeys (Alouatta caraya), p.186.
Porter, L. M. & Garber, P. A. Niche expansion of a cryptic
primate, Callimico goeldii, during polyspecific associa-
tions, p. 191.
Rakhovskaya, M. V., Fernindez-Duque, E. & Di Fiore, A.
The effects of demographic and ecological factors on ter-
ritory size and ranging patterns of Argentinean owl mon-
keys (Aotus azarai), p.195.
Righini, N. & Martinez-Mota, R. Spatial proximity and
association patterns in four groups of wild black howler
monkeys (Alouattapigra) in Southern Mexico, p.199.

Robson, S. L. Encephalization and life history: Lessons
from primate brain growth trajectories, p.201.
Rodman, P. S. & Bossuyt, E J. Fathers and stepfathers: Fa-
milial relations of old and new males within groups of
Callicebus brunneus in southeastern Peri, p.201.
Rodrigues, M. R. & Lindshield, S. L. Scratching the sur-
face: Observations of tool use in wild spider monkeys,
Ross, C. E & Reed, D. A. Modulation of mastication to varia-
tion in food material properties in Cebus capucinus, p.203.
Rutherford, J. G., Layne Colon, D. & Tardif, S. D. Using
ultrasound to determine gestational age and litter size in
marmoset monkeys, pp.205-206.
Schmidt, M. Kinematics and proportions of the three-
segmented limb: How are small primates different from
other small mammals?, p.208.
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. & Fernindez-Duque, E. Com-
parative ranging behavior of eight species of primates in a
western Amazonian rainforest, pp.208-209.
Schrago, C. E. G. & Russo, C. A. M. Assessment of the time
of origin of New World primates and rodents, p.209.
Seltzer, L. J. & Ziegler, T. E. Release of social hormones in
response to visual cues in a pair-bonded primate, p.214.
Sendall, C., Fernindez-Duque, E. & Di Fiore, A. A pre-
liminary study of mate-guarding in wild titi monkeys
(Callicebus discolor), pp.214-215.
Shattuck, M. R. Color vision and food detection in squirrel
monkeys (Saimiri boliviensis), pp.215-216.
Shepston, D. S. Return of Alouatta pigra to the forests
around Nahi, Chiapas, Mexico, p.216.
Shibata, C. & Ford, S. M. The maintenance of social bonds
in adult pairs of captive cotton-top tamarins (Saguinus
oedipus), p.217.
Taylor, A. B. & Vinyard, C. J. Jaw-muscle fiber architecture
in Cebus, p.229.
Thompson, C. L., Jackson, E. M., Stimpson, C. D. &Vin-
yard, C. J. Assessing how experimental and surgical ma-
nipulations during in vivo laboratory research influence
chewing speed in tufted capuchins (Cebus apella), p.231.
Valenta, K. & Fedigan, L. M. The effect of gut passage by
Cebus capucinus on rates of seed germination and time to
germination, p.235.
Vinyard, C. J., Thompson, C. L., Stimpson, C. D., Mork,
A. L., Armfield, B. A., Doherty, A. H., Wasserman, H.
M., Jackson, E. M. & Horne, W. I. Masseter muscle strain
during chewing in tufted capuchins (Cebus apella), p.239.
Walker, S. E. & Davis, L. C. Postcranial features of Cacajao,
with comparisons to Chiropotes and Pithecia, p.242.
Ward, C. V., Plavcan, J. M. & Paulus, F L. Canine tooth
crown, root and facial dimorphism in anthropoid pri-
mates, p.243.
Wheeler, B. C. The function of alarm calling among wild
tufted capuchin monkeys (Cebus apella), p.248.
Williams, S. H., Vinyard, C. J., Glander, K. E., Teaford,
M. E, Deffenbaugh, M. & Thompson, C. L. A telemetry
system for studying jaw-muscle activity in free-ranging

Neotropical Primates 14(1), January 2007

primates: Pilot data from howling monkeys (Alouatta
palliata) at La Pacifica, Costa Rica, p.250.
Wolovich, C. K., Perea-Rodriguez, J. P. & Fernandez-
Duque, E. Food sharing as a form of paternal care in wild
owl monkeys (Aotus azarai), p.252.
Young, J. W. Substrate alters asymmetrical gait dynamics in
common marmosets (C .-' jacchus), p.254.
Ziegler, T. E. What makes a good dad? Neuroendocrine
factors influencing paternal care, pp.255-256.


25"' Annual Conference of the Australasian Primate Society.
March 9-11, Queensland, Australia, University of Queensa-
land St. Lucia Brisbane. For more information consult the
web site: .

The Mind of the Chimpanzee. March 22-25, The Lincon
Park Zoo, Chicago, Illinois. In the tradition of the "Un-
derstanding Chimpanzees" conferences, which started
20 years ago, "The Mind of the Chimpanzee" conference
will bring together the top experts in the fields of chimpan-
zee cognition and conservation as well as the "next genera-
tion" of chimpanzee researchers in order share new research
findings, generate new collaborative research partnerships
and examine how studying chimpanzee cognition impacts
chimpanzee conservation. The Lester E. Fisher Center
for the Study and Conservation of Apes is proud to host
this historic event organized by Dr. Elizabeth Lonsdorf
and Steve Ross. For more information contact mind@lpzoo.org>, or visit the web site: chimpmindconference.org/>.

Annual IACUC Conference. March 26-27, Town and
Country Resort, 500 Hotel Circle, San Diego, CA. The
Public Responsability in Medicine and Reseach will held
the annual IACUC conference. This event will include a
range of keynote addresses, panels, workshops, and didac-
tic sessions, all of which are designed to help build, and
then strengthen, an effective animal care and use pro-
gram. For more information visit: education/2007_IACUC/overview_IACUC07.html>.

The Annual Meeting of the American Association of
Physical Anthropologists. March 27-April 1, Philladel-
phia, PA. For program information contact Program Chair
Dennis H.O'Rourke at the web site: .

Workshop & Symposium on Laboratory Animal Dis-
eases. 18-21 April. Chicago, Illinois. The Midwest Div.
of The Charles Louis Davis, D.V.M. Foundation in Co-
sponsorship with The Biologic Resources Laboratory

(BRL) of The University of Illinois at Chicago will pres-
ent a Workshop and Symposium on Laboratory Animal
Diseases. The Workshop will begin Wednesday the 18'
of April, continuing through Friday the 20th of April.
The Symposium will be held on Saturday April 21t. For
more information contact James E. Artwohl at edu.> or visit the web site: CLDavis.meetings.htm#20071abdisease>.

Callitrichid Workshop 2007. May 12-13, Providence,
Rhode Island. The 6th annual Callitrichid Husbandry Work-
shop will be a free workshop focusing on the education and
development of callitrichid keepers and managers in zoos
and aquariums. Some of the topics to be covered are: Biol-
ogy and Husbandry, Veterinary care, In Situ conservation.
For more information consult the web page: rwpzoo.org/calendar/callitrichid.cfm>.

87th Meeting of the American Society of Mammalogists.
June 6-10, Museum of the Southwestern Biology, Albu-
querque, New Mexico. Some of the topics of the meeting are
Biogeography, Conservation, Population Ecology and Ge-
netics among others. For more information and registration
go to the web site:

10th FELASA Symposium and the XIV ICLAS General As-
sembly & Conference. June 11-14. The FELASA-ICLAS
Joint Meeting 2007, hosted by AISAL, Associazione Itali-
ana per le Scienze degli Animali da Laboratorio, will take
place in Northern Italy, on the shores of Lake Como. The
international meeting will include the 10' FELASA Sym-
posium and the XIV ICLAS General Assembly & Confer-
ence, and will provide a comprehensive overview of the
most recent developments in the field of laboratory animal
sciences and technologies. For more information go to
, or
contact Stefania Sella at .

Training Workshop on Ethical Considerations and Bio-
medical Use of Non Human Primates for Research in
Tropical Diseases. June 18-27, Karen, Nairobi, Kenya.
Non human primates will continue to play a pivotal role
as models of human diseases including tropical infections.
However the use of these animals needs to be carefully
applied so that proper consideration of ethical and welfare
issues is incorporated in all aspects of research involving
non-human primates (NHP). This workshop, supported
by TDR and the Institute of Primate Research, National
Museums of Kenya with coordinated assistance from SSI
coordinators, is a ten-day training course with lectures,
hands-on laboratory sessions and demonstrations. Em-
phasis will be placed on biomedical techniques used in
primates in addressing TDR diseases, including the ethi-
cal and welfare aspects of using NHP in research. Details
of the contents and tentative schedule can be viewed at

Application deadline in 14th April 2007. For more in-

Neotropical Primates 14(1), January 2007

formation visit the web site ipr-nmk/callforapplication.html>.

30th Meeting of the American Society of Primatologists.
June 20-23, 2007, Winston-Salem, NC. Hosted by the
Wake Forest University School of Medicine. For more
information consult the web site: asp2007/index.htm>.

Pan African Sanctuary Alliance 2007 Management
Workshop. June 21-24, Kigali, Rwanda. The Pan African
Sanctuary Alliance (PASA) will focus on issues such as
law enforcement, eco-tourism, and disease control at the
PASA 2007 Management Workshop. PASA was formed
in 2000 to promote unity and cooperation among the
primate sanctuaries of Africa, and its members represent
17 sanctuaries in 12 African countries. The PASA Manage-
ment Workshop has been an annual event since 2000. For
more information go to org/index.htm>.

XII Meeting of the Sociedade Brasileira de Primatologia.
Julho 22-27, Minas Gerais, na Pontificia Universidade
Cat61lica de Minas Gerais (PUC-Minas), Campus Coradao
Eucaristico, em Belo Horizonte. 0 campus se localiza a
Av. Dom Jos6 Gaspar, 500, no Bairro Coracao Eucaristico,
regiao oeste de Belo Horizonte. Para mais informacao visit
o web page: .

Annual Meeting of the Association for Tropical Biol-
ogy and Conservation. July 15-19, Morelia, Michoacan,
M6xico. The meeting will cover a wide array of basic and
applied research topics on tropical biology and conserva-
tion, including: origin, evolution, and maintenance of
tropical biodiversity (TD); structure, dynamics and func-
tioning of tropical ecosystems (TEs); anthropogenic ef-
fects on TD and TEs; and socio-cultural-economical driv-
ers of such effects. Registration, abstracts for symposium,
and abstracts for contributed papers and posters must be
submitted on-line by April 15'. More information and de-
tails may be found on the web page: mx/atbc2007>.

44th Annual Meeting of the Animal Behavior Society. July
21-25, Burlington Sheraton Hotel and Conference Center,
Burlington, Vermont. For more information go to: www.animalbehavior.org/ABS/Program/>.

Laboratory Animal Welfare Training Exchange 2007 Bi-
ennal Conference. August 8-10, Radisson Hotel, down-
town Boston. The main issue will be to relieve pain in
laboratory animals... and yourself. For more information,
fees, schedule and registration go to: org/conference.html>.

6"' World Congress on Alternatives & Animal Use in the
Life Sciences (WC6). August 21-25, Tokyo, Japan. With the
support and organization of the Japanese Society ofAlterna-

tives to Animal Experiments (JSAAE), the Alternative Con-
gress Trust (ACT), and the Science Council of Japan (SCJ).
The WC6 is the first congress to be held in Eastern Asia and
will be a good opportunity to review animal welfare issues
and to strongly encourage research on alternative methods
in this region. Some of the topics of the congress will be:
Animal welfare, Moral Ethical and Cultural issues and public
policies of animal usage, and Knowledge management and
information services. For more information visit the web
site: .

2nd Congress of the European Federation for Prima-
tology. September 3-7, Faculty of Education, Charles
University, Prague. Organized by the Czech Group of
Primatologists, this Congress aims to step in the future
by inviting also those colleagues who are able to present
anthropological topics interesting for primatologists 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
ecology and conservation among others. For more in-
formation and registration go to: webbio/efp/efp_pr',,,,"7.pdf>, or visit the web page:

The 25t' Annual Symposium for Nonhuman Primate
Models for AIDS. September 10-13, 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 cru-
cial nonhuman primate studies to better understand how
HIV and SIV cause disease, and to facilitate the develop-
ment of new methods for the treatment, control and pre-
vention 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 pathogenesis, vaccines, and therapeutic approaches
against primate lentivirus infection and disease; primate
genomics; viral agents associated with simian acquired im-
munodeficiency syndrome; and the mechanisms of natu-
ral resistance in several 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: NHPM2007/>.

III Congreso Mexicano de Primatologia. Octubre
24-27. 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 UNAM. For more information please
visit the web site: .

VIII Curso Latino-Americano de Biologia da Conser-
vacio e Manejo da Vida Silvestre. Novembro 06 a 12 de
Dezembro. Na 61tima d6cada o Curso Latino Americano

Neotropical Primates 14(1), January 2007

de Biologia da Conservagao e Manejo da Vida Silvestre
vem contribuindo para a formacao de profissionais com-
prometidos corn a conservagao da diversidade socioambi-
ental da Am&rica Latina. Durante as cinco semanas de ca-
pacitagao os participants compartilham experiencias com
renomados profissionais que atuam nas diferentes esferas
da Biologia da Conservagao. 0 conteddo programatico
deste curso trata primeiramente de uma abordagem con-
ceitual te6rica, seguido de metodologias e ferramentas tteis
em estudos de campo e por ltimo, 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 Nazarn Paulista
- SP e no municipio de Teodoro Sampaio SP. Para mais
informacao visit o web page: cursos.asp?mes=Novembro&id= 104>.

Animal Training & Behavior Through Positive Reinforce-
ment- Further Challenging and Advanced Issues. Decem-
ber 7-9, Munich, Germany. The seminar will be imparted
by Ken Ramirez, vice president for animal collections and
animal training; he develops and supervises animal-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 op-
erant conditioning, training situations and problem solving
with positive reinforcement. For more information visit:

6.- Gbttinger Freilandtage. Primate Behavior and Human
Universals. December 11-14, 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 weg page: de/welcome.html>.


XXIInd IPS Congress. August 3-8, Edinburg International
Conference Center, Edinburg, Scotland. Sponsored by the
Primate Society of Great Britain. For information consult
the web site: .


the most biologically
diverse countries in the world
due to its rich and varied flora
and fauna and is superseded
only by Brazil and Peru in
terms of primate diversity.
This field guide illustrates and
describes 28 primate species
comprising 43 different taxa,
15 of which are endemic to
Colombia. It is a compilation
of all primate field work done
on Colombian primates both
in and out of country and has
quickly become an important
tool for young primatologists to establish research
priorities for study. The field guide also includes
chapters on primate classification, fossil history,
zoogeography, conservation and phylogeny, and
is a first step towards the necessary conservation
of this beautiful group of animals.

About the author, Thomas Richard DefHer:

Tom Defler is a primatologist who has spent the last
28 years in the Orinoco and Amazonian regions of Colom-
bia, focusing his studies on the ecology and conservation
of primate species in these two regions of the country. His
research began in 1976 with INDERENA of the Minis-
try of Agriculture (now superceded by the Ministry of the
Environment) studying Colombian flora and fauna. He
established two research stations, Capard Biological Sta-
tion and Ecological Station Om&. Defler has written more
than 60 publications on diverse aspects of ecology, primate
taxonomy and natural history. He is currently Professor at
the Instituto Amaz6nico de Investigaciones, at the National
University of Colombia in Leticia.

pauses con mayor diversi-
of Colombia dad biol6gica del mundo debido
a su rica y variada flora y fauna;
s6lo Brasil y Peri la superan en
terminos de diversidad de prima-
tes. Esta gula de campo ilustra y
describe 28 species de primates
conteniendo 43 distintos taxo-
nes, 15 de los cuales son end6-
micos de Colombia. Es una
recopilaci6n de todo el trabajo
de campo que fue realizado sobre
primates colombianos dentro y
fuera del pais, y se ha convertido
ripidamente en una important
herramienta para j6venes prima-
t6logos que desean establecer prioridades de inves-
tigaci6n para sus studios. La gula de campo tam-
bi2n include capitulos sobre clasificaci6n de pri-
mates, historic f6sil, zoogeografia, conservaci6n y
filogenia, y es un primer paso hacia la conservaci6n
necesaria de este hermoso grupo de animals.

Thomas Richard Defter, biograffa de/ author:

Tom Defler es un primat6logo que ha pasado los 1ltimos
28 afios en las regions del Orinoco y Amazonia de Colom-
bia, enfocando sus studios en la ecologia y conservaci6n de
species de primates en estas dos regions del pals. Inici6 sus
investigaciones en 1976 con el INDERENA del Ministerio
de Agriculture (ahora Ministerio del Ambiente), estudiando
la flora y fauna colombiana. Estableci6 dos estaciones de
investigaci6n, la Estaci6n Biol6gica Capardi y la Estaci6n
Ecol6gica Om6. Defler ha escrito mais de 60 publicaciones
sobre distintos aspects de la ecologia, taxonomia e histo-
ria natural de primates. En la actualidad, es professor del
Institute Amaz6nico de Investigaciones, en la Universidad
Nacional de Colombia en Leticia.

Primates of Colombia

Mail and Fax Order Form

Book Title: Primates of Colombia by Thomas Richard Defler. Illustrations by Stephen D. Nash,
Cesar Landazabal Mendoza and Margarita Nieto Diaz. Editor: Jose Vicente Rodriguez-Mahecha.

English ISBN: 1-881173-83-6. Softcover.
Spanish ISBN: 1-881173-73-9. Softcover.

Price: $40.00 (includes UPS Ground shipping within the continental United States). For orders
requiring faster service than UPS Ground, you will be responsible for paying all shipping costs.

Please email or call the phone number listed below for overnight deliveries and wholesale orders.

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

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made payable to Conservation International. Please allow 2-3 weeks for delivery.
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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: Jilio C6sar Bicca-Marques, Departamento de Biodiversidade
e Ecologia, Pontificia Universidade Cat61lica do Rio Grande do Sul,
Av. Ipiranga, 6681 Pr6dio 12A, PortoAlegre, 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/ore-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, Systematics,
Genetics (when relevant for systematics and conservation), Bioge-
ography, 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 Portuguese 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 fundamental for the text (as in species descrip-
tions, 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, including 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 conservation and research programs
(who, what, where, when, why, etc.) and most particularly informa-
tion on geographical distributions, locality records, and protected
areas and the primates that occur in them. Text should be typewrit-
ten, 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 references).

Figures and maps. Articles may include small black-and-white
photographs, high-quality figures, and high-quality maps and tables.
Please keep these to a minimum. We stress the importance of provid-
ing 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 publications,
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 chronologically
and then in alphabetical order. For example, "... (Fritz, 1970; Albert,
1980, 2004; Oates, 1981; Roberts, 2000; Smith, 2000; Albert et al.,
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 TropicalRain 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: 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 "Dissertaqao de Mestrado" and "Tesis de
Maestria" for articles in Portuguese and Spanish respectively.
"Unpublished report" changes to "Relat6rio T6cnico" 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(1), 2007


E d ito rial .................................................................................................................................................... ................................. 1


Density, Habitat Use, and Ranging Patterns of Red Howler Monkeys in a Colombian Andean Forest
Carolina Gdmez-Posada, Jessis Martinez, Paula Giraldo and Gustavo H. Kattan ............................................................................2...
Distribution of the Black Howler Monkey (Alouattapigra) and the Mantled Howler Monkey (A. palliata) in their
Contact Zone in Eastern Guatemala
Am rei Baumgarten and G. Bruce -i ",, ... ........................... ............................................... ..................... ............................11
An Experimental Census Method and Estimates of Population Density of a Black Howler Monkey (Alouatta pigra)
Highland Population in the Sierra Yalijux, Guatemala
Swen C Renner, Regula Rieser and Robert H. Horwich......................................................................................................19

Short Articles

Capuchin Monkey (Cebus apella) Vocalizations in Response to Loud Explosive Noises
Briseida D~go de Resende, Dilmar A. G. Oliveira, Eduardo D. Ramos da Silva and Eduardo B. Ottoni ........................25
Extragroup Copulations Among Brown Howler Monkeys in Southern Brazil
M arcos de Souza Fialho and Eleonore Z. E Setz .......................................... ........... ............... .................................................. 28
Golden Lion Tamarins, Leontopithecus rosalia (Linnaeus, 1766) in the Taquara Municipal Natural Park (Duque de
Caxias, RJ): A Southern Extension of the Known Range
Carlos Henrique de Freitas Burity, Leandro Duarte da Cruz, Vera Lucia Rocha, Nelson Barroso da Conceifao,
Daniel Eduardo da Luz, Durval da Silva Santos, Devylson da Costa Campos andAlcides Pissinatti ................................................. 30
Black-tufted-ear Marmoset Callithrixpenicillata (Primates: Callitrichidae) Following the Army Ant Labidus praedator
(Formicidae: Ecitoninae) in the Cerrado and the Atlantic Forest, Brazil
TadeuArtur de Melo Junior and Fernando Jose Zara ................................................................................... 32

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

P rim ate Societies ........................................................................................................................................................................38

R ecen t P u blication s ...................................................................................................................................................................38

M eetin g s ..................................................................................................................................................................................... 4 4

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