Front Cover
 Back Matter
 Back Cover

Title: Neotropical primates
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00098814/00055
 Material Information
Title: Neotropical primates a newsletter of the Neotropical Section of the IUCNSSC Primate Specialist Group
Abbreviated Title: Neotrop. primates
Physical Description: v. : ill. ; 27 cm.
Language: English
Creator: IUCN/SSC Primate Specialist Group -- Neotropical Section
IUCN/SSC Primate Specialist Group -- Neotropical Section
Conservation International
Center for Applied Biodiversity Science
Publisher: Conservation International
Place of Publication: Belo Horizonte Minas Gerais Brazil
Belo Horizonte Minas Gerais Brazil
Publication Date: January 2008
Frequency: quarterly
Subject: Primates -- Periodicals -- Latin America   ( lcsh )
Primates -- Periodicals   ( lcsh )
Wildlife conservation -- Periodicals   ( lcsh )
Genre: review   ( marcgt )
periodical   ( marcgt )
Spatial Coverage: Brazil
Additional Physical Form: Also issued online.
Language: English, Portuguese, and Spanish.
Dates or Sequential Designation: Vol. 1, no. 1 (Mar. 1993)-
Issuing Body: Issued jointly with Center for Applied Biodiversity Science, <Dec. 2004->
General Note: Published in Washington, D.C., Dec. 1999-Apr. 2005 , Arlington, VA, Aug. 2005-
General Note: Latest issue consulted: Vol. 13, no. 1 (Apr. 2005).
 Record Information
Bibliographic ID: UF00098814
Volume ID: VID00055
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: oclc - 28561619
lccn - 96648813
issn - 1413-4705


This item has the following downloads:

NP15 ( PDF )

Table of Contents
    Front Cover
        Front Cover 1
        Front Cover 2
        Page 1
        Page 2
        Page 3
        Page 4
        Page 5
        Page 6
        Page 7
        Page 8
        Page 9
        Page 10
        Page 11
        Page 12
        Page 13
        Page 14
        Page 15
        Page 16
        Page 17
        Page 18
        Page 19
        Page 20
        Page 21
        Page 22
        Page 23
        Page 24
        Page 25
        Page 26
        Page 27
        Page 28
        Page 29
        Page 30
        Page 31
        Page 32
        Page 33
        Page 34
        Page 35
        Page 36
        Page 37
        Page 38
    Back Matter
        Back Matter 1
        Back Matter 2
        Back Matter 3
    Back Cover
        Back Cover
Full Text

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

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

ISSN 1413-4703 Abbreviation: Neotrop. Primates

Erwin Palacios, Conservaci6n Internacional Colombia, Bogoti DC, Colombia
Liliana Cort6s Ortiz, Museum of Zoology, University of Michigan, Ann Arbor, MI, USA
J6lio Cesar Bicca-Marques, Pontificia Universidade Cat61lica do Rio Grande do Sul, Porto Alegre, Brasil
Eckhard Heymann, Deutsches Primatenzentrum, Gattingen, Germany
Jessica Lynch Alfaro, Washington State University, Pullman, WA, USA
Liza Veiga, Museu Paraense Emflio Goeldi, Beltm, Brazil

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

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 E Ferrari, Universidade Federal do Sergipe, Aracajfi, Brazil
Russell A. Mittermeier, Conservation International, Arlington, VA, USA
Marta D. Mudry, Universidad de Buenos Aires, Argentina
Horicio Schneider, Universidade Federal do Pard, Campus Universitirio de Braganga, Brazil
Karen B. Strier, University of Wisconsin, Madison, WI, USA
Maria Emilia Yamamoto, Universidade Federal do Rio Grande do None, Natal, Brazil
Primate Specialist Group
Chairman, Russell A. Mittermeier
Deputy Chair, Anthony B. Rylands
Coordinator, Special Section on Great Apes, Liz Williamson
Regional Coordinators Neotropics
Mesoamerica, Ernesto Rodriguez Luna
Andean Countries, Erwin Palacios and Eckhard W. Heymann
Brazil and the Guianas, M. Cecilia M. Kierulff
Regional Coordinators Africa
West Africa, John 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: Glenda Fibregas, 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: Aggressive stare of white-fronted capuchin (Cebus albifrons) in upper subtropical forest (2,515 m) in southern Ecuador.

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 15(1), January 2008 1


Katharine Milton1

1 Dept. Environmental Science, Policy &Management, University of California Berkeley, CA 94720-3114, USA, kmilton@
berkeley. edu


To attract seed dispersal agents, most Neotropical tree species produce edible fruits. Animals eat the fruits, often swallowing
the seeds, which may be deposited away from the parent tree. Data show that not all frugivores in a given habitat are equally
attracted to the same fruit species. Though numerous factors can influence fruit choice, the nutrient value of the pulp to the
disperser is clearly of key importance. Here data are presented on the macronutrient content (total crude protein, fat and
nonstructural carbohydrate) of 19 fruit species collected from a single forest site, Barro Colorado Island in central Panama.
The only criterion for analyses was that each species produce fruits consumed by at least two primate species at this site.
Though all fruit species contained a measurable amount of each macronutrient class, there were often striking differences in
the amount contributed per class. Overall, two species (10.5% of the total sample) were highest in protein, five (26%) were
highest in fats and twelve (63%) were highest in nonstructural carbohydrates. Fruit species in the same family or genus did
not necessarily show the same macronutrient pattern. Similar data from a range of Neotropical sites could prove useful in
formulating hypotheses related to community fruit-disperser relationships.

Key Words: Tropical forest, Panama, fruits, nutrients, protein, fats, nonstructural carbohydrates, frugivores, seed dispersal.


Para atraer a los agents dispersores de semillas, muchas de las species de irboles Neotropicales produce frutos comes-
tibles. Los animals comen los frutos, a menudo tragando las semillas, las cuales pueden ser depositadas lejos del irbol
parental. Los datos muestran que no todos los frugivoros en un habitat dado son igualmente atraidos por las mismas species
de frutos. A pesar de que numerosos factors pueden influir la selecci6n de frutos, el valor nutricional de la pulpa para el
dispersor es claramente de fundamental importancia. Aqui se presentan datos del contenido de macronutrientes proteina
cruda total, grasa y carbohidratos no estructurales) de 19 species de frutos colectados en un solo bosque en la Isla de Barro
Colorado, region central de Panami. El dnico criterio para el anAlisis fue que cada especie produjera frutos consumidos por
al menos dos species de primates en dicho sitio. Aunque todas las species de frutos contuvieron una cantidad apreciable
de cada clase de macronutriente, hubo a menudo marcadas diferencias en la cantidad aportada por cada clase. En general,
dos species (10.5% de la muestra total) tuvieron mais alta protein, cinco (26%) mais alto contenido de grasas y doce (63%)
mais alto contenido de carbohidratos no estructurales. Las species de frutos de una misma familiar o gdnero no mostraron
necesariamente el mismo patr6n de macronutrientes. Datos similares de diferentes sitios en el neotr6pico serian muy tiles
para formular hip6tesis acerca de las relaciones de las comunidades de dispersores de semillas.

Palabras Clave: Bosque tropical, Panami, frutos, nutrients, protein, grasas, carbohidratos no estructurales, frugivoros,
dispersi6n de semillas.


The overwhelming majority of tropical tree species rely on animal vectors to disperse seeds to potential germination sites
away from the parent (van der Pijl, 1969; Snow, 1971; Howe and Vande Kerckhove, 1979; Howe, 1993; Stiles, 1993;
Herrera, 2002). To facilitate dispersal, seeds are enveloped in an edible matrix-fleshy mesocarps, pericarps, arils and the
like-that serves as an attractant for potential dispersal agents. Animals enter the tree to eat the fruit, often swallowing the
seeds, which may later be regurgitated or defecated some distance away. Some frugivores (e. g., various bat species) carry
fruits away from the parent tree to consume elsewhere, often dropping quantities of seeds below the feeding perch. Studies
show that not all frugivores in a given habitat are equally attracted to the same fruit species (Snow, 1962; McDiarmid et al.
1977; Howe and de Stevens, 1979; Milton, 1981; Chen et al., 2001; Herrera, 2002; Dew, 2005). Fruit choices at times
appear deliberate in that individuals of a particular species will ignore available ripe fruits of one tree species while feed-

ing heavily on ripe fruits from another. Such behavioral
disparities may influence patterns of seed distribution and
thereby affect forest structure and composition (Clark et al.

Considerable effort has been devoted to examination of
the attractant relationship between fruit species and their
dispersal agents, with particular attention paid to avian dis-
persers. Fruit traits examined include fruit size, color, hard-
ness, nutrient composition, seed size and number, second-
ary metabolites, distribution patterns in space and time,
and accessibility, among others (Denslow and Moermond,
1982; Howe, 1993; Fleming et al., 1993; Corlett, 1996;
Alves-Costa and Lopez, 2001; Herrera, 2002). These traits
must then be integrated with features of the external and
internal morphology and physiology of potential dispersal
agents (Milton, 1981; Witmer and Van Soest, 1989; Stiles,
1993; Alves-Costa and Lopez, 2001; Levey and Martinez
del Rio, 2001; Martinez del Rio and Karasov, 1991). Due
to this complex array of factors, many questions related to
seed-disperser interactions remain unanswered or incom-
pletely understood as information to address them is scarce,
inconclusive or simply non-existent (Herrera, 2002).

As the edible pulp or aril is the "reward" typically offered
to dispersers, its nutrient value would seem to be a critical
element in plant-disperser interactions (e. g., Stiles, 1993,
p. 228: "Birds eat fruits to gain the nutritional rewards in
fruit pulp. The preferences exhibited by birds must, in
part, be influenced by the nutritional components assimi-
lated.") (see also Herrera, 2002). Yet to date, in spite of
numerous elegant papers on aspects of fruit selection and
seed dispersal by a wide range of dispersal agents, there
is little comparative data on the nutrient composition of
a representative array of mammal-consumed wild fruits
from tropical forest sites (Howe, 1993). This absence is
particularly striking for Barro Colorado Island (BCI) in
the Republic of Panama, as this 1600 ha nature reserve has
been the site of numerous studies of mammalian dietary
ecology and there is also an unusual wealth of information
on BCI forest composition and phenology and the mainte-
nance of forest diversity (Knight, 1975; Leigh et al., 1982;
Hubbell and Foster, 1990; Milton, 1991).

Here I present data on the macronutrient content of 19
species of wild fruits collected on Barro Colorado Island.
All species have seeds dispersed by various mammals as well
as other vertebrates and/or invertebrates. Unlike many
studies, fruit species analyzed were not selected because of
the dietary focus of any particular species-rather they are
regarded as a composite sample of macronutrient patterns
of mammal-consumed fruits from the BCI forest. Results
of analyses provide a quantitative basis for characterizing
fruit nutrient profiles for fruits at this site. Similar infor-
mation from a number of Neotropical lowland forest sites
might prove useful in refining hypotheses related to fruit-
disperser relationships, niche-partitioning and the pre-
dicted composition of particular disperser communities as

Neotropical Primates 15(1), January 2008
well as the role played by primate and other mammalian
dispersers in influencing and maintaining forest composi-


Study site
Fruits were collected from adult trees, hemiepiphytes or
lianas growing in the forest on Barro Colorado Island,
Panama. Detailed descriptions of this moist lowland
tropical forest and of climatic features characteristic of this
region can be found in the literature (Allee, 1926; Croat,
1978; Leigh et al., 1982; Hubbell and Foster, 1990).

Study species
The 19 fruit species analyzed were not collected according
to any predetermined plan and can be regarded as an arbi-
trary sample of edible fruit flesh from this forest. The only
criterion for acceptance was that each species produce fruit
flesh of a type generally accepted as functioning to attract
mammalian feeders (van der Pijl, 1969) and known to be
consumed by at least two of the four non-human primate
species on BCI. I typically noted a particular fruiting tree
because some mammal, often a howler monkey (Alouatta
palliata) or spider monkey (Ateles .. o ..' i. was feeding on
its fruits. If sufficient fruit could be obtained for analysis, I
collected a sample. Each sample was composed of fresh, ripe,
undamaged fruit dropped under the parent tree by feeding
monkeys or removed by a climber with a tree pruner.

Ultimately fruits were collected from four hemiepiphytes
(Ficus .. . E costaricana, E obtusifolia, E trigonata), one
liana (Doliocarpus major) and 14 tree species (see Table 1
for list of all species). Four of the 14 tree species are among
the 50 most abundant species in the 50-hectare survey plot
of Hubbell and Foster (Hubbell and Foster, 1990; S. Lao,
personal communication). Six of the fruit species ana-
lyzed are members of the genus Ficus; there are 17 Ficus
species identified thus far on Barro Colorado Island and
one to three species of the other genera represented in my
sample (Wendeln et al., 2000; S. Lao, personal commu-
nication). Wendeln et al. (2000) examined the nutrient
content of 14 species of Ficus fruit from BCI, including the
six species I analyzed. Their work looked at Ficus species
only. The array of families and genera represented in my
study extend their findings on Ficus into a broader com-
parative framework. Fruits analyzed ranged from -1 cm
in diameter when ripe (E costaricana) to -9 cm in diameter
(Gustavia superba). Most species produced fruits -2-3 cm
in diameter and -3-5 cm. in length. The Ficus species con-
tained many dozens of tiny seeds; Gustavia superba fruit
contained an average of 12 seeds per fruit, each seed -2.5
cm in length. The rest of the species contained one to a few
moderate sized (-0.5-1.5 cm long) seeds per fruit.

Treatment of samples
Each sample was composed of the flesh or arils of various
fruits collected at the same time from a single parent tree.

Neotropical Primates 15(1), January 2008

Ripe fruits of a given species can show notable differences
in nutrient composition between trees or even when col-
lected from the same tree on the same or different days
(Howe, 1980; Wheelwright, 1993; Chapman et al., 2003).
Though the exact percentage of protein, fat or carbohy-
drate in a given ripe fruit may vary within or between trees
of a particular species, the profile of the three macronu-
trient groups in relation to one another should generally
remain consistent. In other words, one would not expect
to find that ripe fruits from tree 1 of species X at a given site
would predominate in protein while ripe fruits of tree 2 of
species X at that same site would predominate in fats. For
this reason, analytical results presented here are regarded
as a composite overview of the distribution profile of the
three macronutrient groups both for ripe fruits from that
tree and for ripe fruits of that species on BCI. Fruits of
all species except Ficus spp. were opened shortly after col-
lection and the flesh or aril separated from the seeds. The
Ficus fruits were broken apart to facilitate drying but seeds
were not removed (more on this point below). Samples
were dried in an oven at 55 C until a constant weight was
reached. The dried flesh was then ground in a Wiley mill
using a 20-mesh screen. Samples were stored in sealed
plastic containers placed in a desiccator and later analyzed
for crude protein, crude fat and total nonstructural carbo-
hydrates (TNC) at the Palmer Research Laboratory, Uni-
versity of Alaska, using techniques described below.

Crude protein
Total nitrogen (N) in each sample was determined by the
macro-Kjeldahl technique. Results were then multiplied

by the standard 6.25 conversion factor to get an estimate
of crude protein present in each sample.

Crude fat
The crude fat (= all fats, oils and waxes) content of each
sample was determined by the Randall extractor method
(Randall 1974). In this technique each sample is immersed
in hot ethyl ether for ten minutes, rinsed out of the solvent
and further extracted and rinsed with solvent condensate
for 20 minutes. The ethyl ether is then removed by evapo-
ration and the residue is weighed. The difference in weight
of the sample prior to and following ether extraction is used
as an estimate of the crude fat content of the sample.

Total nonstructural carbohydrates
The modified Weinmann technique as described in Milton
(1979) was used to determine the total nonstructural car-
bohydrate (TNC = sugars, starches and fructosans) content
of each sample.

Results and discussion

Results of analyses are presented in Table 1. All values are
expressed as percent dry weight of fruit pulp. Results are
discussed first in terms of each macronutrient class and then
in terms of the proportion contributed per class per species.

Protein content of the flesh or arils of the 19 species ranged
from a low of 3.2% (Tetragastris panamensis) to a high of
15.2% (Gustavia superba). Mean protein content for the

Table 1. The percent dry weight of each macronutrient group (crude protein, crude fat, total nonstructural carbohydrates, TNC) in flesh
or arils of 19 Panamanian fruit species.
Family Species Crude Protein Crude Fat TNC
Anacardiaceae Spondias mombin 4.3 1.3 40.0
Anacardiaceae Spondias radlkofera 11.7 3.9 24.6
Bombacaceae Quararibea asterolepis 5.4 0.2 31.0
Burseraceae Tetragastris panamensis 3.2 0.2 56.3
Dilleniaceae Doliocarpus major 4.5 3.8 21.1
Lauraceae Beilschmiedia pendula 6.2 25.4 11.9
Lecythidaceae Gustavia superba 15.2 42.3 5.1
Meliaceae Trichilea tuberculata 7.8 38.3 15.6
Moraceae Brosimum alicastrum 9.3 1.2 20.7
Moraceae Ficus bullenei 7.1 3.4 3.7
Moraceae Ficus costaricana 6.9 3.9 6.4
Moraceae Ficus insipida 7.0 5.8 14.5
Moraceae Ficus obtusifolia 4.1 3.6 8.7
Moraceae Ficus trigonata 5.6 6.4 10.5
Moraceae Ficus yoponensis 7.5 6.0 11.3
Myristicaceae Virola nobilis 4.5 42.5 18.6
Palmae Astrocaryum standleyanum 4.8 1.4 45.5
Palmae Scheelea zonensis 3.6 22.3 15.1
Rubiaceae Faramea occidentalis 4.1 0.1 38.8

combined samples was 6.5% 3.0%. This is a low protein
content when compared with that of tree leaves from this
same forest. On Barro Colorado Island, young tree leaves
averaged 19.5% 7.2% protein (Kjeldahl technique; n =
6 species; K. M., unpubl. data; see also Milton 1979 for
estimates of leaf protein using summed amino acid tech-
nique). Data compiled by Leung (1969) show that insects
(i. e., flying ants, beetles, crickets, caterpillars, grasshop-
pers, locusts and termites) average 16.2% 8.5% protein.
Some insect larvae contain > 42% protein (Jenkins and
Milton, 1993). When compared both to young leaves
and insects, most ripe fruits are low in protein. The actual
protein content of fruits analyzed may be even lower than
estimates suggest as using nitrogen as a proxy for protein
may over-estimate the protein content of fruit pulp (Levey
and Martinez del Rio, 2001).

The crude fat content of the 19 species ranged from a low
of 0.1% (Faramea occidentalis) to a high of 54.2% (Virola
surinamensis). Mean fat content for the combined samples
was 11.8% 16.5%. Though most species were decidedly
low in fat, five species contained a substantial (> 22% dry
wt.) amount (Table 1). Leaves are generally quite low in
fat. Young leaves from the Barro Colorado forest average
2.4% 2.3% crude fat (Randall technique, n = 10 species;
K. M., unpubl. data). Insects examined by Leung (1969,
see list above) averaged 9.5% 9.9% fat. Some insect
larvae may have fat contents > 42% (Jenkins and Milton,
1993). Though most fruit species in the BCI sample were
not high in fat, a few species were notably high.

Total nonstructural carbohydrates
The total nonstructural carbohydrate content (TNC) of the
19 species ranged from a low of 5.1% (Gustavia superba)
to a high of 56.3% (Tetragastrispanamensis). Mean TNC
content for the combined samples was 21.0% 14.8%.
Leaves are generally low in TNC. Mature leaves from the
Barro Colorado forest average 3.7% 3.5% TNC and
young leaves 4.0% 2.2% (Weinmann technique; n = 6
species for each category; Milton, 1979). Data from Leung
(1969) show that insects average 2.7% 1.6% TNC.
Therefore, in contrast to both leaves and insects, many ripe
fruit species contain a notable amount of nonstructural

Nutrient content by species
Though all species contained some measurable proportion
of each macronutrient class, there were often striking dif-
ferences in the amount contributed per class per species.
In terms of the three nutrient classes, two species (10.5%
of the total sample) were highest in protein, five (26%)
were highest in fats and twelve (63%) were highest in non-
structural carbohydrates. Species predominating in TNC
showed a wide range of values. In contrast, results sug-
gest that when a given plant taxon uses fat as the principal
caloric attractant, it will use a high percentage, as all five
species in this category had > 22% fat. Gustavia superba,

Neotropical Primates 15(1), January 2008
with a fat content of 42.3%, is clearly in the high fat group
but unlike the other four species predominating in fat,
Gustavia had the highest protein content (15.2%) and
one of the two lowest TNC contents in the entire sample.
Therefore, it was an anomalous species in comparison with
the other eighteen and has its own distinct pattern of nutri-
ent allocation. The two species predominating in protein
were both Ficus species. It is likely that the protein content
of the flesh of all Ficus species was confounded by the pro-
tein content of the many indigestible fig seeds in each fruit
and for this reason overestimated. Fig seeds are degraded
by chemicals used in analyses but, if swallowed intact, (that
is, not crushed by the teeth of the feeder) apparently are
not degraded by the action of digestive enzymes in the guts
of most fig-eating animals and are excreted intact (Hladik
and Hladik, 1969; Morrison, 1980). On BCI, Ficus fruits
are made up of approximately 50% pulp and 50% seeds
dry weight and the two components show somewhat dif-
ferent nutrient profiles, with flesh higher in water-soluble
carbohydrates than seeds and seeds higher in protein and
fiber than flesh (Wendeln et al., 2001).

Howler monkeys and some other fig-eating mammals crush
some, but by no means all, fig seeds when eating the fruits
(Hladik and Hladik, 1969; Wendeln et al., 2001; K.M.,
pers. obs.). For this reason, it is difficult to provide an esti-
mate of the proportion of pulp versus seed nutrient intake
for many fig-eating animals. All values presented in Table
1 for Ficus species include both flesh and seeds; the precise
nutrient contribution of one or the other component in
a given animal's diet would depend on an array of factors
particular to that feeder. With one exception, the TNC
content of E insipida, in comparison to all other species
in the analyses, Ficus species were not outstandingly high
in any nutrient group in spite of the fact that all estimates
likely include augmentation from seed macronutrients.

The nutrient patterns present in fruit species from particu-
lar locales presumably reflect a long evolutionary interplay
between tree species and seed dispersal agents (Snow, 1971;
McKey, 1975; Howe and Estabrook, 1977; Howe, 1993;
Herrera, 2002). The manufacture of fruit flesh is costly
and theoretically each parent tree is seeking to maximize
returns from such investments. If we view these 19 fruit
species as a type of "grab sample" of macronutrient pat-
terns present in vertebrate-consumed fruit species in this
forest, it is clear that the majority of species predominate
in nonstructural carbohydrates. In terms of caloric value,
TNC may not always dominate per unit mass ingested, but
it predominates as a chemical constituent in terms of the
three macronutrient groups. If we assume that, over evo-
lutionary time, each plant species has been free to elaborate
fruit nutrients in whatever proportions seem most expedi-
ent for seed dispersal success, it is clear that most species in
this sample have "chosen" to emphasize TNC over protein
or fat. A priori, there is apparently no physiological reason
why all fruit flesh could not be high in protein or fat rather
than TNC if either of these were more favored mode of

Neotropical Primates 15(1), January 2008

nutrient allocation, since there are fruit species, both on
BCI and elsewhere, showing such patterns (Snow, 1962;
McDiarmid etal., 1977; Styles, 1993; Herrera, 2002). On
a community-wide basis, soluble carbohydrates appear to
be the most commonly utilized disperser attractant.

The dominance of nonstructural carbohydrates in wild fruit
flesh is seen in data from other Neotropical locales. For
example, in a list of 40 wild fruit species eaten by frugi-
vores in Guatemala, 38 were highest in TNC and two in fat
(Coelho et al., 1976). Similarly, in most Neotropical plant
communities for which data are available, lipids appear
restricted to a small subset of species producing fleshy fruits.
For example, only 20% of the fleshy-fruited species in a
Costa Rican plant community had a lipid content > 10%
dry weight (Stiles 1993). Protein content appears low for
most wild fruit species, regardless of locale. A mean protein
content of 8.9% 0.6% was reported for 122 fruit species
in the Neotropics while an overall mean protein content of
5.5% 0.4% was reported for fruit species in Europe and
5.2% 0.4% for species in North America (Stiles 1993).
Several reasons can be suggested for the use of nonstructural
carbohydrates rather than protein or fat as the predominant
fruit attractant. Generally, nonstructural carbohydrates are
the most rapidly and inexpensively digested of the three
nutrient groups (Maynard and Loosli, 1969). This factor
may be of special importance to a wide array of smaller-
bodied frugivores, since they need to turn over consider-
able food energy each day to meet metabolic demands. In
herbivorous mammals, as body size decreases, metabolic
costs per unit mass increase exponentially while gut capacity
increases proportionate to body mass (Parra, 1978). Thus,
in general, smaller homeotherms should be greatly attracted
to foods offering high and rapid energetic returns.

Flesh from six of the 19 fruit species used in these analy-
ses, and likewise collected by KM on BCI, were analyzed
for sugar composition by I. and H. Baker, using meth-
odologies presented by Baker and colleagues (1998). In
all cases (100%) fruit flesh predominated in glucose and
fructose and sucrose was uniformly low (Milton, 1999).
In striking contrast, data on the sugar composition of a
wide range of cultivated fruits show that sucrose is gener-
ally the predominant fruit sugar (Milton, 1999). Sugar
composition of flesh could have important implications
for the fruit preferences of particular seed dispersal agents
(Freeman and Worthington, 1989; Ko et al., 1998; Mar-
tinez del Rio et al., 1989; Martinez and Karasov, 1991).
The digestion of the hexoses glucose and fructose does not
require manufacture of a sucrase, necessary if sucrose were
the principal sugar attractant. This metabolic "savings"
may benefit many frugivores, particularly smaller bodied
frugivores. Fats typically take considerably longer to digest
than nonstructural carbohydrates and are more costly to
digest in terms of their specific dynamic effect. In addi-
tion, most animals find it necessary to maintain a balance
between energy and protein in the diet. The source of the
energetic calories can affect the amount of protein required

to maintain this balance. Experimental data show that for
each 4% of extra fat input into the diet, the protein content
must go up 1% by weight to maintain a constant protein
to non-protein caloric ratio (Crampton and Lloyd, 1959).
A certain threshold in body size and/or digestive specializa-
tions may be required to utilize quantities of fatty foods
efficiently (Stiles, 1993; Martinez del Rio and Karasov,
1990; Levey and Martinez del Rio, 2001; Dew, 2005).

As noted, there are already various sources of high qual-
ity protein available in tropical forests, including young
leaves, insects and other animal matter. For this reason,
tree species producing protein-rich flesh would not be
offering a particularly novel attractant. As most vertebrates
do not require large quantities of protein in the diet each
day, smaller-bodied species in particular might rapidly
become satiated on protein-rich flesh which would lower
the number of fruits eaten per visit and thereby depress
the number of seeds dispersed. Protein ingested in excess
of what the body actually requires also tends to be wasted
in so far as its specific functions are concerned, since only
the most marginal amounts can be stored. And dietary
protein must be catabolized, a process that is energetically
expensive because of the high specific dynamic effect of
protein digestion (Maynard and Loosli, 1969). The strik-
ing exception to the low protein pattern in the BCI sample
was Gustavia superba. It was by far the largest fruit in the
sample-softball-sized- and its unique nutrient profile
(and large seeds) suggests that it may seek to attract large
terrestrial frugivores as seed dispersal agents. Offering
notable protein and energy as a reward may help ensure the
specialized dispersal services required for unusually large
seeds. Fruits rarely comprise the entire diet of mammals
and birds, as fruit pulp provides primarily energy (Rode
and Robbins, 2000). Not only do fruits generally lack a
sufficiency of protein and various other nutrients animals
require, a high carbohydrate-low protein intake is not
desirable over the long term as it greatly elevates energy
metabolism (Rode and Robbins, 2000). To obtain an
optimal balance of energy and nutrients, most vertebrates
must therefore move away from fruiting trees each day to
seek out other types of foods. A fruit species that provided
feeders with complete nutrition would likely disperse few
seeds as animals would cluster around such trees until the
fruit crop was exhausted. In the case of Gustavia superba,
a high protein content might not be disadvantageous in
terms of seed dispersal, as trees of this species produce only
a few ripe fruits at any given time and all trees are small
in size. Large frugivores would thus be forced to move
about the forest each day to obtain sufficient food and in
doing so would move Gustavia seeds away from the parent.
Rodent predation on Gustavia seeds is high at some sites
(Sork 1987) and for this reason, movement of seeds well
away from parent trees may be of particular value.
In choosing to emphasize TNC in fruit flesh, species may
also be seeking to maximize their possibilities for future
reproductive success. As no plant can predict where a
given seed will land, it seems most expedient to use the

disperser attractant with the widest appeal. If seed disper-
sal agents present in the parent habitat are absent in new
habitats where seeds arrive and germinate, other frugivo-
rous species should still be attracted to sugar-rich fruits.
This might not happen so easily with a high protein or fat
reward as most dispersal agents depend largely on other
sources (leaves, insects, larvae, other vertebrates) for these
nutrients. It therefore appears that on BCI, nonstructural
carbohydrate is the most "popular" reward offered seed dis-
persal agents. By providing a source of readily assimilated
energy, largely in the form of fructose and glucose, sugary
flesh gives dispersal agents the fuel required for an active
life style that should facilitate the movement of seeds away
from the parent plant. Sugary fruits also give dispersers
the energy needed to move about the forest in search of
foods containing other nutrients they require. Many pri-
mates preferentially eat fruits early in the day and leaves
later in the day (Chivers, 1977; Milton, 1980). This feed-
ing pattern suggests that many frugivores actively seek out
energy-rich food sources at their first opportunity and feed
heavily from them in order to ensure the energy required
for their other daily activities. Such a feeding pattern, in
turn, should enhance seed dispersal efficiency.

As noted, the six Ficus species in this sample were not nota-
bly high in any nutrient group. In keeping with this find-
ing, data suggest that most Neotropical frugivores are not
attracted to fig fruits because of their high nutrient value.
On BCI, spider monkeys (Ateles ._ ..".. for example, will
ignore fruiting fig trees if other more preferred fruit species
are available (Milton, pers. obs.). Neotropical Ficus species
often produce large ripe fruit crops at times of year when
many other fruit species are in short supply (Foster, 1978;
Milton, 1980; Terborgh, 1983; Milton, 1993). It would
appear that many Neotropical frugivores are attracted to
Ficus fruit largely because it is the "only game in town" at
some times of year (Terborgh, 1983). Various Neotropical
mammals that show strong dependence on fig fruits
throughout the year appear to have either an energetically
inexpensive life style (e.g., howler monkeys, Alouatta pal-
liata; Milton, 1980) or a technique for eating fig fruits such
that nutrient returns are maximized and the intake of bulky
indigestible material is kept low (e.g., fruit bats; Artibeus
jamaicensis, Morrison, 1980; Wendeln et al., 2001).


Chemical analyses presented in this paper were carried out
by the Palmer Plant and Soil Analysis Laboratory, Alaska
Agricultural Experiment Station under the supervision of
Gary Michaelson. I thank David Rhoades for suggesting
this laboratory and the Smithsonian Tropical Research
Institute for providing funds for the analyses. Robin
Foster, Carol Augspurger and Gene Hammell provided
useful comments on much of this material.

Neotropical Primates 15(1), January 2008


Allee, W. C. 1926. Measurements of environmental fac-
tors in the tropical rain forest of Panama. Ecology 7:
Alves-Costa, C. P. and Lopes, A. V. 2001. Using artificial
fruits to evaluate fruit selection by birds in the field. Bio-
tropica 33: 713-117.
Baker, H. G., Baker, I., and Hodges, S. A. 1998. Sugar
composition of nectars and fruits consumed by birds
and bats in the tropics and subtropics. Biotropica 30:
Chen, J., Deng, X.-B., Bai, Z.-L., Yang, Q., Chen, G.-Q.,
Liu, Y. and Liu, Z.-Q. 2001. Fruit characteristics and
Muntiacus muntijak vaginalis (Muntjac) visits to indi-
vidual plants of Choerospondias Biotropica 33:
Chivers, D. J. 1977. The feeding behaviour of siamang
(\, ' syndactylus). In: Primate ecology: studies of
feeding and ranging behaviour in lemurs, monkeys and apes,
T. H. Clutton-Brock (ed.), pp.255-382. Academic Press,
New York.
Clark, C. J., Poulsen, J. R. and Parker, V. T. 2001. The role
of arboreal seed dispersal groups on the seed rain of a
lowland tropical forest. Biotropica 33:606-620.
Corlett, R. T. 1996. Characteristics of vertebrate-dispersed
fruits in Hong Kong. J. Trop. Ecol. 9: 501-510.
Crampton, E. W. and L. E. Lloyd. 1959. Fundamentals of
nutrition. W. Freeman Co., San Francisco.
Croat, T. B. 1978. Flora of Barro Colorado Island. Stanford
University Press, Palo Alto, California.
Danish, L., Chapman, C. A., Hall, M. B., Rode, K. D.
and Worman, C. O'D. 2006. The role of sugar in diet
selection in redtail and red colobus monkeys. In: Feeding
ecology in apes and other primates, G. Hohmann, M. M.
Robbins and C. Boesch (eds.), pp.473-488. Cambridge
University Press, Cambridge.
Denslow, J. S. and Moermond, T. C. 1982. The effect of
accessibility on rates of fruit removal from tropical shrubs:
an experimental study. Oecologia 54: 170-176.
Dew, J. L. 2005. Foraging, food choice and food process-
ing by sympatric ripe-fruit specialists: Lagothrix lagothri-
chapoeppigii and Ateles belzebuth belzebuth. Int. J. Prima-
tol. 26: 1107-1135.
Fleming, T. H., Venable, D. L., and Herrera M., L. G.
1993. Opportunism vs. specialization: the evolution
of dispersal strategies in fleshy-fruited plants. Vegetatio
107/108: 107-120.
Foster, M. S. 1978. Total frugivory in tropical passerines: a
reappraisal. J. Trop. Ecol. 19: 131-154.
Freeman, C. E., Worthington, R. D. 1989. Is there a dif-
ference in the sugar composition of cultivated sweet fruits
of tropical/subtropical and temperate origins? Biotropica
21: 219-222.
Herrera, C. M. 2002. Seed dispersal by vertebrates. In:
Plant-animal Interactions, C. M. Herrera and 0. Pellmyr
(eds.), pp.185-208. Blackwell Science Ltd., Oxford.

Neotropical Primates 15(1), January 2008
Hiadik, A. and Hladik, C. M. 1969. Rapports trophiques
entire vegetation et primates dans la foret de Barro Colo-
rado (Panama). Terre et vie 1: 25-117.
Howe, H. E 1993. Specialized and generalized disper-
sal systems: Where does the paradigm stand? Vegetatio
107/108: 3-14.
Howe, H. F 1980. Monkey dispersal and waste of a
Neotropical fruit. Ecology 61: 944-959.
Howe, H. E and Estabrook, G. F 1977. On intraspecific
competition for avian dispersers in tropical trees. Am.
Nat. 111: 817-832.
Howe, H. E and de Steven, D. 1979. Fruit production, mi-
grant bird visitation, and seed dispersal of Guarea glabra
in Panama. Oecologica 39: 185-186.
Howe, H. F and Vande Kerckhove, G. A. 1979. Fecun-
dity and seed dispersal of a tropical tree. Ecology 60:
Hubbell, S. P. and Foster, R. B. 1990. Structure, dynamics
and equilibrium status of old-growth forest on Barro Col-
orado Island. In: Four Neotropical Rainforests, A. Gentry
(ed.), pp.522-541. Yale University Press, New Haven.
Izhaki, I. and Safriel, U. N. 1989. Why are there so few
exclusively frugivorous birds? Experiments on fruit diges-
tion. Oikos 54: 23-32.
Janzen, D. H. 1978. How to be a fig. Annual review of ecol-
ogy and systematics 10: 13-51.
Jenkins, C. and Milton, K. 1993. Diet and survival among
the Hagahai of Papua New Guinea. In: Tropical forests,
people and food, C. M. Hladik, A. Hladik, 0. F Linares,
H. Pagezy, A. Semple and M. Hadley (eds.), pp.281-293.
Man and the Biosphere Series, Vol. 13, pp.281-293.
Parthenon Pub. Group, Pearl River, New York.
Karasov, W. H. 1990. Digestion in birds: Chemical and
physiological determinants and ecological implications.
Studies in avian biology 13: 391-415.
Knight, D. H. 1975. A phytosociological analysis of species-
rich tropical forest on Barro Colorado Island, Panama.
Ecological monographs 45: 259-284.
Ko, I. W. P., Corlett, R. T. and Xu, R-J. 1998. Sugar com-
position of wild fruits in Hong-Kong, China. J. Trop.
Ecol. 14: 381-387.
Leigh E. G., Rand, A. S. and Windsor, D. M. 1982. Ecol-
ogy of a Tropical Forest. Smithsonian Press, Washington,
Levey, D. J. and Martinez Del Rio, C. 2001. It takes guts
(and more) to eat fruit; lessons from avian nutritional
ecology. Auk 118 : 819-831.
Leung, W. T. W. 1969. Food composition tables for use
in Africa. FAO-HEW, Public Health Service Publication.
Bethesda, Maryland.
Martinez del Rio, C., Karasov, W. H., and Levey, D. J.
1989. Physiological basis and ecological consequences of
sugar preferences in cedar waxwings. Auk 106: 64-71.
Martinez del Rio, C. and Karasov, W H. 1990. Digestion
strategies in nectar- and fruit-eating birds and the sugar
composition of plant rewards. Am. Nat. 136: 618- 637.
Maynard, L. A. and Loosli, J. K. 1969. Animal nutrition.
McGraw-Hill, New York.

McDiarmid, R. W., Ricklefs, R. E., and Foster, M. S. 1977.
Dispersal of Stemmadenia donnell-smithii (Apocynaceae)
by birds. Biotropica 9: 9-25.
McKey, D. 1975. The ecology of coevolved seed dispersal
systems. In: Coevolution of animals and plants, L. E. Gil-
bert and P. H. Raven, (eds.), pp.159-191. University of
Texas Press, Austin.
Milton, K. 1979. Factors influencing leaf choice by howler
monkeys: a test of some hypotheses of food selection by
generalist herbivores. Am. Nat. 114: 362-378.
Milton, K. 1980. The foraging strategy of howler mon-
keys: a study in primate economics. Columbia University
Press, New York.
Milton, K. 1981. Food choice and digestive strategies of
two sympatric primate species. Am. Nat. 117: 476-495.
Milton, K. 1991. Annual patterns of leaf and fruit pro-
duction by six Neotropical Moraceae species. J. ofEcol.
79: 1-26.
Milton, K. 1999. Nutritional characteristics of wild Pri-
mate foods; Do the natural diets of our closest living rela-
tives have lessons for us? Nutrition 15: 488-498.
Morrison, D. W. 1980. Efficiency of food utilization by
fruit bats. Oecologia 45: 270-273.
Parra. R. 1978. Comparison of foregut and hindgut fer-
mentation in herbivores. In: The ecology of arboreal fo-
livores, G. G. Montgomery (ed.), pp.205-230. Smithso-
nian Press, Washington, DC.
Pijl, L. van der. 1969. Principles of dispersal in higher
plants. Springer-Verlag, Berlin.
Randall, E. L. 1974. Improved method for fat and oil anal-
ysis by a new process of extraction. Journal of Association
of Official American Chemists 57: 1165-1168.
Snow, D. W. 1962. The natural history of the oil-bird
Steatornis caripensis in Trinidad W.I. Part 2. Population,
breeding ecology and food. Zoologica 47: 199-221.
Snow, B. K. 1971. Evolutionary aspects of fruit-eating by
birds. Ibis 113: 194- 202.
Stiles, E. W. 1993. The influence of pulp lipids on fruit
preferences by birds. Vegetatio 107/108: 227-235.
Wendeln, M. C., Runkle, J. R., and Kalko, E. K. V. 2000.
Nutritional value of 14 fig species and bat feeding prefer-
ences in Panama. Biotropica 32: 489- 501.
Witmer, M. C and Van Soest, P. J. 1998. Contrasting di-
gestive strategies of fruit-eating birds. Functional Ecology
12: 728-741.
Terborgh, J. 1983. Five new world primates: A study in
comparative ecology. Princeton University Press, Princ-
eton, N.J.
Wheelwright, N. T. 1993. Fruit size in a tropical tree spe-
cies: variation, preference by birds, and heritability. Veg-
etatio 107/108: 163-174.

Neotropical Primates 15(1), January 2008


Alfred L Rosenberger1, Luke J. Matthews2

'Department of A .-. '...:*..'.. Brooklyn College, CUNY Brooklyn, NY 11210, USA
'Department ofA .-'. .....'.. New York University, New York, NY 10003, USA


We summarize our re-examination and extension of the Groves (2001) parsimony analysis of Woolly monkeys, genus
Lagothrix, which led him to conclude that the species flavicauda is not most closely related to lagotricha but to Ateles, the
Spider monkeys. As a consequence, Groves further proposed that the Yellow-tailed woolly monkey should be assigned to a
separate genus, Oreonax, previously erected by Oldfield Thomas (1927). Our analysis, while closely following his methods,
samples a greater diversity of species and sub-species representing all the living ateline genera and makes minor revisions in
Groves' data matrix of craniodental characteristics. With this broader analysis we show that Groves' cladistic results cannot
be replicated except by duplicating his study using only a restricted range of taxa. A more wide ranging taxonomic sam-
pling fails to link consistentlyflavicauda and other ateline species, in any particular cladogram topology, while the overall
craniodental morphology of flavicauda does not separate it from lagotricha. Groves' cladistic conclusion is likely to be an
artifact stemming from a chance combination of the particular taxa used as a study group and selection of characters that
may not be appropriate in intergeneric comparison. There is thus no justification for recognizing Oreonax as a distinct
genus, and its usage should be rejected.

Key Words: Primates, Atelidae, Lagothrix, Oreonax, Lagothrix flavicauda, Yellow-tailed woolly monkey, cladistics, parsi-
mony, classification.


Presentamos abreviadamente nuestra re-examinaci6n y extension del anAlisis de parsimonia de Groves
(2001) de los monos lanudos, genero Lagothrix, que le llevaron a concluir que la especie flavicauda
no esta mias cercanamente relacionada con lagotricha pero si con Ateles, los monos arafia. En consecuencia, Groves propuso
que el mono lanudo de cola amarilla deberia ser asignado a un genero aparte, Oreonax, previamente erigido por Oldfield
Thomas (1927). Nuestro analisis cuidadosamente sigui6 sus m6todos, e incluye una mayor diversidad de species y sube-
species que representan todos los generos vivientes de Atelinos, y hace revisiones menores en la matriz de Groves de datos de
caracteristicas craneodentales. Con este mias amplio anAlisis mostramos que los resultados cladisticos de Groves no pueden
ser replicados except al duplicar su studio utilizando solamente un rango restringido de taxa. Un muestreo taxon6mico
de rango much mias amplio falla en articular consistentemente aflavicauda y otras species de atelinos, en cualquier cla-
dograma topol6gico en particular, mientras que la morfologia craneodental general deflavicauda no la separa de lagotricha.
La conclusion cladistica de Groves es probablemente una idea derivada de una combinaci6n azarosa de los taxa particulars
utilizados como un grupo y la selecci6n de caracteres que parecen no ser apropiados para las comparaciones entire generos.
Por lo tanto no hay justificaci6n para reconocer Oreonax como un genero distinto, y su uso deberia ser rechazado.

Palabras Clave: Primates, Atelidae, Lagothrix, Oreonax, Lagothrixflavicauda, mono lanudo de cola amarilla, cladistica,
parsimonia, clasificaci6n.

Introduction ture of platyrrhine biodiversity. Contemporaries of Hersh-
kovitz working under a similar paradigm, such as Napier
The alpha taxonomy of living New World monkeys is expe- and Napier (1967) and Napier (1976) who, like Groves,
riencing an extreme makeover. The turning point behind surveyed the systematics of all primates, recognized 64 and
this new perspective may be taken as the publication of 67 platyrrhine species, respectively. Groves recognizes 112.
Groves' (2001) book "Primate Taxonomy." After decades The difference is remarkable, and it has large implications
of consolidation and stability beginning in the late 1940s, for systematics, evolutionary theory and conservation. The
largely based on the contributions of Philip Hershkovitz, difference results largely from elevating subspecies-level taxa
recent treatments have promulgated quite a different pic- to full species rank. As far as we are aware, this movement

Neotropical Primates 15(1), January 2008

toward split taxonomies (i. e., as in the traditional distinc-
tion of lumperss" and splitterss") has not been scrutinized
scientifically by primatologists. Groves's platyrrhine classifi-
cation (2001, and references therein) is also a departure in
the number of modern platyrrhine genera recognized, three
more than the authoritative and widely accepted works of
the Napiers (Napier and Napier, 1967; Napier, 1976) and
Hershkovitz (e.g., 1977). One of these is Oreonax, the
genus which Groves (2001) revived for a species of Woolly
monkey, flavicauda, that had fallen into synonymy decades
ago. The purpose of this report is to explain that the analy-
sis which led Groves (2001) to raise the rank of the species
L. flavicauda to the genus level is flawed, making his phylo-
genetic interpretation and taxonomic decision unjustified.

Groves' study was based on an algorithmic cladistic analy-
sis of 20 craniodental characteristics involving five ate-
line species, one (none were identified by species name)
taken from each living genus Alouatta, Lagothrix, Ateles,
Brachyteles and a pair of species representing Lagothrix
(explicitly lagotricha and flavicauda). In a complimentary
analysis, he also added the Brazilian subfossils Caipora and

Protopithecus, but this aspect did not add meaningfully to
the results. The character state data was analyzed under
parsimony criteria using PAUP (Swofford, 1993). Groves
found that the species flavicauda was more closely related
to Ateles sp. than to L. lagotricha. Having thus discovered
that Lagothrix is non-monophyletic, Groves elected to
remove flavicauda to the genus Oreonax.


As explained more fully elsewhere (Matthews and Rosen-
berger, in press), we replicated Groves' (2001) study and
his results, and then extended his methodology to include
more taxa belonging to the genera he studied in order to
see if it produced a stable outcome that was consistent
with his findings. We also made slight modifications in
his dataset that are fully justified and explained in Mat-
thews and Rosenberger. The taxonomic distribution of the
original set of craniodental characters are listed in tabular
form in these sources. This direct test is a more challenging
one than a critique in which we would have developed an
entirely independent analysis, or if we had introduced new

Figure 1. Four different parsimonious cladograms (from Matthews and Rosenberger, in press) based on
the data and methods of Groves (2001), as explained in the text. All use five ateline species as the study
group, one each fim io .'i:.:. L:gothrix and Ateles and two for Lagothrix, flavicauda and one other. Fig.
1 a is a tree that we generated which replicates Groves's result, his basis for resurrectingflavicauda as Genus
Oreonax Thomas. The three alternatives, and several others (Matthews and Rosenberger, in press), were
obtained using different combinations of five species drawn from the four ateline genera. The absence of
a stable, replicable cladistic result indicates the Groves's tree is an artifact of taxonomic sampling.

types of data. We showed that a large variety of minimal-
length, incongruent parsimonious cladograms can be gen-
erated by varying the number and taxonomic composition
of species included as in-group taxa, the very ones whose
interrelationships are being tested.


There is no solid evidence that Lagothrix, as classically con-
stituted, is paraphyletic; there is no solid evidence thatfla-
vicauda is closer cladistically to Spider monkeys than to the
Common woolly monkey. Figure 1 shows a sample of cla-
dograms that we generated by analyzing the Groves (2001)
characters with a more recent version of PAUP* (Swof-
ford, 2002); additional examples appear in Matthews and
Rosenberger (in press). Several points need to be made: 1)
when replicating Groves' protocols as precisely as possible,
we obtain the same tree (Fig. 1A), 2) when adding new
ateline taxa to the sample, but still maintaining his 5 spe-
cies / 20 character matrix, the results diverge from Groves'
result and the monophyly and interrelationships of the ate-
line genera begin to vary in unstable ways (Fig. 1B-1D),
3) the driving force behind these variations appears to be
the taxonomic composition of the five species defined as
the in-group, or study group. Cladograms resembling the
outcome Groves found occurred only under specific condi-
tions, when they included L. lagotricha and Alouatta senicu-
lus in the study group. For example, when we substituted
L. lagotricha with another Woolly monkey taxon (poeppigii,
cana or lugens) in combination with flavicauda, the mono-
phyly of flavicauda and Ateles sp., the crux of Groves' new
findings, was broken.

Discussion and Conclusion

These results reflect an underappreciated shortcoming of
algorithmic parsimony analysis and possibly of cladistic
analyses in general (including likelihood and Bayesian
approaches), that topologies generated from some charac-
ter sets are highly sensitive to the taxonomic composition
of the study sample. Collins (2003) found essentially the
same thing in his molecular study of ateline intergeneric
relationships. The outcomes varied as he sampled more
and different species within the genera. Similar results were
also obtained by Sargis (2007) and Silcox (2007) using
morphological data in studies involving the relationships of
primates to other orders of mammals. Thus the constraints
that drive taxonomic sampling to skew cladistic analyses
with programs like PAUP appear to occur irrespective of
data type and taxonomic level.

Regarding Oreonax, we offer some additional thoughts
on why Groves' (2001) study may have been problematic
from the outset. First there is a practical issue in the nature
of the craniodental material Groves used that raises some
questions. Lagothrix flavicauda is poorly represented in
museum collections (Mittermeier et al., 1977). Groves'
samples of skulls were the only two specimens in the col-

Neotropical Primates 15(1), January 2008

elections of the American Museum of Natural History
(AMNH) and in the United States. They are curiously
different in morphology from the holotype and others in
the Natural History Museum (British Museum of Natural
History, London), a young adult male and female (Fig. 2)
plus a juvenile. The AMNH individuals are aged, with
very heavily worn teeth and some anatomical peculiarities
in jaw and cranial shape. Skins, with the typical flavicauda
yellow proximal tail, are associated with these and the
London specimens, so the species identifications are prob-
ably correct. However, unlike the AMNH material, the
London skulls do not differ much from samples of lagotri-
cha in craniodental anatomy. After consulting with Groves
(pers. comm.) some years ago regarding the morphology
of the AMNH specimens, we believed it would be wise
to split Lagothrix into two subgenera (see Hartwig et al.,
1996). Groves went a step further as a consequence of his
parsimony analysis. We can only offer that genus Lagothrix
may be somewhat variable craniodentally (as is Ateles, but
neither case is well documented). While this is not a sat-
isfactory status report on research materials or intrageneric
variation, we hope it encourages a more comprehensive
reinvestigation of the systematics of Lagothrix and study of
the full sample of cranial material, including the skulls in
Lima, Peru (Mittermeier et al., 1977).

Figure 2. Craniodental morphology of the (right) adult male,
the holotype, and (left) female specimen of Lagothrixflavicauda
in the Natural History Museum (BMNH). The morphology is
basically indistinguishable from L. lagotricha.

Neotropical Primates 15(1), January 2008

A second point relates to the data Groves (2001) employed.
As we discuss to some extent in Matthews and Rosenberger
(in press), several of his anatomical characters were prob-
lematic, or were coded in ambiguous ways. We tested this
by asking three other primate morphologists to score a
sample of ateline skulls using Groves' system, and used their
determinations in separate PAUP runs. All three produced
different cladograms. At another level, some of the features
Groves employed, whether or not they are well defined or
the characters states easily verified by others, would appear
to be low-level population variations that we think would
not be reliable as cladistic indicators at the generic level
(Matthews and Rosenberger, in press). Finally, we raise
a philosophical question. Systematists understand that
naming a genus unlike identifying and naming a species,
which has a biological reality is an artful practice, though
it is not without mutually agreed guidelines. We do not
believe Groves' (2001) decision to recognize Oreonax fol-
lowed standard practice, irrespective of the difficulties of
his parsimony analysis. Conventionally, living primate
genera are "defined" by morphology (implicitly or explic-
itly involving functional/adaptive uniqueness), often with
an emphasis on the skull and dentition (even if the taxon
is first identified by molecular means and cladistics). Dif-
ferential diagnoses of ateline genera are not plentiful, as
these formalistic descriptions are usually published only
as a part of broadly based, synthetic taxonomic revisions.
Napier (1976) and Groves (2001), for example, offer such
diagnoses and provide at least a brief paragraph of details
that allow for identification and diagnosis of each genus by
the skull and teeth. Napier (1976) also presents generically
distinctive features pertaining to body shape, limbs, hands
and feet, and external genitalia. It is worth noting that
although she had access to the type series Oldfield Thomas
(1927) used in naming Oreonax flavicauda when writing
her catalogue of primates in the Natural History Museum,
Napier retained this species in genus Lagothrix, citing the
authority of Fooden's (1963) revision.

Groves (2001) also provides apt craniodental and somatic
descriptions of Alouatta, Ateles, Brachyteles and Lagothrix.
However, he offers no similar differentia for Oreonax, no
anatomical description or list of features that either char-
acterizes Oreonax or distinguishes it from any other platyr-
rhines. In fact, we would argue that the description he
gives for his concept of Lagothrix perfectly encompasses
flavicauda as well as lagotricha. Instead of an explicit mor-
phological diagnosis, Groves (2001:194) presents the cla-
distic analysis of traits and concludes:
...the yellow-tailed woolly monkey should be separated
generically from Lagothrix, because there are no derived
character states that appear to unite them; and from all
other atelines, because the bootstrap value for the clade
uniting it with Ateles, its putative closest relative, is not
high. Its taxonomic position is isolated.

In other words, Groves' (2001) decision to separate flav-
icauda from Lagothrix is based on his inability to establish

the monophyly of Woolly monkey species on the basis of
his characters and method. However, by the same logic,
because his study was so limited in scope, it could also
be argued that the one unidentified species of Ateles that
grouped with flavicauda should also thus be a candidate
for generic status, either monotypically or grouped with
flavicauda. Applying Groves' logic to our complimentary
multi-species study would suggest that numerous new
genera ought to be erected, since in many cases the clado-
grams (Fig. 1) yielded novel clusters and irresolute branch-
ing sequences. Also, the "isolated" status of the flavicauda
linkage, which Groves evidences by low PAUP bootstrap
values, has no biological or phylogenetic meaning. The low
numerical value means that the characters analyzed do not
consistently link flavicauda with Ateles sp., more probably
because the linkage is wrong or the evidence for it is weak;
not because it is sending a biological signal of uniqueness
that has classificatory relevance. Essentially, flavicauda has
been misclassified because a heuristic measure of statistical
support has been misconstrued as a biological and phylo-
genetic characteristic.

The reality is that among all the modern platyrrhines there
is no mistaking the dense, woolly brownish coat, stocky
body build, thick-looking limbs and tail, and naked hel-
meted face of a Woolly Monkey from any other platyr-
rhine. These same characteristics make it hard to tell apart
flavicauda from lagotricha unless one examines the under-
side of the tail and pubic region, which tends to be a light
golden brown inflavicauda. It has yet to be proven thatfla-
vicauda differs in biologically meaningful ways from lagot-
richa that would justify separating it at the genus level. We
also wonder if the small allopatric population of Yellow-
tailed woolly monkeys can even be shown convincingly
to be a different species from its congeners using modern
morphological and molecular approaches. We submit that
there is no rationale for dividing Woolly monkeys into two
genera, Lagothrix and Oreonax, and the notion should be


Financial support was provided by the Tow Faculty Travel
Fellowship, Brooklyn College (City University of New
York) and through a PSC CUNY Research Award to ALR.
Institutional support for this research was provided by
the New York Consortium in Evolutionary Primatology


Collins, A. C. 2003. Atelinae phylogenetic relationships:
The trichotomy revived? Am. J. Phys. Anthropol. 124:
Fooden, J. 1963. A revision of the woolly monkeys (genus
Lagothrix). J. Mammal. 44: 213-247.
Groves, C. 2001. Primate Taxonomy. Smithsonian Institu-
tion Press, Washington DC.

Neotropical Primates 15(1), January 2008

Hartwig, W. C., Rosenberger, A. L., Garber, P. A. and Nor-
conk, M. A. 1996. On atelines. In: Adaptive Radiations of
Neotropical Primates, M. A. Norconk, A. L. Rosenberger
and P. A. Garber (eds.), pp.427-431. Plenum Press, New
Hershkovitz, P. 1977. Living New World Monkeys (Platyr-
rhini), with an Introduction to Primates. Vol. 1. University
of Chicago Press, Chicago.
Matthews, L. J. and Rosenberger, A. L. (in press). An
object lesson for primate systematics: parsimony analysis
(PAUP*) and the taxonomy of the yellow-tailed woolly
monkey, Lagothrix flavicauda. Am. J. Phys. Anthropol.
Mittermeier, R. A., de Macedo-Ruiz, H., Luscombe, B.
A. and Cassidy, J. 1977. Rediscovery and conservation
of the Peruvian yellow-tailed woolly monkey (Lagothrix
flavicauda). In: Primate Conservation, Prince Rainier III
and G. H. Bourne (eds.), pp.95-116. Academic Press,
New York.
Napier, J. R. and Napier, P. H. 1967. A Handbook ofLiving
Primates. Academic Press, London.
Napier, P. H. 1976. Catalogue of Primates in the British
Museum (Natural History). Part I: Families Callitrichi-
dae and Cebidae. British Museum (Natural History),
Sargis, E. J. 2007. The postcranial morphology of Ptilocer-
cus lowii (Scandenti, Tupaiidae) and its implications for
primate supraordinal relationships. In: Primate Origins,
M. J. Ravosa and M. Dagosto (eds.), pp.51-82. Springer,
New York.
Silcox, M. T. 2007. Primate taxonomy, Plesiadapiformes,
and approaches to primate origins. In: Primate Ori-
gins, M. J. Ravosa and M. Dagosto (eds.), pp.143-178.
Springer, New York.
Swofford, D. L. 1993. PAUP: f' ...-. analysis using
parsimony. Version 3.1.1. Smithsonian Institution Press,
Washington DC.
Swofford, D. L. 2002. PAUP*: .' '...-. analysis using
parsimony (*and other methods). Version 4.0 lOb. Sinauer
Associates, Sunderland MA.
Thomas, 0. 1927. A remarkable new monkey from Peru.
Ann. Mag. Nat. Hist. 19: 156-157.

Neotropical Primates 15(1), January 2008


Amira Elvia Apaza Quevedo1, Luis Fernando Pacheco2'3, Alejandra Irene Roldin3 y Maria Sol Aguilar

'INPA (Instituto Nacional de Pesquisas da Amazonia), PDBFF (Projeto Dindmica Bioldgica de Fragmentos Florestais), Av.
AndreAratjo, 2936-Petripolis, CEP 69083-000, Manaus, Brasil. amiraelvia@yahoo.es
2 Centro de Postgrado en Ecologiay Conservacidn, Instituto de Ecologia, Universidad Mayor de San Andres, Casilla 10077,
Correo Central, La Paz, Bolivia
3 Centro de Estudios en Biologia Tedrica y Aplicada (BIOTA), Avenida Aranjuez, Condominio Los Sauces de Aranjuez No. 2, La
Paz, Bolivia


Desde enero de 2000 a marzo de 2001 evaluamos aspects de la variaci6n temporal del comportamiento, dieta y tamano de
grupo de una poblaci6n de Ateles chamek en un bosque hdmedo montano de los Yungas Bolivianos (Estaci6n Biol6gica de
Tunquini EBT), asi como la disponibilidad de frutos consumidos por este primate (desde abril de 2000). Encontramos
correlaciones negatives entire las combinaciones del porcentaje de tiempo dedicado a actividades de descanso y desplaza-
miento y entire alimentaci6n y desplazamiento, pero ninguna correlaci6n entire las combinaciones de los porcentajes de
tiempo dedicado a otras actividades. La dieta de este primate fue altamente frugivora con un bajo consumo de hojas y otras
parties vegetables. Las species mis consumidas fueron Protium montanum, Casearia mariquitensis, Podocarpus sp., Anomos-
permum sp., species del gdnero Ficus, una especie indeterminada (nombre vernacular "canel6n") y species de la familiar
Lauraceae. La median mensual del nmero de individuos por grupo no present un pico evidence pero si una tendencia a
una relaci6n positive con la disponibilidad de frutos distintos al gdnero Ficus (agrupados como no-Ficus). En cuanto a la dis-
ponibilidad de frutos, se observ6 un incremento en la oferta de frutos no-Ficus hacia el inicio de la 6poca hdmeda, mientras
que la oferta de Ficus se di6 en todos los meses aunque de manera irregular. Finalmente, encontramos una relaci6n negative
entire la disponibilidad de frutos no-Ficus y el tiempo invertido en el desplazamiento.

Palabras Clave: Ateles chamek, dieta, disponibilidad de frutos, Estaci6n Biol6gica de Tunquini.


From January 2000 to March 2001 we evaluated aspects of temporal variation of activities, diet, and group size of an Ateles
chamek population in an area of humid mountain forest (Estaci6n Biol6gica Tunquini EBT) in the Bolivian Yungas. We
also evaluated (from April 2000) the availability of fruit species consumed by these primates. We found negative correlations
between the proportion of time dedicated to resting and traveling, as well as between feeding and traveling, but no correla-
tions were found between other combinations. The diet of these primates was highly frugivorous with a low consumption
of leaves and others items. The species most consumed were Protium montanum, Casearia mariquitensis, Podocarpus sp.,
Anomospermum sp., various species of the genus Ficus, an undetermined species (known by the common name "canel6n"),
and various species of the family Lauraceae. The monthly median of the number of individuals per group did not present
an evident peak, but did present a tendency towards a positive relation with non-Ficus fruits. Regarding fruit availability, we
observed an increase of non-Ficus species around the beginning of the wet season, whereas Ficus species had fruits available
during all months of the study. Finally, we found a negative relationship between the fruit availability of non-Ficus species
and the time spent in traveling.

Key Words: Ateles chamek, diet, fruit availability, Estaci6n Biol6gica de Tunquini.


Ateles chamek Humboldt se distribuye en bosques altos
amaz6nicos no perturbados de Brasil, Perd y Bolivia
(Anderson, 1997). Ateles tiene una dieta especializada
en frutos carnosos maduros y muchos aspects de su
ecologia se relacionan con la disponibilidad de los frutos
como recurso alimentario (Wallace, 1998). En Bolivia,
la mayoria de los studios sobre Ateles chamek han sido
desarrollados en bosques amaz6nicos y de pie de monte
(Pacheco y Simonetti, 1998, 2000; Wallace, 1998; Wal-
lace etal., 1998; Mendez, 1999; Tarifa, 2000; Palomeque,
2001; Aguilar, 2002). Estos studios reportan una dieta
basada principalmente en un gran ndmero de frutos
(Mendez, 1999; Palomeque, 2001; Aguilar, 2002) y una
fuerte dominancia de aquellos frutos que se encontraron
en mayor disponibilidad en su ambiente (Aguilar, 2002;
Wallace, 2005). En relaci6n a la asignaci6n de tiempo
a diferentes actividades del comportamiento de este pri-
mate, Wallace (2005) encuentra variaciones estacionales en
las actividades de movimiento, descanso y alimentaci6n,
estando esta lltima relacionada positivamente con el peso
de los frutos de las species mais dominantes. Tambidn
se indica que el tamafio de grupo en algunas poblaciones
de A. chamek tiende a disminuir o la observaci6n de los
grupos tiende a ser menos frecuente durante la 6poca seca
(Mendez 1999; Aguilar 2002). Esto posiblemente estd
relacionado con la disponibilidad de frutos como reportan
otros studios para el gdnero en Surinam y Perd (van Roos-
malen, 1985; Chapman, 1990).

En Bolivia A. chamek tambidn habitat bosques himedos
montanos y bosques nublados de Yungas, entire 1000 y 2700
msnm (Ribera, 1995; Rios, 1999); sin embargo no cono-
cemos studios anteriores sobre su ecologia en estos ambi-
entes. Los bosques himedos montanos se encuentran entire
los menos conocidos y mais amenazados del Neotr6pico
(Carrizosa, 1990; Dodson y Gentry, 1991; Henderson et
al., 1991) y estan siendo altamente fragmentados debido a
la expansion de actividades agricolas (Ribera, 1995), consti-
tuyendo ello una seria amenaza para las poblaciones de A.
chamek ya que esta especie no suele habitar ambientes per-
turbados (Johns y Skorupa, 1987; Wallace, 1998). Por otra
parte, Ateles es considerado un buen dispersor de semillas
debido a su dieta y fisiologia alimentaria y podria estar cum-
pliendo un rol important en la dinamica de regeneraci6n de
estos bosques, como ocurre en otros habitats (Klein y Klein,
1977; van Roosmalen, 1985; White, 1986; van Roosmalen
y Klein, 1988; Pacheco y Simonetti, 1998, 2000; Wallace,
1998; Aguilar, 2002).

El present studio da a conocer aspects sobre la variaci6n
mensual en 1) las actividades desarrolladas por A. chamek,
2) la composici6n de su dieta, 3) el tamafio de sus grupos,
y 4) la disponibilidad de frutos consumidos por Ateles, en
un bosque hilmedo montano de Yungas. Estos bosques
abarcan serranias de gran relieve topografico y caracteris-
ticas climiticas diferentes de otros bosques, por lo cual es

Neotropical Primates 15(1), January 2008

interesante evaluar aspects bisicos de la ecologia de Ateles
en estos ambientes, de forma que puedan ser comparados
con studios en tierras mris bajas.

Area de Estudio

El area de studio abarc6 aproximadamente 200 ha del cerro
Hornuni, entire 1,600 y 1,960 msnm, en los alrededores de
la Estaci6n Biol6gica Tunquini (EBT), Parque Nacional y
Area Natural de Manejo Integrado Cotapata (PNANMI
Cotapata), Bolivia (67043'- 6862'W, 16010'- 1620'S).
La topografla del lugar es muy abrupta con pendientes de
por lo menos 600 en algunas laderas. Los datos climiticos
de la EBT, registrados a 1,840 msnm indican una precipi-
taci6n annual de 2,500 mm y un promedio de temperature
mensual de 17.2 oC (Bach et al., 2003). La precipitaci6n
es mayor durante los meses de noviembre a abril, aunque se
present durante todos los meses del afio. La altura prome-
dio del dosel del bosque es de 25 m, con emergentes como
Ficus spp. y Aniba spp. que alcanzan hasta 30 m. Algu-
nas species representatives son: Hyeronima alchorneoides,
Morus insignis, Licania trianda, Meriania axinaeoides,
Spondias mombin, Hedyosmum angustifolium, Virola spp,
Inga spp. Ladenbergia spp. y Bathysa spp. El sotobosque
es dominado por la familiar Rubiaceae, destacindose Con-
daminea corymbosa, Faramea cf. multiflora, Ladenbergia
cf. multiflora y Palicourea aff. semirasa, ademis de species
de otras families como Piper spp., Allophylus punctatus,
Trichilia clausseni, Morus insignis, Miconia spp. Tambidn
en el sotobosque son abundantes dos species hemiepifitas:
Blechnum ensiforme y Philodendron ornatam (Ribera, 1995;
Bach et al., 2003; Paniagua et al., 2003).


El registro de actividades se llev6 a cabo durante 10 dias
por mes desde enero de 2000 hasta marzo de 2001 excepto
marzo y mayo de 2000). La disponibilidad de frutos se
registry entire abril de 2000 y marzo de 2001. Para detectar
grupos de Ateles se establecieron tres sendas perpendicu-
lares a la pendiente a 1,600, 1,700 y 1,900 msnm. Cada
senda tenia una longitud aproximada de 2 km y fue atrave-
sada por tres sendas de aproximadamente 300 m en el sen-
tido de la pendiente. La detecci6n de los grupos se realize
mediante contact visual, auditivo (vocalizaciones) y por
movimientos de las ramas durante recorridos diarios de
b6squeda intensive (desde las 7:30 a las 18:30) a lo largo de
las sendas. Consideramos como "grupo" a aquellos indi-
viduos observados en conjunto en cada encuentro; nues-
tras observaciones sugieren que todos los "grupos" prob-
ablemente pertenecian a una sola unidad social. En ese
sentido, nuestros "grupos" no representan unidades sociales
independientes y corresponderian a los subgrupos de otros
studios de estructura social de Ateles (Izawa et al., 1979;
Chapman, 1990; Wallace, 1998; Mendez, 1999).

Cada encuentro con un grupo de A. chamek constituy6
un period de observaci6n. Para el muestreo de compor-

Neotropical Primates 15(1), January 2008

tamiento se utiliz6 la tecnica de barrido (scan sampling,
Martin y Bateson, 1986), registrando en cada barrido el
ndmero de individuos observados realizando cada activi-
dad. Se hicieron barridos cada 5 minutes y cada period
de observaci6n fue mantenido mientras al menos un
individuo de Ateles permanecia a la vista. Las actividades
muestreadas fueron (modificado de Wallace, 1998): Movi-
miento: cuando el animal se desliza, balance, salta, sube,
desciende o trepa en las ramas de un mismo irbol o irboles
contiguos al arbol donde se realize el primer registro de un
period de observaci6n; Alimentaci6n: cuando el animal es
observado ingiriendo cualquier recurso alimenticio; Des-
canso: cuando el animal esti sentado o recostado y no se
mueve entire las ramas; Desplazamiento: cuando el animal
se mueve hacia y entire irboles lejanos al drbol done se
realize el primer registro de un period de observaci6n y
generalmente el desplazamiento es ripido, y Otros: cuando
el animal realize cualquier actividad fuera de las anterior-
mente mencionadas, tales como: acicalamiento, juego,
vocalizaciones, defecar, etc. Durante los registros de "ali-
mentaci6n" se anot6 tambidn el recurso consumido (flor,
fruto, hoja, brote y otros), nombre comin de la especie y,
siempre que fuera possible, se obtuvieron muestras bothni-
cas para su correspondiente identificaci6n en el Herbario
Nacional de Bolivia (LPB).

Para establecer la composici6n de la dieta se calcul6 el
porcentaje mensual de registros de alimentaci6n para dos
categories: frutos y no-frutos (cualquier otra parte vegetal
distinta de fruto). La categoria frutos incluy6 tres grupos
de species: 1) Ficus, 2) species 1, speciess con un minimo
de 10 registros) y 3) species 2 speciess con menos de 10
registros). Para el registro mensual de disponibilidad de
frutos se establecieron diez parcelas de 0,1 ha (10 x 100 m)
cada una, en ireas donde fueron observados los primeros
grupos de A. chamek, pero distribuidas de manera dispersa
en el irea de studio. En cada parcela se marcaron los irbo-
les con DAP > 10 cm y mediante el m6todo de estimaci6n
de abundancia visual del dosel (Chapman et al., 1992) se
realize un seguimiento mensual de la fenologia de frutos
maduros de cada individuo. Para esta estimaci6n se asign6
una categoria de abundancia de frutos maduros en la copa
de cada drbol segin la siguiente escala: 0 = 0%, 1 = 1 25%,
2 = 26 50%, 3 = 51 -75%, 4 = 76% 100%. Finalmente
se calcul6 la disponibilidad mensual de frutos maduros
por especie (D) mediante el indice propuesto por Wallace
(1998). Asi, para una especie dada:

D = (DAP x categoriafenologica)

Para el anAlisis de disponibilidad de frutos solamente se
utilizaron datos de species que estuvieron presents en
los registros de la dieta de A. chamek, excluyendo aquellas
reportadas por los guias locales pero sin evidencia en
campo, y s6lo se consideraron los arboles con DAP > al
menor DAP que present actividad reproductive dentro
de cada especie y durante el muestreo fenol6gico. Ficus

present una fenologia irregular a lo largo del ano y valores
muy altos de disponibilidad de frutos debido a la enorme
cantidad que produce en relaci6n a las otras species, lo
cual podria determinar el comportamiento fenol6gico de
la muestra en un anAlisis conjunto con las demis species,
enmascarando asi el comportamiento de estas 61timas. Por
tal raz6n se consider convenient analizar la disponibili-
dad mensual de frutos en dos grupos de species: a) espe-
cies que pertenecen al gdnero Ficus y b) species no-Ficus,
que incluyen al resto de las species.

Para expresar la variaci6n temporal mensual del compor-
tamiento de A. chamek se obtuvo el porcentaje de tiempo
dedicado a cada actividad en un mes dado asi:


(registros actividad i)
(registros todas las actividades)

x 100

i = movimiento, alimentacidn, descanso, desplazamiento y

Con los datos mensuales para cada actividad se realizaron
correlaciones de Spearman entire pares de variables de activi-
dad para determinar posibles relaciones entire ellas (Zar,

La variaci6n en el tamano de los grupos de A. chamek fue
reportada como la median y los cuartiles del ndmero de
individuos observados por grupo en cada mes. Para evaluar
la relaci6n entire la disponibilidad de frutos, la variaci6n
temporal del comportamiento y el tamano de grupo se
realizaron correlaciones de Spearman (Zar, 1996) entire
disponibilidad de frutos no-Ficus y el tiempo asignado a
cada actividad, asi como con el tamano de grupo desde
abril/2000. Se excluy6 de este analisis al grupo Ficus porque
present una fenologia poco sincr6nica a nivel poblacio-
nal (Milton et al., 1982; van Schaik, 1986; Ragusa-Netto,
2002), y por tanto, una ausencia de un pico fenol6gico
padr6n. Asi, un analisis de correlaci6n con nuestros datos
en esta variable, no produciria una correlaci6n o produciria
una correlaci6n ficticia.


Durante el studio se obtuvieron 78 encuentros periodss
de observaci6n) con grupos de Ateles, resultando en 1,068
barridos y 3,775 registros de organizaci6n temporal, para
un total de 89 horas con 45 minutes de observaci6n direct
(Fig. 1). Se encontr6 una correlaci6n negative entire los
porcentajes de tiempo asignados a alimentaci6n y a despla-
zamiento (R,= -0.7, n= 13, p= 0.02), asi como para la
combinaci6n entire desplazamiento y descanso (R= 0.65,
n= 13, p= 0.03). Los analisis de combinaciones entire las
demas actividades no presentaron relaciones estadistica-
mente significativas (Tabla 1 y Fig. 2).

Ateles chamek consumi6 por lo menos 29 species vege-
tales en la zona de studio con un 96.1 % de frutos y s6lo

Neotropical Primates 15(1), January 2008

un 3.9% correspondiente a hojas u otras parties vegetables
(Tabla 2). Se descartaron del anAlisis de la dieta 12 regis-
tros para los cuales no se pudo identificar la parte vegetal
ni la especie consumida. Es important resaltar que por lo
menos tres species de Ficus (E _1p,'wopl'llI, E trigona, E
cf. mathewsii) fueron consumidas casi todos los meses y que
en general, la dieta de Ateles estuvo dominada por pocas
species: Protium montanum, una especie indeterminada
de nombre comdn "Canel6n", Anomospermum sp., Case-
aria mariquitensis, mris de tres species dentro de la familiar
Lauraceae y dos del gdnero Podocarpus (Tabla 2).

No se observ6 un patr6n claro en el ndmero de individuos
por grupo a lo largo del studio (Fig. 3). Sin embargo,
existe una correlaci6n de tendencia positive (no signifi-
cativa estadisticamente) entire el tamafio de grupo y la
disponibilidad de frutos no-Ficus (R= 0.60; p= 0.08; n=
11; Tabla 1). Distintas species de Ficus presentaron domi-
nancia para la mayoria de los meses (Fig 4). Otras espe-
cies con frutos disponibles por largos periods de tiempo
pertenecen al gdnero Cecropia y diferentes species de la
familiar Lauraceae (Tabla 3). La disponibilidad de frutos
no-Ficus se increment6 hacia el period de transici6n entire
la 6poca seca y la hdmeda (septiembre-diciembre, Fig. 4),
la cual coincide con un mayor ndmero de species fruc-
tificando (Tabla 3). La disponibilidad de frutos no-Ficus
s6lo mostr6 una correlaci6n significativa con la actividad
de alimentacidn (R= 0.7, n= 11, p= 0.02; Tabla 1).

Tabla 1. Correlaciones de Spearman entire variables del compor-
tamiento de Ateles chamek, disponibilidad de frutos y tamafio de
grupo: a) entire disponibilidad de frutos no-Ficus y porcentaje
de tiempo asignado a cada actividad, b) entire los porcentajes de
tiempo asignado a cada actividad y c) entire disponibilidad de
frutos no-Ficus y ndmero de individuos por grupo.

Variables R p
no-Ficus alimentacidn -0.70 0.02
no-Ficus movimiento 0.15 0.65
no-Ficus descanso -0.45 0.17
no-Ficus otros 0.14 0.69
no-Ficus desplazamiento 0.45 0.16
alimentacidn movimiento -0.13 0.71
alimentacion descanso 0.43 0.19
alimentacidn otros -0.46 0.15
alimentacion desplazamiento -0.68 0.02
movimiento descanso -0.08 0.81
movimiento otros 0.19 0.57
movimiento desplazamiento -0.29 0.39
descanso otros -0.31 0.36
descanso desplazamiento -0.65 0.03
otros desplazamiento 0.19 0.57
no-Ficus tamano de grupo 0.60 0.08

E/00 F M A M J J A S 0 N D E/01 F M
M encuentros o barridos

Figura 1. Ndmero de encuentros y muestreos de barrido men-
suales obtenidos para Ateles chamek en la Estaci6n Biol6gica
de Tunquini (PN-ANMI Cotapata), Bolivia. En fondo gris
se representan meses en 6poca hdmeda y en blanco meses en
6poca seca.


. . ... . .

U movniento 0 ahmentaci6n S descanso Ef desplazamiento Z otros

Figura 2. Porcentaje de tiempo asignado por Ateles chamek
a cada actividad en la Estaci6n Biol6gica de Tunquini (PN-
ANMI Cotapata). Nimero de registros por mes: E=455,
F=36, A=313, J=350, J=44, A=194, S= 82, 0=106, N=427,
D=251, E=211, F= 118 y M= 1188. Areas en blanco indican
meses sin muestreo.


6 6*

E/00 F M A M J J A S O N D E/01 F M
Mediana [= 25%-75% = Mm-Max

Figura 3. Mediana mensual del tamafio de grupo de Ateles
chamek en la Estaci6n Biol6gica de Tunquini (PN-ANMI
Cotapata). En fondo gris se representan meses en 6poca
hdmeda y en blanco meses en 6poca seca.

o 0. -30

S A00 M J A S O N D F01 F M
i Ficus no-Ficus -- -x-- -desplazamiento -- alimentaci6n

Figura 4. Relaci6n entire la disponibilidad de frutos Ficusy no-Ficus con-
sumidos por Ateles chamek y las actividades de desplazamiento y aliment-
acidn, en la Estaci6n Biol6gica de Tunquini (PN-ANMI Cotapata). En
fondo gris se representan meses en 6poca hdmeda y en blanco meses en
6poca seca.

u -o

^0 U
U -o
0 U

u u

; U

o -o
* U

a, S

< N


g a

.2 .t


-C -
u u
M0 -o



U u

- S

U5 U

-o N

* -S

oU U

Neotropical Primates 15(1), January 2008


-c cc 00

-c f-
--. oO 0

uN 0
2 o ~

'C 0
c-i o ~

',D r-l c-q 2

N C c

0-0 *^-i
s i 0


CL C L C, LC, L mm mm


o== = = =_ o 0 0.
I- c' 3 ' 3 3 3 3 3


"-i.~-~- o >- E t .-c U
2 ,
-71 C- >4'
20^ on 2 ^,^ ^ iS

^ -&ii iii '
u uu u SS^ u ^^ uu -

U: -s-- s- 1 I
? 1 , i
^ ^ ^ ? S S ' tS ^ s ^ '

Neotropical Primates 15(1), January 2008



- CO

0 0 0 ^ 0 ^ 0 ^ 0 0 00
N C l


cc c c c6

0 ~~~~ ~ ~ [ 0 00 0'

oo f o
N1 r-q I


NO r-\ (Q

No N1 in it

-c C N c -c N 5 -

c C c c c c c o6

m rM m l\'S ^; n rM^ o

cc N c 0 [
N N R Cl

0 0 00ci 0 C_|

00 , ^ S
0 c -c 0 C0 0 0 O IN

-c tI N

N Cl^f
0 ^ 0 0 0 0 0 _; 0 0 O 0 N

N 0 IN Cl^

N-s Cl
0 0 0 0 0 0 0 0 0 p

- N


oll I '
In\ 00

Neotropical Primates 15(1), January 2008


Algunas correlaciones entire variables encontradas en este
studio de la variaci6n temporal de actividades de Ateles
chamek apoyan lo reportado por Wallace (2005). Asi, la
correlaci6n negative entire los porcentajes de tiempo asigna-
dos a las actividades de descanso y desplazamiento podria
indicar que la actividad descanso actda como reserve para
que otras actividades necesarias como desplazamiento
puedan ser desarrolladas. Como sugiere Wallace (2005), la
correlaci6n negative entire el tiempo asignado a las activi-
dades de desplazamiento y alimentaci6n, podria explicarse
por el hecho de que Ateles se alimenta principalmente de
recursos distribuidos en parches y energ6ticamente variables.
Esto sugiere que este primate podria estar asignando mais
tiempo a desplazarse entire various irboles con frutos de alta
calidad en terminos de contenido energ6tico, para com-
pensar la energia gastada en sus desplazamientos; futures
studios deberian enfocarse en llevar a cabo anAlisis de la
composici6n quimica de los frutos para comprobarlo. En
concordancia con lo anterior y considerando que los meses
con mayor disponibilidad de frutos no-Ficus incluirian por
lo menos algunas species energ6ticamente mas ricas que
las que fructifican en meses de menor disponibilidad, se
esperaria mayor asignaci6n de tiempo al desplazamiento
en dichos meses. Sin embargo, en nuestro studio no
encontramos correlaci6n entire la disponibilidad de frutos
no-Ficus y el porcentaje de tiempo utilizado para despla-
zamiento; ademis encontramos una tendencia a una mayor
asignaci6n de tiempo a alimentaci6n cuando la disponibili-
dad de frutos no-Ficus es menor. En este sentido, se puede
sugerir que en meses de menor disponibilidad de estos frutos
en el bosque, Ficus esti disponible. Si bien Ficus constitute
un recurso disperse espacialmente y de bajo valor energ6tico
(Morrison, 1978; Lambert y Marshal, 1991; Borges, 1993),
los irboles de Ficus produce abundantes frutos maduros
durante un period relativamente corto (Lambert y Mar-
shal, 1991). Por lo tanto Ateles tenderia a congregarse en
estos irboles para alimentarse por periods mas largos ya
que dispone de mias frutos maduros que en irboles de espe-
cies cuya maduraci6n de frutos es mias prolongada y, en con-
secuencia, disponen de menor cantidad de frutos maduros
en cada visit. De esta manera, Ficus parece ser un recurso
muy important y podria constituir un recurso clave, tal
como se sugiere para otros habitats (Terborgh, 1986; Ahu-
mada et al., 1998; Wallace, 1998; Aguilar, 2002).

Otros studios han reportado que las variaciones en el
tamafio de subgrupos se relacionan con la disponibilidad
de frutos (van Roosmalen, 1985; Chapman, 1990; Wal-
lace, 1998). Debido a que en nuestro studio no tenemos
certeza de haber observado grupos pertenecientes a una sola
unidad social, no podemos concluir respect a la tendencia
a una relaci6n positive entire estas variables. Las variaciones
en el tamafio de grupo en nuestro studio podrian estar
reflejando caracteristicas propias de diferentes unidades
sociales o de una muestra incomplete de una unidad social.
Por lo tanto, nuestros resultados reflejan solamente una

situaci6n particular para el irea estudiada con respect a
las relaciones entire disponibilidad de frutos y tamafio de

Se ha postulado que el incremento en la actividad de des-
canso durante periods de escasez de frutos podria estar
asociado a un incremento en el consume de hojas y la con-
secuente dificultad de digestion de esta parte vegetal (Ste-
venson et al., 2000; Wallace, 2005). En nuestro caso, meses
con menor disponibilidad de frutos no-Ficus ocurren tanto
al final del period humedo, como al comienzo del period
seco, pero no registramos un period en que se evidence
mayor consume de hojas por Ateles. En cuanto a la fru-
givoria, nuestro studio concuerda con el elevado consume
de frutos carnosos previamente reportado paraAteles (Klein
y Klein, 1977; van Roosmalen, 1985; Chapman, 1988),
asi como con la dominancia de pocas species en la dieta
(van Roosmalen, 1988; Wallace, 2005). Mis del 50% de
los registros de alimentaci6n correspondent al gdnero Ficus
y aproximadamente 21% corresponde s6lo a siete species,
cada una en diferentes meses. Como caracteristica coming,
estas species se encuentran generalmente dispersas y pre-
sentan copas grandes y abundante producci6n de frutos
(Ficus, Lauraceae, Protium montanum, "Canel6n", Podocar-
pus spp, Anomospermum sp y Casearia mariquitensis).

Nuestro studio no incluy6 un period para la habituaci6n
de los animals, pudiendo ello enmascarar los resultados
obtenidos en el comportamiento durante los primeros meses.
Sin embargo, aunque se podria pensar que se obtuvieron
pocos encuentros y tambidn cortos periods de observaci6n
durante los meses mias secos, por ser los primeros meses de
evaluaci6n, el ndmero de encuentros no incrementa lineal-
mente hacia el final del studio. Por tanto, es altamente
probable que la baja frecuencia de encuentros durante esos
meses corresponda a la menor disponibilidad de frutos
no-Ficus en el bosque (Mendez, 1999, Aguilar, 2002). Es
possible que durante estos meses los animals se hayan refu-
giado en zonas fuera del area de b6squeda, las cuales no eran
accesibles por las abruptas pendientes de la zona.

Este studio es pionero para la especie en este tipo de ambi-
entes y brinda un panorama general acerca de la ecologia
conductual de Ateles chamek en un bosque humedo mon-
tano. Es interesante enfatizar que algunas de las car-
acteristicas de poblaciones de Ateles estudiadas en bosques
amaz6nicos de tierras bajas, e incluso en Centroam&rica,
se mantienen en nuestra region de studio: 1) una dieta
altamente frugivora, 2) dominancia de pocas species en
la dieta, 3) tendencies a un intercambio del tiempo asig-
nado a las actividades de alimentaci6n y desplazamiento, asi
como entire descanso y desplazamiento, y 4) una correlaci6n
negative entire la disponibilidad de frutos y el porcentaje
de tiempo asignado a la alimentaci6n. Aquellos aspects
de nuestro studio que parecen contradecir los patrons
conocidos sobre la ecologia de Ateles, tales como el bajo por-
centaje de flexibilidad de la dieta (no existe un period de
mayor consume de hojas) y la poca variaci6n estacional en

el tamano de grupo, podrian tener explicaciones distintas,
por el hecho de haberse desarrollado en un ambiente muy


Este trabajo fue financiado por Wildlife Conservation
Society y John D. & Catherine T. MacArthur Foundation.
Una version preliminary del trabajo fue comentada por R.
Wallace, M. A. Pizo, P. Rubim y J. Shortt. Agradecemos
a los guias de campo P. Calisaya, F Condori y D. P&rez,
asi como la ayuda del personal del Herbario Nacional de
Bolivia en la identificaci6n del material vegetal.


Aguilar, M. S. 2002. Dispersi6n de semillas por tres espe-
cies de primates, Ateles chamek, Cebus apella y Saimiri
sciureus en un bosque de la estaci6n biol6gica del Beni,
Bolivia. Tesis de licenciatura, Universidad Mayor de San
Andres, La Paz, Bolivia.
Ahumada, J. A., Stevenson P. R. y Quii6nez, M. J. 1998.
Ecological responses of spider monkeys to temporal varia-
tion in fruit abundance: The importance of flooded forest
as a keystone habitat. Primate Conserv. 18: 10-14.
Anderson, S. 1997. Mammals of Bolivia, taxonomy and
distribution. Bull. Amer. Mus. Nat. Hist. 102: 560-561.
Bach, K., Schawe, M., Beck, S., Gerold, G., Gradstein,
S. R. y Moraes, M. 2003. Vegetaci6n, suelos y clima en
los diferentes pisos altitudinales de un bosque montano
de Yungas, Bolivia: primeros resultados. Ecol. Boliv. 38:
Borges, R. 1993. Figs, malabar giant squirrels, and fruit
shortage within two tropical Indian forests. Biotropica 25:
Carrizosa, U. J. 1990. La Selva Andina. En: Selva y Futuro,
U. J. Carrizosa y J. C. Hernindez (eds.), pp.151-184. El
Sello, Bogoti.
Chapman, C. 1988. Patch use and patch depletion by the
spider and howling monkeys of Santa Rosa National
Park, Costa Rica. Behaviour 105: 99-116.
Chapman, C. A. 1990. Ecological constraints on group size
in three species of Neotropical primates. Folia Primatol.
55: 1-9.
Chapman, C. A., Chapman, L. J., Wrangham, R. W., Hunt,
K., Gebo, D. y Gardner, L. 1992. Estimators of fruit
abundance of tropical trees. Biotropica 24: 527-531.
Dodson, C. y A. H. Gentry. 1991. Biological extinction in
western Ecuador. Ann. Miss. Bot. Gard. 78: 273-295.
Henderson, A., Churchill, S. P. y Luteyn, J. 1991.
Neotropical plant diversity. Nature 351: 21-22.
Izawa, K., Kimura, K. y Samper-Nieto, A. 1979. Grouping
of the wild spider monkeys. Primates 20: 503-512.
Johns, A. D. y Skorupa, J. P. 1987. Responses of rain-forest
primates to habitat disturbance: a review. Int. J Primatol.
8: 157-191.
Klein, L. L. y Klein D. B. 1977. Feeding Behaviour of
the Colombian Spider Monkey. En: Primate Ecology, T.

Neotropical Primates 15(1), January 2008

H. Clutton-Brock (ed.), pp.153-181. Academic Press,
Lambert, E y Marshal, A. G. 1991. Keystone characteris-
tics of bird-dispersed Ficus in a Malaysian lowland rain
forest. J Ecol. 79: 793-809.
Martin, P. y Bateson, P. 1986. Measuring behaviour: an in-
troductory guide. Cambridge University Press, New York.
Mendez, P.E 1999. La influencia de la distribuci6n y abun-
dancia de frutos sobre el tamano de subgrupo de Ateles
chamek en la EBB, Bolivia. Tesis de licenciatura, Univer-
sidad Mayor de San Andres, La Paz, Bolivia.
Milton, K., Windsor, D. M., Morrison, D. W. y Estribi M.
A. 1982. Fruiting phenologies of two Ficus species. Ecol-
ogy 63: 752-762.
Morrison, D.W. 1978. Foraging ecology and energetic
of the frugivorous bat Artibeus jamaicensis. Ecology 59:
Pacheco, L. F. y Simonetti, J. A. 1998. Consecuencias de-
mogrificas para Inga ingoides (Mimosidae) por la p&rdida
de Ateles paniscus (Cebidae), uno de sus dispersores de
semillas. Ecol. Boliv. 31: 67-90.
Pacheco, L. F. y Simonetti, J. A. 2000. Genetic structure of
a mimosoid tree deprived of its seed disperser, the spider
monkey. Conserv. Biol. 14: 1766-1775.
Palomeque, de la C. E. S. 2001. Estimaci6n y preferencia
de habitat del marimono (Ateles chamek) en el Area de
Chalalhn (Parque Nacional y Area Natural de Manejo
Integrado Madidi). Tesis de licenciatura, Universidad
Mayor de San Andr5s, La Paz, Bolivia.
Paniagua, N., Maldonado, C. y Chumacero, C. 2003.
Mapa de vegetaci6n de los alrededores de la Estaci6n Bi-
ol6gica de Tunquini, Bolivia. Ecol. Boliv. 38: 15-26
Ragusa-Netto, J. 2002. Fruiting phenology and consump-
tion by birds in Ficus -.. .... (MIQ.) (Moraceae).
Braz. J Biol. 62: 339-346.
Ribera, M. 0. 1995. Aspectos ecol6gicos del uso de la tierra
y conservaci6n en el Parque Nacional y Area Natural de
Manejo Integrado Cotapata. En: Caminos de Cotapata,
C. B. Morales (ed.), pp.1-28. Institute de Ecologia, La
Paz, Bolivia.
Rios, B. 1999. Evaluaci6n de m6todos indirectos para de-
tectar presencia de mamiferos medianos y grandes en
habitats de montana. Tesis de licenciatura. Universidad
Mayor de San Andr5s, La Paz.
Stevenson, R. P., Quifones, J. M. y Ahumada A. J. 2000.
Influence of fruit availability on ecological overlap among
four Neotropical primates at Tinigua National Park, Co-
lombia. Biotropica 32: 533-544.
Tarifa, T. 2000. Abundancia, distribuci6n y estado de con-
servaci6n del marimono (Ateles chamek) en la Reserva de
la Biosfera Estaci6n Biol6gica del Beni, Bolivia. En: Biodi-
versidad, conservacidn y manejo en la region de la Reserva de
la Biosfera Estacidn Bioldgica del Beni, Bolivia, 0. Herrera-
MacBryde, F. Dallmeier, B. MacBryde, J. A. Comiskey
y C. Miranda (eds.), pp.275-296. SI/MAB Series No4,
Smithsonian Institution, Washington DC.
Terborgh, J. 1986. Keystone plant resources in the tropical
forest. En: Conservation biology: the science of scarcity and

Neotropical Primates 15(1), January 2008

diversity, M. E. Soule (ed.), pp.330-344. Sinauer, Sun-
derland, Massachusetts.
van Roosmalen, M. G. M. 1985. Habitat preferences,
diet, feeding strategy and social organization of the black
spider monkey (Ateles paniscus paniscus) in Surinam. Acta
Amazonica 15: 9-238.
van Roosmalen, M. G. M. y Klein, L. L 1988. The spider
monkeys, genus Ateles. En: Ecology and behavior of
Neotropical primates, R. A. Mittermeier, A. B. Rylands,
A. Coimbra-Filho y G. A. B. Fonseca (eds.), pp.445-537.
WWF, Washington DC.
van Schaik, C. P. 1986. Phenological changes in a Suma-
tran rain forest. J Trop. Ecol. 2: 327-347.
Wallace, R. B. 2005. Seasonal variations in diet and forag-
ing behavior of Ateles chamek in a southern Amazonian
tropical forest. Int. J. Primatol. 26: 1053-1075.
Wallace, R. B. 1998. The behavioral ecology of black
spider monkeys in north-eastern Bolivia. Tesis de Doc-
torado, University of Liverpool Press, Liverpool.
Wallace, R. B., Painter, R. L. E. y Taber A. B. 1998. Pri-
mate diversity, habitat preferences, and population densi-
ties estimates in Noel Kempff Mercado National Park,
Santa Cruz department, Bolivia. Am. J Primatol. 46:
White, F 1986. Census and preliminary observations on
the ecology of the black-faced spider monkey (Ateles pa-
niscus chamek) in Manu National Park, Perd. Am. J Pri-
matol. 11: 125-132.
Zar, J. H. 1996. Biostatistical analysis. Prentice-Hall, USA.


J. Berton C. Harris
Diego G. Tirira
Pedro J. Alvarez L
Vicente Mendoza I.


Capuchins in the genus Cebus (Cebidae) are medium-
sized, gregarious, and generally arboreal monkeys found
in Central and South America (Emmons and Feer, 1997;
Eisenberg and Redford, 1999). Like other Cebus, the
white-fronted capuchin (Cebus .alb.fo:u Humboldt, 1812)
is omnivorous, consuming seeds, fruit, invertebrates, and
vertebrates (Defler, 1979a; Eisenberg and Redford, 1999;
Tirira, 2007). Cebus alb.p'os can occupy a home range
as large as 120 ha and the species is known to wander to
varied habitat types in search of widely separated food
resources (Defler, 1979a; Terborgh, 1983; Haugaasen and
Peres, 2005).

Capuchins are mostly lowland species although Cebus
apella and C. ,IbfJ ovs have been recorded up to 2,130 m
a.s.l. and 2,000 m a.s.l. respectively (Tate, 1939; Hershko-
vitz, 1949; Emmons and Feer, 1997). Cebus Ilb.Jo,:s is
known from humid to dry forests in isolated areas of north-
ern Colombia and Venezuela, in western Ecuador, and in
the upper and middle Amazon basin of Colombia, Ecua-
dor, Peru, Bolivia, and Brazil west of the Rios Tapaj6s and
Negro (Emmons and Feer, 1997; Eisenberg and Redford,
1999; Tirira, 2007). Cebus .zIbJo,:s includes several sub-
species whose distributions and morphological differences
have been incompletely studied. Historically, 11 subspe-
cies were recognized (Cabrera, 1957), but more recently,
Groves (2005) recognized only five, two of which occur in
Ecuador, C. a. aequatorialis and C. a. yuracus. The latter
is most likely involved in our records in southern Ecuador.
Cebus aIlb.fovs aequatorialis is endemic to humid to dry
tropical and subtropical forests west of the Andes in Ecua-
dor (Albuja and Arcos, 2007; Tirira, 2007); its type locality
is Rio de Oro, Manabi province (Cabrera, 1957). Cebus
,Ilb.j oso yuracus is found in humid tropical and subtropical
forests east of the Andes in Ecuador and northeastern Peru,
between the Marafi6n and Putumayo rivers (Emmons and
Feer, 1997; Tirira, 2007); its type locality is Montalvo, Rio
Bobonaza, Pastaza province, Ecuador (Cabrera, 1957).

The highest altitude records for C. aIlb.fj'o across its dis-
tribution are as follows. In Colombia, the highest known
record of the species is 2,000 m a.s.l. in the department
of Tolima (Defler, 2003). The highest confirmed record
in Perd comes from the eastern foothills of the upper Rio



Neotropical Primates 15(1), January 2008
Comainas, Cordillera del C6ndor (Amazonas depart-
ment) at 1,738 m a.s.l. (Schulenberg and Awbrey, 1997;
this record is of C. a. yuracus) and there are unconfirmed
sightings from the headwaters of the Rio Poyeni, Cordillera
de Vilcabamba (Apurimac department) at 2,050 m a.s.l.
(Alonso et al., 2001). In Venezuela, the species is known
up to 850 m a.s.l. (Linares, 1998). In Bolivia, the altitu-
dinal range of the species has not been well documented
but there are no specimen records at altitudes greater than
2,000 m a.s.l. (Anderson, 1997). In Ecuador, the highest
previously reported altitudes for C. a. yuracus are 1,600 m
a.s.l. from Sardinayacu in Sangay National Park (0203'S,
7809'W; Albuja, 1996) and 1,550 m a.s.l. in Coangos,
Morona-Santiago province (03o29'S, 7814'W; Schu-
lenberg and Awbrey, 1997). C. a. aequatorialis was once
reported from 1,660 m a.s.l. at Gualea, Pichincha province
(Allen, 1914). Allen (1914) indicated that the altitude of
Gualea is "7,000 ft" (2,330 m a.s.l.). As Gualea actually
lies at 1,660 m a.s.l., this elevation must be erroneous and
no future authors have included this high elevation record.
Here we report C. a. yuracus (subspecies inferred by range)
from 2,425-2,515 m a.s.l. in montane upper subtropical
forest in the Cordillera de Sabanilla, Zamora-Chinchipe
province, Ecuador. This record extends the known altitu-
dinal range of the species by 515 m and for C a. yuracus
by 777 m.

Study Site and Methods

We observed C ,albf o:us in mature forest at 2,425-2,515
m a.s.l. (4o29'S, 7907'W) in the Tapichalaca Biological
Reserve, a 2,870 ha protected area administered by the Joc-
otoco Conservation Foundation located above the town of
Valladolid, Zamora-Chinchipe province, adjacent to Podo-
carpus National Park. The forest in this area, characterized
as upper subtropical forest (Krabbe et al., 1999), has an
average canopy height of approximately 10 m with 20 m
emergent crowns and it receives about 4 m of rainfall annu-
ally. The canopy is characterized by Moraceae (Ficus sp.),
Euphorbiaceae (Croton sp.), Lauraceae, and Rubiaceae, and
the understory is largely made up of bamboo (Chusquea
sp.), Piperaceae, and Melastomataceae. The vegetation at
Tapichalaca is covered in abundant mosses and liverworts
as well as vascular epiphytes in families such as Bromeli-
aceae, Araceae, and Orchidaceae (Krabbe et al., 1999).

Results and Discussion

On 15 January 2007 at 11:00 (EST), PJAL and VMI
observed one individual C ,albJf ons for 5 minutes at
2,425 m a.s.l. Then on 31 January, approximately 500 m
up the same trail, JBCH and VMI observed three indi-
viduals for 2.5 hr (from 9:00-11:30) at 2,515 m a.s.l. The
weather on both observation days was partly cloudy and
warm (approx. 160 C). We photographed and made audio
recordings of the group on 31 January. The capuchins fed
on Persea sp. (Lauraceae) and Croton sp. (Euphorbiaceae)
fruits, and unidentified small items inside bromeliads

Neotropical Primates 15(1), January 2008

(probably arthropods). We were unable to sex the capu-
chins but all individuals appeared to be adults (Fig. 1). The
capuchins spent all of the observation time in trees. The
following observations apply to the group of three capu-
chins we observed on 31 January. The capuchins made
frequent, although quiet vocalizations, as is typical for
the species (Defler, 1979b). The three individuals often
made soft yah danger calls (Defler, 1979b) throughout the
2.5 h observation period, and were likely using the calls
to communicate the presence of the human observers (T.
Defler, pers. comm.). The capuchins vocalized frequently
but the calls were quiet and could only be heard within
approximately 30 m of the vocalizing individual. Once,
a capuchin approached a human observer and stared and
shook branches for 30 sec; this behavior is a common sign
of aggression in C. ,alb jons (Defler, 1979b; Fig. 1). The
dense vegetation and the abundance of epiphytes made it
difficult to monitor all of the individuals constantly but
it seemed no capuchin was ever farther than 20 m from
another. During the observation period the group of capu-
chins traveled approximately 150 m.

Cebus ,z/bJou:s is most often observed in groups of 7-35
individuals (Defler, 1979a,b; Defler, 1982; Terborgh,
1983; Eisenberg and Redford, 1999), and our observations
of much smaller groups are unusual for the species. We are

Figure 1. Aggressive stare of white-fronted capuchin (Cebus albi-
frons) in upper subtropical forest (2,515 m) in southern Ecuador.

unsure why the species apparently lives in such small groups
in the Tapichalaca area but we hypothesize that small group
size may confer some advantage in the high altitude habi-
tat. Additionally, C. ,Idbf o:s may inhabit Tapichalaca in
low numbers because the high Andes are at the very limit
of the species' distribution. This pattern of rarity at the
edge of a distribution is shown in many widely-distributed
animals (Brown, 1984). It is also possible that we observed
subgroups of larger capuchin groups in the area. Subgroups
are sometimes formed in Cebus when seasonal changes
cause resources to be dispersed (Izar, 2003; Lynch Alfaro,
2007). The high altitude environment at Tapichalaca may
provide more dispersed food resources than lower elevation
habitats which may result in similar Cebus subgrouping.
More research is needed to better understand small group
size of highland capuchins.

Cebus albf} o, has never before been recorded at Tapicha-
laca, even though the area has been frequented by experi-
enced observers since 1997. We searched for the species
for several days following these observations but could not
relocate any capuchins. In addition, we conducted exten-
sive field work in the area for the following seven months
and never re-encountered the species. The small number
of records of C. ,I/bfoJ:s can partially be explained by the
species' large home range size and quiet, inconspicuous
behavior, but the species is likely to be rare in the area. We
hypothesize that C. ,Ib/.oJ:s is a low density resident in
upper subtropical forest in the Tapichalaca region.


We are grateful to the Jocotoco Conservation Founda-
tion and the World Land Trust for funding and logistical
support. We thank the JF office staff and the Tapichalaca
guardaparques, as well as R. Carpio, T. Defler, M. Juinia,
J. Lynch Alfaro, R. Ridgely, and F Sornoza for their sup-
port. The manuscript benefited from the comments of A.
Chenault, M. Harris, L. Reid, and B. Scheffers.

J. Berton C. Harris, Pedro J. Alvarez L., and Vicente
Mendoza I., Fundaci6n de Conservaci6n Jocotoco, Av.
Los Shyris N37-146 y El Comercio, Quito, Ecuador,
e-mail: , Diego G. Tirira,
Fundaci6n Mamiferos y Conservaci6n, Quito, Ecuador,
e-mail: .


Albuja, L. 1996. Diagn6stico faunistico para la actual-
izaci6n del plan de manejo del Parque Nacional Sangay.
Plan maestro para la protecci6n de la biodiversidad me-
diante el fortalecimiento del Sistema Nacional de Areas
Protegidas. Proyecto INEFAN-GEE INEFAN. Quito,
Albuja, L. and Arcos, D. R. 2007. Evaluaci6n de las pobla-
ciones de Cebus .IbjoJns cf. aequatorialis en los bosques

suroccidentales ecuatorianos. Revista Politecnica 27(4),
Serie Biologia 7: 58-67.
Allen, J. A. 1914. New South American monkeys. Bull.
Am. Mus. Nat. Hist. 33: 647-655.
Alonso, L. E., Alonso, A., Schulenberg, T. S., and Dallmeier,
E (eds.). 2001. Biological and social assessments of the
Cordillera de Vilcabamba, Peru. Rapid Assessment Pro-
gram (RAP). Working Papers 12 and SI/MAB Series 6.
Conservation International. Washington, DC.
Anderson, S. 1997. Mammals of Bolivia, taxonomy and
distribution. Bull. Am. Mus. Nat. Hist. 231: 1-652.
Brown, J. H. 1984. On the relationship between abundance
and distribution of species. Am. Nat. 124: 255-279.
Cabrera, A. 1958. Catilogo de los mamiferos de Am&rica
del Sur. Revista del Museo Argentino de Ciencias Naturales
"Bernardino Rivadavia" Ciencias Bioldgicas 4: 1-308.
Defler, T. R. 1979a. On the ecology and behavior of Cebus
,Idbo.:s in eastern Colombia: I. Ecology. Primates 20:
Defler, T. R. 1979b. On the ecology and behavior of Cebus
,IbfoJ.s in eastern Colombia: II. Behavior. Primates 20:
Defler, T. R. 1982. A comparison of intergroup behavior in
Cebus alb.poIts and C. apella. Primates 23: 385-392.
Defler, T. R. 2003. Primates de Colombia. Conservaci6n In-
ternacional. Serie de guias tropicales de campo 4. Bogotai
Eisenberg, J. and Redford, K. 1999. Mammals of the Neo-
tropics, Vol. 3. The University of Chicago Press, Chicago.
Emmons, L. H. and Feer, F 1997. Neotropical rainforest
mammals. A field guide. 2nd edition. The University of
Chicago Press, Chicago.
Groves, C. 2005. Order Primates. In: Mammalspecies of the
world, a taxonomic and geographic reference, 3rd edition,
D. E. Wilson and D. M. Reeder (eds.), pp.111-184, The
John Hopkins University Press, Baltimore.
Haugaasen, T. and Peres, C. T. 2005. Primate assemblage
structure in Amazonian flooded and unflooded forests.
Am. J. Primatol. 67: 243-258.
Hershkovitz, P. 1949. Mammals of northern Colombia.
Preliminary report 4: monkeys (Primates), with taxo-
nomic revisions of some forms. Proceedings of the U. S.
NationalMuseum 98: 323-427.
Izar, P. 2003. 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.
Krabbe, N., Agro, D. J., Rice, N. H., Jacome, N., Na-
varrete, L., and Sornoza M., E 1999. A new species of
antpitta (Formicariidae: Grallaria) from the southern Ec-
uadorian Andes. Auk 116: 882-890.
Linares, 0. 1998. Mamiferos de Venezuela. Sociedad Con-
servacionista Audubon de Venezuela, Caracas.
Lynch Alfaro, J. W. 2007. Subgrouping patterns in a
group of wild Cebus apella nigritus. Int. J. Primatol. 28:
Schulenberg, T. S. and Awbrey, K. (eds.). 1997. The Cordil-
lera del C6ndor region of Ecuador and Peru: A biological

Neotropical Primates 15(1), January 2008

assessment. Rapid Assessment Program (RAP). Working
Papers 7. Conservation International, Washington, DC.
Tate, G. H. H. 1939. The mammals of the Guiana region.
Bull. Am. Mus. Nat. Hist. 76: 151-229.
Terborgh, J. 1983. The behavioral ecology of five species of
Amazonian primates. Princeton University Press, Princ-
eton, New Jersey.
Tirira, D. G. 2007. Guia de campo de los mamiferos del Ec-
uador. Publicaci6n especial sobre los mamiferos del Ecua-
dor 6, Ediciones Murcidlago Blanco, Quito, Ecuador.

Mauricio de Almeida Noronha
Jose de Sousa e Silva Jfnior
Wilson Roberto Spironello
Dayse Campista Ferreira


The Mau&s marmoset, Mico mauesi (Mittermeier et al.,
1992), was described in the last decade of the 20th cen-
tury and originally placed in the genus C .-' The spe-
cies was described from a specimen at the Museu Paraense
Emilio Goeldi holotypee: MPEG -22177) and observations
of captive and wild animals. At the time of its description
the species was only known to occur at the type locality, on
the left bank of the Rio Mau5s-Acu (03o23'S, 5746'W),
just across from the city of Maues in the state of Amazonas,
Brazil. Based on distribution patterns of other species of
Amazonian marmoset, Mittermeier et al. (1992) predicted
that the new species would occur throughout the entire
medium to lower Mau5s-Urarii-Abacaxis interfluve and
that its range would be limited by these three rivers.

During primate surveys conducted in and around the
Canuma-Urarii-Abacaxis interfluve, Silva Jr. and Noronha
(1995, 2000) registered the presence of M. mauesi at two
localities on the right bank of the lower Rio Abacaxis:
Abacaxis (03o55'S, 5845'W) and Santa Maria (03o54'S,
5846'W). Three specimens were collected at Santa Maria
and deposited in the zoological collection of the Museu
Paraense Emilio Goeldi (MPEG 23962, 23963, 23964).
These findings (Silva Jr. and Noronha, 1995, 2000) and
data from the literature (Hershkovitz, 1977; Vivo, 1988,
1991), confirm the northern and western limits of the
species' range as proposed by Mittermeier et al. (1992).
According to the range map produced by these authors,
the distribution of M. mauesi should be limited to the east
by the Rio Mauds-Acu and extend as far south as 05 S.
The objective of this study is to update knowledge of the
geographic distribution of M. mauesi through the collec-
tion and analysis of new field data.

Neotropical Primates 15(1), January 2008

Data on the occurrence and habitat of the mauns mar-
moset were collected during a field trip to the lower Rios
Madeira and Tapaj6s interfluve (Noronha, 2004). The
area surveyed during the study spanned the headwaters of
the Rios Mauns, Abacaxis and Sucunduri. Data were col-
lected using two methods: direct sightings and interviews
with local informants. Mico mauesi was identified based
on the diagnostic characters described by Mittermeier et al.
(1992) and comparisons with material from the zoological
collections of the Museu Paraense Emilio Goeldi and the
National Institute of Amazonian Research (MPEG-23962,
23963, 23964; INPA-4082, 4109, 4105, 4106).

Results and discussion

During the study, eight new occurrence records were
obtained for M. mauesi, seven via direct sightings and one
in an interview. Marmosets with the traits described by
Mittermeier et al. (1992) were seen at sites 4, 5, 6, 7, 8, 9

and 10 (Fig. 1.). All records were obtained in terra firme
primary forest, secondary growth and igapd, suggesting
that the habitat requirements of this species are similar to
other members of the genus (Digby et al., 2005; Mitter-
meier et al., 1992; Silva Jr. and Noronha, 1995, 2000).
These new records demonstrate that the geographic distri-
bution delineated by Mittermeier et al. (1992) underesti-
mated the true range of the species. Although northern
and western limits were predicted correctly, authors were
over cautious with respect to the eastern and southern
limits. Data from this study extend the range east to the
Rio Mauds-Acu; south almost as far as the Igarapd do Suru-
bim (06053'S, 59003'W) along the corridor formed by the
Rios Tapaj6s and Sucunduri; west to the Rio Sucunduri,
reaching the mid and upper Sucunduri-Abacaxis interfluve
(the northernmost record was the Cantagalo site: 06033'S,
59005'W). Interviews with local residents suggest that the
species may extend as far south as the region of Igarapd do
Urucd (0709'S, 5856'W), on the right bank of the Rio

Figure 1. Geographic range of Mico mauesi, based on Mittermeier et al. (1992), Silva Jr. and Noronha (1995, 2000) and this
study: (1) left bank of the Rio Mau&s-Acu (0323'S, 57o46'W), across from Mauds (type locality), municipality of Mauds;
(2) Abacaxis (03o55'S, 58045'W), right bank of the Rio Abacaxis, municipality of Nova Olinda do Norte; (3) Santa Maria
(03o54'S, 58046'W), right bank of the Rio Abacaxis, municipality of Nova Olinda do Norte; (4) right bank of the Rio Sucun-
durf (0648'S, 59o04'W), municipality of Apuf; (5) right bank of the Rio Abacaxis (06o41'S, 58o50 'W) municipality of Mauds;
(6) left bank of the Rio Tapaj6s (0634'S, 5835'W), municipality of Mauds; (7) Cantagalo, right bank of the Rio Sucundurf
(06'33'S, 5905'W), municipality of Borba; (8) left bank of the Rio Tapaj6s (06034'S, 5824'W), municipality of Mauds; (9)
mouth of the Igarap6 do Surubim, right bank of the Rio Sucunduri (0653'S, 5903'W), municipality of Apuf; (10) Floresta
Nacional do Pau Rosa, right bank of the Rio Abacaxis (04o40'S, 5815'W), municipality of Mau&s; (11) mouth of the Igarap6
do Uruci (0709'S, 58056'W), left bank of the Rio Sucunduri, municipality of Apuf.

ty 56*
_,/Parank do
rarA R_ Mau6s A94

4'. IFL Madeim o2 4'

R_ AJmwaids
4- R_ Paranari

lip 6'. R_ 7 *- IRL TaT4 6'

Occurrence records of
Mico mauesi
oE This study
Other studies

According to Hershkovitz (1977), Mico humeralifer is
expected to occur in the Sucunduri-Tapaj6s-Juruena inter-
fluve. Rylands (1994) suggested that this species' range
could extend as far south as 8 S. This study however, does
not support these expectations. It is possible that Mico
mauesi's range extends between the Rios Sucunduri and
Juruena, reaching as far south as the transition between
dense tropical forest, open tropical forest and savanna
(BRASIL, 1975) at around 7o40'S, or even as far as the
Serra do Sucunduri at 8010'S. These predictions coin-
cide with the southern limit for M. humeralifer proposed
by Rylands (1994). According to Rylands (1981), south
of this latitude the species M. melanurus should occur
between the Rios Aripuana and Juruena. This hypothesis
is partially supported by the occurrence of M. melanurus
in the Serra do Sucunduri (08o34'S, 59008'W) and on the
right bank of the Rio Bararati (08o21'S, 5837'W). Mico
melanurus was the only member of the genus occurring
at these localities (M. A. Noronha, unpubl. data). Our
results show that the geographic distribution of M. mauesi
is large relative to that of other species of the genus in the
Madeira-Tapaj6s interfluve. We did not find any area of
contact between M. mauesi and any other species of Mico,
suggesting that the species is parapatric with respect to its
congeneric neighbors M. humeralifer, M. melanurus, M.
acariensis and M. saterei.

This study quadrupled the number of recorded localities for
M. mauesi, and revealed a range over twice that proposed
by Mittermeier et al. (1992). The improved knowledge
of this taxon's geographic distribution also provides infor-
mation about its occurrence within protected areas. The
mau5s marmoset is probably present in the Floresta Nacio-
nal do Pau Rosa (994,800 ha), Parque Nacional do Juru-
ena (2,002,565 ha) and the Floresta Nacional de Mau5s
(438,440 ha), and if we assume that the southern limit of
its range is close to the Serra do Sucunduri, it also occurs
in the Reserva de Desenvolvimento Sustentivel do Bararati
(113,606 ha), Parque Estadual do Sucunduri (808,312 ha)
and the Floresta Nacional do Apui (185,946 ha).


This study was supported by Fundagao 0 Boticario de Pro-
tegao a Natureza.

Mauricio de Almeida Norofia, Rua dos Jatobis, 142,
69085-380, Coroado III, Manaus, Amazonas, Brasil,
e-mail: , Jos6 de Sousa e Silva
Junior, Museu Paraense Emilio Goeldi, Coordenacao de
Zoologia, caixa postal 399, 66040-170, Belkm, Pari, Brazil,
e-mail: , Wilson Roberto
Spironello, Instituto Nacional de Pesquisas da Amaz6nia
(INPA), Coordenacao de Pesquisas em Silvicultura Tropi-
cal, CP 478, 69060-001, Manaus, AM, Brazil, e-mail:
, and Dayse Campista Ferreira,
Rua dos Jatobis, 142, 69085-380, Coroado III, Manaus,
Amazonas, Brasil, e-mail: .

Neotropical Primates 15(1), January 2008


Digby, L. J, Ferrari, S. E and Saltzman, W. 2005. Cal-
litrichines: The role competition in cooperatively breed-
ing species. In: Primates in perspectives, C. J. Campbell, A.
E Fuentes, K. C. Mackinnon, M. Panger and S. Bearder
(eds.), pp.85-106. Oxford University Press.
Hershkovitz, P. 1977. Living New World monkeys. Part 1:
(Platyrrhini), with an introduction to Primates. Chicago
University Press, Chicago.
Mittermeier R. A., Schwarz, M. and Ayres, M. 1992. A
new species of marmoset, genus C .-' Erxleben,
1977 (Callitrichidae, Primates) from the Rio Mau&s
region, state of Amazonas, central Brazilian Amazonia.
Goeldiana, Zoologia 14: 1-17.
Noronha, M. A. 2004. Estado atual de conservagao e dis-
tribuigao geogrnfica do Mico saterei Silva Jr. & Noronha,
M.A. 1998 (Primates: Callitrichidae), na Amaz6nia Cen-
tral, Brasil. MSc thesis, Instituto de Florestas, Universi-
dade Federal Rural do Rio de Janeiro, Rio de Janeiro.
Rylands, A. B. 1981. Preliminary field observations on the
marmoset, C .-' humeralifer intermedius (Hersh-
kovitz, 1977), at Dardelos, Rio Aripuana, Mato Grosso.
Primates 22 (1): 46-59.
Rylands, A. B. 1994. Sagui-de-Santardm. In: Livro vermelho
das espicies de mamiferos brasileiros ameafados de extinfao,
G. A. B. Fonseca; A. B. Rylands; C. M. R. Costa; R. B.
Machado; Y. L. R. Leite (eds), pp.63-72. Fundagao Bio-
diversitas, Belo Horizonte, MG., Brazil.
Silva Jdnior, J. S. da and Noronha, M. A. 1995. A new
record for C .-' mauesi Mittermeier, Schwarz &
Ayres, 1992. Neotrop. Primates 3(3): 79-80.
Silva Jdnior, J. S. da and Noronha, M. A. 2000. Resultados
de uma pequena expedigao primatol6gica a Amaz6nia
Central (Primates, Platyrrhini). A Primatologia no Brasil
Vivo, M. de 1988. Sistemdtica de C- Erxleben, 1777.
PhD thesis, Universidade de Sao Paulo, Sao Paulo.
Vivo, M. de 1991. Taxonomia de C .- Erxleben, 1777
(Callitrichidae, Primates). Fundagao Biodiversitas, Belo
Horizonte, Minas Gerais, Brasil.

Mauricio de Almeida Noronha
Wilson Roberto Spironello
Dayse Campista Ferreira

The black-tailed marmoset (Mico melanurus) was first
described by E. Geoffroy in Humboldt, 1812. He pro-
vided a brief description of the species but only noted
Brazil as the type locality. Allen (1916, in Vivo, 1991)
later determined the type locality to be Cuiabi, in the state
of Mato Grosso. Mico melanurus was considered part of
the group C .-' argentata sensu Hershkovitz (1977)
and Vivo (1988, 1991), and associated with the bare-

Neotropical Primates 15(1), January 2008

eared marmosets Mico saterei and M. acariensis. Diagnos-
tic characters include pigmented ears and ear-surfaces, an
absence of ear tufts, white or yellowish-white spots on the
upper thighs, and a completely black tail (Vivo, 1991).
Vivo (1991) noted color variations among the different
individuals collected. Specimens from Circeres, Palmei-
ras, Rio Aricid, Santo Ant6nio do Leverger, Chapada dos
Guimaraes, Aripuana and Agua Dulce (Chaco-Paraguay)
were very similar, with brown torsos and anterior limbs,
while those collected in Corumbi and a specimen from the
Rio Paraguay were noticeably lighter.

The black-tailed marmoset has an ample geographic range,
extending from the headwaters of the Rios Madeira,
Mamord and Guapor6, to the Rios Aripuana and Juruena
(van Roosmalen et al., 2000). Prior to this study, the spe-
cies was only known to occur in Bolivia, Paraguay and the
states of Mato Grosso and Mato Grosso do Sul in Brazil.
Fonseca et al. (1994) predicted that the northernmost limit
of the species' range would probably occur in the area
between the Rios Aripuana and Juruena, near the Serra do
Sucunduri or the headwaters of the Rio Sucunduri at 8S,
in the Brazilian state of Amazonas.

During a recent (June and July, 2006) primate survey of
the Aripuana-Juruena interfluve in the state of Amazonas,
we observed the brown form of Mico melanurus in both
terra firme and campinarana forests at two locations near
the Serra do Sucunduri. At the first site, three groups
were spotted on the right bank of the Rio Sucunduri in
the Floresta Estadual do Sucunduri (08o34'S, 59008'W).
A specimen was collected and deposited in the vertebrate
collection of the National Institute of Amazonian Research
(INPA- lote 2002). At the second site, a troop was seen
on the right bank of the Rio Bararati, a left bank tribu-
tary of the Rio Tapaj6s, in the Parque Estadual do Sucun-
duri (08o21'S, 5837'W). These new records confirm the
northern limit of the species' range proposed by Fonseca et
al. (1994) (Fig. 1.), and if predicted geographic ranges for
the species M. acariensis (van Roosmalen et al., 2000) and
M. mauesi are correct, indicate possible contact zones with
these species further north, on the left and right banks of
the Rio Sucunduri, respectively.

1 I1
Figure 1. Geographic range of Mico melanurus, based on Vivo (1991) and records from this study: (1) N. Chiquitos, Rio Quiser,
Santa Cruz, Bolivia; (2) Aripuana, Mato Grosso, Brazil; (3) Ciceres, Mato Grosso; (4) Chapada dos Guimaraes, Mato Grosso; (5) Rio
Aricd, Mato Grosso; (6) Santo Ant6nio do Leverger, Mato Grosso; (7) Palmeiras, Mato Grosso; (8) Corumbi, Mato Grosso do Sul;
(9) Urucum, Mato Grosso do Sul; (10) 48 km E de Agua Dulce, Chaco, Paraguay; (11) Floresta Estadual do Sucunduri, right bank
of the Rio Sucunduri, Amazonas, Brazil (08o34'S, 5908'W); and (12) Parque Estadual do Sucunduri, right bank of the Rio Bararati,
Amazonas (08o21'S, 58037'W).

66,V54 W


This study was supported by the Juruena-Apui Expedition,
World Wildlife Fund Brazil (WWF), the Instituto Brasil-
eiro de Meio Ambiente e Recursos Naturais Renoviveis
(IBAMA) and the Secretaria de Estado de Meio Ambiente
e Desenvolvimento Sustentivel do Amazonas.

Mauricio de Almeida Norofia, Rua dos Jatobis, 142,
69085-380, Coroado III, Manaus, Amazonas, Brazil.
e-mail: , Wilson Roberto
Spironello, Instituto Nacional de Pesquisas da Amaz6nia,
Coordenacao de Pesquisas em Silvicultura Tropical, CP
478, 69060-001, Manaus, AM, Brazil. e-mail: inpa.gov.br>, Dayse Campista Ferreira, Rua dos Jatobis,
142, 69085-380, Coroado III, Manaus, Amazonas, Brazil.
e-mail: .


Fonseca, G. A. B. da, Rylands, A. B., Costa, C. M. R.,
Machado, R. B. and Leite, Y. L. R. 1994. Livro vermelho
dos mamiferos brasileiros ameacados de extincao. Belo Hori-
zonte, Fundagao Biodiversitas.
Hershkovitz, P. 1977. Living New World monkeys (Platyr-
rhini) vol. 1. University of Chicago Press, Chicago.
van Roosmalen, M. G. M, van Roosmalen, T, Mittermeier,
R. A. and Rylands, A. B. 2000. Two new species of mar-
moset, genus C .-' Erxleben, 1777 (Callithrichidae,
Primates) from the Tapaj6s/Madeira interfluvium, South
Central Amazonia, Brazil. Neotrop. Primates, 8(1): 2-18.
Vivo, M. de 1988. Sistemitica de C Erxleben, 1777.
PhD thesis, Universidade de Sao Paulo, Sao Paulo.
Vivo, M. de 1991. Taxonomia de C .- Erxleben, 1777
(Callitrichidae, Primates). Fundagao Biodiversitas, Belo
Horizonte, Minas Gerais, Brasil.


Rodrigo Anzolin Begotti
Laura Frey Landesmann

Aldm da perda e fragmentagao do habitat, principals amea-
gas a sobrevivencia de mamiferos no Brasil (Costa et al.,
2005), alguns primatas da Mata Atlantica term enfrentado
o problema da introdugao accidental ou intencional de espe-
cies competidoras. Este 6 o caso de alguns calitriquideos dos
estados do Rio de Janeiro e porgao centro-sul de Sao Paulo,
onde representantes do genero C principalmente
C jacchus e C. penicillata tem sido introduzidos (Olmos
e Martuscelli, 1995; Costa et al., 2005; Bicca-Marques et
al., 2006; Ruiz-Miranda et al., 2006). Estas espcies pos-
suem alto potential de colonizagao de habitats florestais,

Neotropical Primates 15(1), January 2008

hibridizagao e competigao corn representantes da fauna
local, al6m da possivel transmissao de doengas (Costa et
al., 2005; Bicca-Marques et al., 2006; Ruiz-Miranda et al.,
2006; Mendes Pontes et al., 2007). Das 25 espcies de pri-
matas que ocorrem na Mata Atlbntica, 16 se encontram em
risco de extingao por um conjunto destas ameagas (Costa et
al., 2005; Mendes Pontes et al., 2006).

Apesar da grande preocupagao gerada pela introdugao destas
espcies (Bicca-Marques et al., 2006), poucos estudos tem
enfocado a auto-ecologia dos sagiiis em ambientes recente-
mente colonizados. Ruiz-Miranda et al. (2006), por exem-
plo, observaram que a presenga de grupos introduzidos e
hibridos de C. jacchus e C. penicillata provocou mudangas
no comportamento de forrageio e defesa de territ6rio em
grupos de micos-leoes-dourados (Leontopithecus rosalia,
Callitrichidae) no municipio de Silva Jardim, Estado do
Rio de Janeiro. Neste trabalho relatamos events relacio-
nados a predagao de ninhos de aves por um grupo hibrido
de C. jacchus e C penicillata introduzidos em uma area
residential arborizada, adjacent a pequenos fragments
florestais na regiao metropolitan de Sao Paulo.

O estudo foi realizado no Condominio Nova Higien6po-
lis, municipio de Jandira, localizada na Grande Sao Paulo
(23o33'S, 46o52'0). A area de estudo apresenta temper-
atura mrdia annual de 20C e precipitagao entire 1,500 e
1,600 mm. 0 clima segundo a classificacao de Kbppen 6
do tipo Cfb (temperado de inverno seco). A area se encon-
tra dentro da distribuigao do sagiii-da-serra-escuro (Cal-
lithrix aurita), endemico da Mata Atlhntica e classificado
na categoria "Vulnerivel" na lista de mamiferos ameagados
do Brasil (Olmos e Martuscelli, 1995; Costa et al., 2005).
Nao hi relatos de moradores e nenhum individuo desta
esp6cie foi encontrado na area e adjacencias. As observa-
goes foram realizadas entire os dias 27 e 28 /10/2006 atrav5s
do m6todo de amostragem instantinea (Altmann, 1974)
totalizando 21 horas, estando o grupo estudado habituado
a presenga humana. Uma femea e um macho adults, um
macho subadulto, uma femea jovem e dois infants com-
punham o grupo estudado.

Dois events de predagao de ninhos de aves foram regis-
trados. 0 primeiro event ocorreu por volta das 15:20 do
dia 27. 0 grupo estava movimentando-se entire arbustos
e arvores quando detectou um ninho de avoante (Zenaida
auriculata, Columbidae) a aproximadamente 6 m de
altura. Imediatamente, os individuos adults (o macho
carregando os dois infants) se aproximaram, afugentando
a ave que incubava os ovos (provavelmente dois). Estes
foram rapidamente consumidos, estando a femea adulta a
frente. A jovem femea se encontrava na mesma irvore e
somente se aproximou depois que os adults se afastaram
do ninho. Ela permaneceu pr6xima ao ninho por cerca de
30 min., manipulando os fragments das cascas dos ovos
predados. 0 individuo de Z. auriculata que incubava os
ovos permaneceu nas proximidades observando a movi-
mentagao. 0 segundo event ocorreu por volta das 9:30

Neotropical Primates 15(1), January 2008
do dia 28, ap6s deslocamento de aproximadamente 200 m
por irvores e cabos telef6nicos, quando o grupo comegou
a ser atacado em v6os rasantes por um individuo de sabia-
barranco (Turdus leucomelas, Turdidae). 0 grupo, entao,
dissipou-se. Nesta ocasiao, os infants estavam sendo car-
regados pelo macho subadulto acompanhado pela femea
jovem. Os adults se dirigiram para outra irvore, quando
foram atacados por um sabi -do-campo (Mimus saturni-
nus, Mimidae) em v6o rasante. Para tentarem se abrigar
dos ataques de ambas as aves, a femea e o macho adults se
refugiaram em uma arvore na qual se encontrava, a aproxi-
madamente 8 m de altura, um ninho de M. saturninus, que
foi imediatamente atacado. A ave que incubava os ovos
defended o ninho por meio de bicadas e peitadas nos sagiiis
por aproximadamente 20 min. at6 que eles se afastaram
sem predar qualquer ovo ou filhote. No mesmo period,
em duas ocasi6es, o grupo foi atacado em locais distintos
por individuos de T leucomelas.

A predagao de ninhos 6 citada na literature como sendo
um recurso eventualmente utilizado por sagiiis (Miranda
e Faria, 2001; Lyra-Neves et al., 2007). Mendes Pontes e
Shares (2005) citam a predagao de filhotes em ninhos de
pombo-domestico (Columba livia domestic, Columbidae)
e sabii-laranjeira (T rufiventris, Turdidae), al6m do relato
de interagoes agonisticas entire esta iltima e os sagiiis. Ha
registros de C. jacchus predando ovos e filhotes de ninhos
de T leucomelas e de mais 10 esp6cies na Estagao Ecol6gica
do Tapacuri em Pernambuco (Lyra-Neves et al., 2007). 0
fato justificaria o comportamento de defesa empregado
pela ave frente a simples aproximacao do grupo de sagiiis
da irvore onde estava o ninho. Os mesmos autores ressal-
tam o comportamento de predagao sempre efetuado pelos
individuos adults e dominantes do grupo. A predagao de
ninhos pode interferir negativamente na riqueza e diversi-
dade da avifauna (Argel de Oliveira, 1995), embora seus
efeitos sejam dificeis de serem mensurados por estudos
comportamentais. Em areas urbanas, Sinclair et al. (2005)
encontraram maior abundincia de mamiferos predadores
de ninhos em manchas lineares e estreitas de vegetagao.
A regiao metropolitan de Sao Paulo apresenta poucas
areas verdes, sendo a maioria destas composta por jardins
e arborizagao de ruas e avenidas. Assim, a comunidade
de aves em ambientes urbanos pode ser seriamente com-
prometida pela introdugao de sagiiis oriundos do trifico
de animals. Aldm disso, a avifauna destes ambientes e de
pequenas manchas de vegetagao native localizadas pr6xi-
mas is areas de soltura destes animals sofrem o sinergismo
dos efeitos da fragmentagao, caga, incid&ncia constant de
incendios e o aumento das taxas de predagao de ninhos.


Somos imensamente gratos a Robert Frey, Sylvia Landes-
mann Frey e Marc L. Frey pelo apoio logistico. Tambdm
agradecemos a Mauro Galetti Rodrigues, Carlos 0. A.
Gussoni e aos editors pelos valiosos comentarios e sug-
est6es para as verses preliminares deste artigo.

Rodrigo Anzolin Begotti, Avenida 4A n 926, Bela Vista,
Rio Claro, SP, CEP13506-770, e-mail: yahoo.com.br> and Laura Frey Landesmann, Rua
Turquesan0 140, Condominio Nova Higien6polis, Jandira,
SP, e-mail: .


Altmann, J. 1974. Observational study of behavior: sam-
pling methods. Behaviour 49: 227-267.
Argel de Oliveira, M. M. 1995. Aves e vegetagao em um
bairro residential da cidade de Sao Paulo (Sao Paulo,
Brasil). Rev. Brasil. Zool. 12(1): 81-92.
Bicca-Marques, J. C., Silva, V. M. e Gomes, D. F. 2006.
Ordem Primates. Em: Mamiferos do Brasil, Reis, N. R.,
Peracchi, A. L., Pedro, W. A., Lima, I. P. (eds.), 437 p.
Costa, L. P., Leite, Y. L., Mendes, S. L. e Ditchfield, A. D.
2005. Mammal Conservation in Brazil. Cons. Biol. 19(3):
Lyra-Neves, R. M., Oliveira, M. A., Telino-Jdnior, W. R.,
e Santos, E. M. 2007. Comportamentos interespecificos
entire C .-' jacchus (Linnaeus) (Primates, Callitrichi-
dae) e algumas aves de Mata Atlintica, Pernambuco,
Brasil. Rev. Brasil. Zool. 24 (3): 709-716.
Mendes Pontes, A. R. e Soares, M. L. 2005. Sleeping sites
of common marmosets (C .-' jacchus) in defaunat-
ed urban forest fragments: a strategy to maximize food
intake. J. Zool. 266: 55-63.
Mendes Pontes, A. R., Malta, A. e Asfora, P. H. 2006. A
new species of capuchin monkey, genus Cebus Erxleben
(Cebidae, Primates): found at very brink of extinction in
the Pernambuco Endemism Center. Zootaxa 1-12.
Mendes Pontes, A. R., Normande, I. C., Fernandes, A. C.,
Ribeiro, P. F. e Soares, M. L. 2007. Fragmentation causes
rarity in common marmosets in the Atlantic Forest of
northeastern Brazil. Biodiv. Conserv. 16(4): 1175-1182.
Miranda, G. H. e Faria, D. S. 2001. Ecological aspects of
black-pincelled-marmoset (C .-' penicillata) in the
Cerradao and dense Cerrado of the Brazilian Central Pla-
teau. Braz. J. Biol. 61(3): 397-404.
Olmos, E e Martuscelli, P. 1995. Habitat and distribution
of the Buffy-tufted-ear marmoset C .-' aurita in
Sao Paulo State, Brazil, with notes on its natural history.
Neotrop. Primates 3(3): 75-79.
Ruiz-Miranda, C. R., Affonso, A. G., Morais, M. M.,
Verona, C. E., Martins, A. e Beck, B. 2006. Behavioral
and ecological interactions between reintroduced golden
lion tamarins (Leontopithecus rosalia Linnaeus, 1766) and
introduced marmosets (C .- spp, Linnaeus, 1758) in
Brazil's Atlantic coast forest fragments. Brazilian Archives
of Biology and Technology 49(1): 99-109.
Sinclair, K. E., Hess, G. R., Moorman, C. E., e Mason, J.
H. 2005. Mammalian nest predators respond to green-
way width, landscape context and habitat structure.
Landscape and Urban Pl,'mizng 71: 277-293.


Oscar Humberto Marin-Gdmez


El genero Aotus comprende un grupo de primates amplia-
mente distribuidos en el neotr6pico que habitan en bosques
primaries y secundarios hasta los 3,200 msnm (Wright,
1994). En Colombia se encuentran 5 species de este
genero, tres de ellas agrupadas en el complejo Aotus lemuri-
nus, conformado por las speciess gemelas" A. lemurinus,
A. zonalis y A. griseimembra (Defler, 2003). A. lemurinus
se conoce como mico de noche andino o marteja y habitat
en todo tipo de bosques, fragments de sucesi6n secun-
daria y plantaciones de caf6 de la cordillera de los Andes de
Colombia entire los 1,000 a 1,500 msnm (Defler, 2003).

La mayoria de los primates tienen una dieta que consiste
de frutos, hojas, insects y en menor proporci6n de flores
y nectar (Terborgh, 1983; Wright, 1994). El nectar aporta
nutrients y energia que no se encuentran en otros recur-
sos alimentarios, por lo que su consume ocurre en los pri-
mates como recurso alternative cuando la disponibilidad
de frutos es baja (Janson et al., 1981; Garber, 1988; Ferrari
& Strier, 1992). Los frutos de un gran ndmero de species
de plants de los bosques neotropicales son consumidos
por primates, los cuales pueden actuar como dispersores de
sus semillas (Chapman, 1995, Lambert & Garber, 1998;
Stevenson, 2000), y polinizadores potenciales al no dafiar
las flores (Prance 1980; Assumpcao, 1981; Janson et al.,
1981; Terborgh, 1983; Ferrari & Strier, 1992). El con-
sumo de nectar es raro dentro del genero Aotus (Wright,
1994) y s6lo existen reports paraA. trivirgatus y A. vocifer-
ans (Janson et al., 1981, Puertas et al, 1992). En esta nota
report el consumo de nectar de las flores de Inga edulis por
A. lemurinus y discuto su papel como un possible poliniza-
dor de esta plant en un paisaje ganadero.


Realic6 las observaciones durante el mes de marzo del
2006 en un agroecosistema ganadero, especificamente
en un potrero arborizado con dominancia de irboles de
I. edulis, cercanos a un fragmento de bosque de la reserve
natural "La Montafia del Ocaso", ubicada en el municipio
de Quimbaya, Departamento del Quindio, Colombia
(4o34'81.8"N, 75o51'87., ',.) a una altura entire 975 y
1,100 msnm (CIBUQ, 2006). /. edulis es conocido local-
mente como guamo y es una especie ampliamente utilizada
como irbol de sombrio en la zona cafeteria Colombiana.
Los arboles tienen una altura entire 6 y 20 m y presentan
dos periods de floraci6n al afio, durante los meses de julio
a noviembre y de febrero a junio, mientras que la fructi-
ficaci6n es constant a lo largo del afio con picos en los

Neotropical Primates 15(1), January 2008
meses de diciembre, febrero y junio (Marin, 2006). Las
flores duran un dia, tienen el perianto reducido, numerosos
estambres de color blanco y secretan abundante nectar que
se acumula en el tubo formado por los filamentos estami-
nales; el fruto es una legumbre con numerosas semillas
cubiertas por un arilo dulce y es consumido por loros y
mamiferos (Marin, 2006). Las flores de I. edulis presentan
su antesis a partir de las 13:00, hasta finalizar por com-
pleto a las 19:00, period durante el cual las flores secretan
pequefias cantidades de n&ctar, siendo su producci6n con-
tinua durante la noche y decreciendo en el dia. Los picos de
producci6n de nectar ocurren a las 21:00, 00:00 y 03:00,
durante este period las flores presentan un promedio de
160 pl de nectar, un porcentaje de sucrosa de 16.6 y un
contenido energ6tico de 505 Julios/flor (Marin, 2006).


A las 00:30 del 14 de marzo observe dos individuos de A.
lemurinus alimentindose de frutos de I. edulis. Los monos
prefirieron los frutos maduros que estaban parcialmente
abiertos, ya que despu5s de agarrarlos por el peddnculo con
una mano, con la otra desprendian parcialmente su cubierta
para dejar expuestas las semillas y consumer el arilo que las
cubria. Las semillas fueron dejadas caer al suelo. Ambos
individuos consumieron entire 7 y 10 frutos en un period
de 20 minutes, y despu5s se desplazaron a otro drbol donde
los perdi de vista. Alas 02:20 observe en el mismo drbol dos
individuos, al parecer los mismos observados inicialmente.
Sin embargo esta vez no consumieron frutos, sino que se ali-
mentaron de nectar; los monos probaron entire 5 y 7 flores
en tres minutes y luego se movieron hacia otro drbol de la
misma especie. El 17 de Marzo a las 01:00 observe dos
individuos de A. lemurinus desplazindose entire las ramas
altas de los irboles de I. edulis y emitiendo constantemente
una series de sonidos guturales de baja intensidad, que al
parecer son emitidos frecuentemente cuando los animals
se encuentran cerca de un recurso alimentario (Moynihan,
1964, citado por Defler 2003). Los monos s6lo consumi-
eron el nectar, para lo cual con una mano acercaron una
inflorescencia y probaron el nctar con la lengua tras aproxi-
mar el hocico al tubo floral. De esta forma contactaron
las estructuras reproductivas con en el hocico y la frente,
aunque la cantidad de polen depositado no fue conspicua.
Durante 35 minutes probaron entire 3 y 5 flores por minuto.
Para un individuo contabilic6 un total de 27 flores probadas
legitimamente (haciendo contact con las estructuras repro-
ductivas). Esta actividad finaliz6 cuando empez6 a lover.


Es possible que los individuos de A. lemurinus observados
tengan su lugar de reposo en el bosque, y que su presencia
en el potrero obedezca a la abundancia de alimento nectarr
o frutos) ofrecido por I. edulis en esta zona. Los frutos de
los irboles del genero Inga son un recurso important en la
dieta de varias species de primates neotropicales, en ireas
de bosque mnis no en zonas abiertas (Lambert & Garber,

Neotropical Primates 15(1), January 2008

1998; Stevenson, 2000), sin embargo este es el primer
report del consume de nectar de las plants de este gdnero
por parte de monos.

Los sistemas de polinizaci6n en mamiferos no voladores
incluyen generalmente a species de tamafo pequefo como
roedores, marsupiales y prosimios (Gautier & Marsels,
1994). En Africa hay evidencia indirecta de la polinizaci6n
por los primates de los generos Cercopithecus y Cercocebus en
dos species de leguminosas, Parkia sp. y Daniella pynaertti
(Gautier & Marsels, 1994), mientras que en el Neotr6pico
hay una mayor cantidad de registros de consumo de nectar
por primates (Terborgh, 1983), los cuales pueden causar
pequefos dafos a las flores o destruirlas totalmente (Ter-
borgh 1983, Garber 1988), o por el contrario pueden no
hacer dafo alguno a la flor y consumer el nectar de forma
legitima, actuando como posibles polinizadores (Prance,
1980; Assumpcao, 1981; Janson et al., 1981; Terborgh,
1983; Ferrari & Strier, 1992).

Janson et al. (1981) mencionaron que las flores erectas dis-
puestas en inflorescencias terminales, de perianto fusionado
en forma de copa, con estambres numerosos, largos y exer-
tos, la producci6n sincr6nica de abundante nectar y la ten-
dencia a producer frutos grandes, son rasgos que facilitan
la transferencia de polen por mamiferos no voladores en el
Neotr6pico. Estas caracteristicas coinciden parcialmente
para todas las species del gdnero Inga, en particular para
las species de flores grandes como I. edulis. Sin embargo
las flores de estas plants presentan un perianto reducido y
el nectar se acumula en un tubo formado por los filamentos
estaminales (Koptur, 1983). Las plants de este gdnero no
muestran un syndrome de polinizaci6n hacia un solo tipo
de vector, pues son polinizadas por insects, aves y murci&
lagos (Koptur, 1983; Marin 2006). Aunque las flores de /.
edulis presentan antesis crepuscular y produce la mayor
cantidad de nectar en la noche, presentan una baja tasa de
visits por parte de animals nocturnos, principalmente
de insects (escarabajos y polillas), que actdan mais como
ladrones de polen y de nectar (Marin, 2006). El nectar se
acumula durante la noche y es disponible en el dia a una
gran variedad de insects y aves, los cuales constituyen sus
principles "polinizadores" (Marin, 2006).

Al parecer los A. lemurinus no son visitantes florales fre-
cuentes de esta especie, pero por su comportamiento de
forrajeo, especialmente al tomar el nectar, probar varias
flores de un mismo drbol y moverse entire irboles vecinos
en busca de flores y frutos, podrian estar desempefando un
papel important como polinizadores potenciales de esta
plant, aunque esto debe validarse con observaciones mas
detalladas en otras localidades de Colombia. La evidencia
de polinizaci6n por primates corresponde generalmente a
observaciones indirectas de este process, dependiendo la
efectividad de la polinizaci6n de la morfologia floral, la
frecuencia de uso, el contact con las parties reproducti-
vas, la carencia de dafo a las flores, el trasporte de polen y

el acceso limitado a un determinado ndmero de visitantes
(Gautier & Marsels, 1994). Es probable que en este caso
A. lemurinus favorezca mais la polinizaci6n cruzada de I/.
edulis que la geitonogamia, pues esta 61tima no es conve-
niente ya que todas las plants de este genero son autoin-
compatibles (Koptur, 1984). I. edulis es una especie opor-
tunista con rasgos florales que permiten la polinizaci6n por
una amplia diversidad de visitantes florales como insects,
aves y mamiferos (Marin, 2006). Son irboles comunes
que constituyen una fuente important de alimento para la
fauna y puedan atraer a una gran variedad de visitantes del
bosque, por lo cual deben considerarse como una especie
de interns en los proyectos silvopastoriles, para el manejo
de los agroecosistemas o en la adecuaci6n de corredores
biol6gicos en la zona cafeteria; estos drboles son utilizados
en la region por Alouatta seniculus para desplazarse entire
parches de vegetaci6n (Marin, obs. pers).


A Pablo Stevenson y Jairo P&rez-Torres por sus valiosos
aportes, a Diego G6mez y Diana Mendez por sus comen-
tarios al escrito, y al Centro de Investigaciones en Biodiver-
sidad (CIBUQ) y al program de Licenciatura en Biologia
y Educaci6n Ambiental de la Universidad del Quindio por
el apoyo logistico.

Oscar Humberto Marin-G6mez, Licenciatura en Biologia
y Educaci6n Ambiental, Universidad del Quindio AA. 460,
e-mail: .


Assumpcao, C. T. 1981. Cebus apella and Brachyteles arach-
noides (Cebidae) as potential pollinators ofMabeafistulif-
era (Euphorbiaceae). J. Mammal. 62(2): 386-388
Chapman, C. A. 1995. Primate seed dispersal: coevolution
and conservation implications. Evol. Anthrop. 4: 74-82.
CIBUQ. 2006. Reserva Natural "La Montafa del Ocaso".
Consultado en Noviembre 23, 2006. URL: http://www.
Defler, T. R. 2003. Primates de Colombia. Serie Guias de
Campo # 4. Conservaci6n Internacional de Colombia,
Bogota DC.
Ferrari, S. F and K. B. Strier. 1992. Exploitation of Mabea
fistulifera nectar by Marmosets (C .-( flaviceps) and
Muriquis (Brachyteles arachnoids) in South East Brazil. J.
Trop. Ecol. 8(3): 225-239
Garber, P. A. 1988. Foraging decisions during nectar feed-
ing in Tamarin monkeys Saguinus mystax and Saguinus
fuscicollis (Calitrichidae: Primates) in Amazonian Peru.
Biotropica 20: 100-106
Gautier-Hion, A. and F Marsels. 1994. Mutualism be-
tween a leguminous tree and large African monkeys as
pollinators. Behav. Ecol. Sociobiol. 34: 203-210

Janson, C, J. Terborgh and L. H. Emmons. 1981. Non-
flying mammals as pollinating agent in the Amazonian
forest. Biotropica 13: 1-6
Koptur, S. 1983. Flowering phenology and floral biology
of Inga (Fabaceae: Mimosoideae). Systematic Botany 8(4):
Koptur, S. 1984. Outcrossing and pollinator limitation of
fruit set: breeding systems of
Neotropical Inga trees (Fabaceae: Minosoideae). Evolution
38: 1130-1143.
Lambert, J. E. and P. A. Garber. 1998. Evolutionary and
ecological implications of primate seed dispersal. Am. J.
Primatol. 45: 9-28.
Marin, 0. H. 2006. Composici6n, actividad de forrajeo y
"polinizaci6n" de las aves e insects que visitan el guamo
Inga edulis Martius (Fabales: Mimosaceae) en dos agro-
ecositemas del Quindio. Tesis de grado. Licenciatura en
Biologia y Educaci6n Ambiental. Facultad de Educaci6n.
Universidad del Quindio.
Prance, G. T. 1980. A note on the probable pollination of
Combretum by Cebus monkeys. Biotropica 12: 239
Puertas, P., R. Aquino y F Encarnaci6n. 1992. Uso de ali-
mentos y competici6n entire el mono nocturno Aotus vo-
ciferans y otros mamiferos, Loreto, Perd. Folia Amazdnica
4(2): 147-156
Stevenson, P. R. 2000. Seed dispersal by woolly monkeys
(Lagothrix lagothricha) at Tinigua National Park, Colom-
bia: Dispersal distance, germination rates, and dispersal
quantity. Am. J. Primatol. 50: 275-289.
Terborgh, J. 1983. Five new world Primates. A study in
comparative ecology. Princenton University, New Jersey.
Wright, P. C. 1994. The behavior and ecology of the owl
monkey. In: Baer, J. F, R. C.


Tropical Conservation Science is a peer-reviewed open
access new e-journal that publishes original research papers
and state of the art reviews of broad interest to the field
of conservation of tropical forests and of other tropical
ecosystems. The 1st issue was released on March 3rd. You
can consult the issue in www.tropicalconservationscience.
org. The global community of scientists, advanced gradu-
ate students and conservationists working in the tropics are
invited to submit manuscripts for possible publication in
future issues of the e-journal.


International Primate Protection League founder and
director Shirley McGreal was awarded the Order of the

Neotropical Primates 15(1), January 2008

British Empire in December 2007 for her longstanding
commitment to protecting primates. As one of the highest
honors that can be bestowed to an individual by the Queen
of England, the Order of the British Empire observes "dis-
tinguished service to the arts and sciences, public services
outside the Civil Service and work with charitable and wel-
fare organizations of all kinds." Dr. McGreal, involved in
primate rescue since the early 1970s, is known for uncover-
ing illicit trade in primates and doggedly pushing for pros-
ecution of the perpetrators.
Animal Welfare Institute


A high level workshop of experts of .1.. ;:, biology, neu-
roscience, and cognition that will address how the primate
mind relates to other minds through empathy, imitation
and other social cognition. It will be held in Erice, Silicy,
Italy from June 4-7, 2009. For more information and reg-
istration go to http://www.emory.edu/LIVING_LINKS/


Guadalajara Zoo celebrating its 20 years of service and com-
mitment to conservation, the Wild Fauna and Companion
Animal Mexican Institute and the Autonomous Univer-
sity of Guadalajara have the pleasure of inviting you to its
Conferences Cycle about: Primates, A Priority For World
Wide Conservation, which will be held at the Autonomous
University of Guadalajara and Guadalajara Zoo facilities
from November 26-29, 2008. The conference cycle will
address topics such as conservation in the field, veterinar-
ian medicine in primates, agreements and the role of zoos
in primate conservation. If you are interested please e-mail
mmartinez@zooguadalajara.com.mx or zoogdl@zoogua-


The primate p ulr.. .1..,- workshop will be held at Colorado
State University, during September 13th and 14th, 2008. The
topics that the workshop will focus are an introduction to
and overviews of major infectious disease studies (incl. FIV,
prion diseases, tuberculosis, highly pathogenic avian influ-
enza and other "select agents"); and non-human primate
pathology case presentations. If you are interested please
contact Helle Bielefeldt helle.ohmann@colostate.edu

Neotropical Primates 15(1), January 2008


This course will be held at the German Primate Center,
from October 13-16, 2008. The registration deadline is
September 30th,2008. For more information visit the web
site http://www.euprim-net.eu/network/courses/course7.
htm or contact Deike Terruhn terruhn@dpz.eu .


IPS Research Grants: Mary Blair "Habitat modification
and gene flow in Saimiri oerstedii"; Kimberly Dingess
"Social monogamy in the Bolivian gray titi monkey, Cal-
licebus donacophilus: Conflict or cooperation?"; Michelle
Brown "Predicting intergroup relationships"; Macken-
zie Bergstrom "Dominance among female white-faced
Capuchins"; Robin Bernstein "Hormonal correlates of
divergent growth trajectories in male anubis and hama-
dryas baboons"; Cynthia Thompson "Why fight? Adress-
ing the ultimate causes of aggressive intertroop encounters
in white faced saki monkeys (Pithecia pithecia)"; Lauren
Brent "Sociability, coalitionary support and stress in
female rhesus monkeys". Charles Southwick Conservation
Education Commitment Award: Pierrot Mbonzo (PASA);
Mugabe Rbert and Byamukama Lawrence (Jane Good-
all Institute). Lawrence Jacobsen Education Development
Award: Sian SaraWaters "Barbary Macaque Conservation
in the Rif (BMCRif) Education and Awareness Rais-
ing Among School Age Children"; Allai Orimba "Grass
root Empowerment for the Conservation of Papio anubis
(Olive Baboon) in Kajulu Hills-Kenya"; Tricia Parish
"Identifying the CITES Appendix I listed Asian slow loris
(Nycticebus): a training program for enforcement officials
and rescue centers in Southeast Asia"; Corrin La Combe
"Completing the Conservation: Modifcation and Evalu-
ation of Hooklock Gibbon (Hoolock hoolock) Conserva-
tion Education Program in Bangladesh". IPS Captive Care
Grants: Doug Cress/Claire Coulson (PASA) "Enclosure
Project for Endangered Guenons at the CERCOPAN Pri-
mate Sanctuary in Nigeria"; Noruki Morimura "Reducing
Aggression in Multi-male and Multi-female Chimpanzee
Group Formation at the Chimpanzee Sanctuary UTO,
Japan"; Suzanne Turnock "The effect of environmental
enrichment on the locomotive, feeding and social behav-
iour of spider monkeys (Ateles '. .. '..i in the Zool6gico
Nacional, Honduras". IPS Conservation Grants: Juliet
Wright "The Lebialem Hunters' Beekeeping Initiative in
Southwest Cameroon"; Cuozzo Frank "An Ethnoprima-
tological Approach to Assessing Feral Animal Predation
of Endangered Wild Lemurs and Domestic Livestock at
Beza Mahafaly, Madagascar"; Hsio Sara "Evaluation and

monitoring of crop raiding mitigation strategies in vil-
lages around Budongo Forest Reserve, Uganda"; Riccardo
Pansini "Stress behaviour and personalities of wild vervet
monkeys (Chlorocaebus aethiops)"; James Robins "Assess-
ing the impact of anthropogenic activities on the distri-
bution of proboscis monkeys (Nasalis larvatus) in Danau
Sentarum National Park, West Kalimantan, Indonesia.";
Suzanne Hagell "Assessing Landscape Connectivity for
Spider Monkeys (Ateles _. ..' .., in Southwestern Nicara-
gua"; Tumukunde Alex "Expectations of Local Commu-
nities towards Ecotourism and its Potential for Conserva-
tion of Mountain Gorillas and Forest Habitats of Bwindi
Impenetrable National Park". Martha J Galante Award:
Ndeloh Etiendem.


Manipulative Monkeys: The Capuchins of Lomas Barbu-
dal, by Susan Perry and Joseph H. Manson. 2008. Har-
vard University Press. 368pp. ISBN: 978-0674026643.
Manipulative Monkeys takes us into a Costa Rican forest
teeming with simian drama, where since 1990 primatolo-
gists Susan Perry and Joseph H. Manson have followed the
lives of four generations of capuchins. What the authors
describe is behavior as entertaining -and occasionally as
alarming- as it is recognizable: the competition and coop-
eration, the jockeying for position and status, the peaceful
years under an alpha male devolving into bloody chaos,
and the complex traditions passed from one generation to
the next. Interspersed with their observations of the mon-
keys' lives are the authors' colorful tales of the challenges
of tropical fieldwork -a mixture so rich that by the book's
end we know what it is to be a wild capuchin monkey or
a field primatologist. And we are left with a clear sense of
the importance of these endangered monkeys for under-
standing human behavioral evolution. Contents: 1. All
in a day's work; 2. The social intelligence debate and the
origins of the Lomas Barbudal monkey project; 3. The
challenges of foraging and self-medication; 4. Predators,
prey and personality; 5. Capuchin communication; 6. Two
females' political careers; 7. The career of and alpha male;
8. The strategies of incoming alpha males; 9. Lethal aggres-
sion and the importance of allies; 10. Capuchin mothers,
infants and babysitters; 11. Innovation and tradition in the
creation of blood-testing rituals; 12. Social learning and the
roots of culture; 13. Conservation of tropical dry forest.

Conservation andSustainable Use: A Ha book ofTechniques,
by E.J. Milner-Gulland and J. Marcus Rowcliffe. 2008.
Oxford University Press. 320pp. ISBN: 978-0198530350.
Conservation and Sustainable Use provides a practical and
integrated approach to carrying out research on the con-
servation of exploited species. It is relevant to both tropical

and temperate biomes and is applicable to all exploited spe-
cies, including mammals, fish and plants. It describes both
the practical (field) and theoretical (modeling) techniques
for obtaining and interpreting information, integrating bio-
logical, social, economic and institutional analyses. It also
demonstrates how to translate information into effective
action through appropriate interventions, from legislation to
changing people's attitudes. Contents: 1. Introduction; 2.
Techniques for surveying exploited species; 3. Understand-
ing natural resource users' incentives; 4. Assessing current
sustainability of use; 5. Developing predictive models;
6. Choosing management approaches; 7. Implementing
management for long-term sustainability.

Economics ofPoverty, Environment and Natural Resource Use,
edited by Rob B. Dellink and Arjan Ruijs. 2008. Springer.
218pp. ISBN: 978-1402083037. This book contributes to
an improved understanding of the economic dimensions
of environmental and natural-resource management and
poverty alleviation. The ten chapters of the book offer an
overview of the current knowledge concerning the relation
between poverty, environment and natural-resource use.
Three sides of the debate receive particular attention. First,
the relation between resource use and poverty is discussed
from a theoretical point of view. Second, it is questioned
whether payments for environmental services or consider-
ing values of resources can be an effective tool for stimulat-
ing both sustainable resource use and poverty alleviation.
Third, alternative strategies to break the land degradation
poverty cycle are discussed. Contents: 1. Introduction A.
Ruijs, R. B. Dellink, and D. W. Bromley; 2. Poverty traps
and resource dynamics in smallholder agrarian systems
- C. B. Barrett; 3. Water resource management and the
poor P. Hellegers, K. Schoengold and D. Zilberman; 4.
The role of measurement problems and monitoring PES
schemes G. Meijerink; 5. Can ecotourism be an alterna-
tive to traditional fishing? An analysis with reference to the
case of the Saloum Delta (Senegal) 0. Sarr, J. Boncoeur,
M. Travers and M. C. Cormier-Salem; 6. Effects of poverty
on deforestation: distinguishing behavior from location
- A. Pfaff, S. Kerr, R. Cavatassi, B. Davis, L. Lipper, A.
Sanchez and J. Timmins; 7. Willingness to pay for system-
atic management of community forest for conservation of
non-timber forest products in Nigeria's rainforest region:
implications for poverty alleviation N. A.Chukwuone
and C. E.Okorji; 8. Traditional institutions and sustainable
livelihood: evidences from upland agricultural communi-
ties in the Philippines M. Omura; 9. Farmers investing
in sustainable land use at a tropical forest fringe, the Philip-
pines M. R. Romero and W. T. de Groot; 10. A bargain-
ing model of migration: getting the permission of the farm
household A. Mensah-Bonsu and K. Burger.
H,.. Remote Sensing of Tropical and Sub-tropical
Forest, edited by M. Kalacska and A. Sanchez-Azofeifa.
2008. CRC. 352pp. ISBN: 978-1420053418. While fre-
quently used in temperate environments, hyperspectral
sensors and data are still a novelty in the tropics. Exploring
the potential of hyperspectral remote sensing for assessing

Neotropical Primates 15(1), January 2008

ecosystem characteristics, Hyperspectral Remote Sensing of
Tropical and Sub-Tropical Forests focuses on the complex
and unique set of challenges involved in using this tech-
nology and the data it provides. Contents: 1. Tropical dry
forest phenology and discrimination of tropical tree spe-
cies using hyperspectral data K. L. Castro-Esau and M.
Kalacska; 2. Remote sensing and plant functional groups:
physiology, ecology and spectroscopy in tropical systems -
M. Alvarez-Afiorve, M. Quesada and E. de la Barrera; 3.
Hyperspectral dara for assessing carbon dynamics and bio-
diversity of forest R. Lucas, A. Mitchell and P. Bunting;
4. Effects of soil type on plant growth. Leaf nutrient/Chlo-
rophyll concentration and leaf reflectance of tropical tree
and grass species J. C. Calvo-Alvarado, M. Kalacska, G.
A. Sanchez-Azofeifa and L. S. Bell; 5. Spectral expression
on gender: a pilot study with two dioecious neotropical
tree species J. P. Arroyo-Mora, M. Kalacska, B. L. Cara-
ballo, J. E. Trujillo and 0. Vargas; 6. Species classification
of tropical tree leaf reflectance and dependence on selection
of spectral bands B. Rivard, G. A. Sanchez-Azofeifa, S.
Forey and J. C. Calvo-Alvarado; 7. Discriminating Sirex
noctilio attack in pine forest plantations in south Africa
using high spectral resolution data- R. Ismail, 0. Mutanga
and F Ahmed; 8. Hyperspectral remote sensing of exposed
wood and deciduous trees in seasonal tropical forests S.
Bohlman; 9. Assessing recovery following selective logging
of lowland tropical forests based on hyperspectral imagery
- J. PArroyo-Mora, M. Kalacska, R. L. Chazdon, D. L.
Cirvco, G. Obando-Vargas, and A. A. S. Hernindez; 10.
Technique for reflectance calibration of airbone hyperspec-
tral spectrometer data using a broad multiband radiom-
eter T. Miura, A. R. Huete, L. G. Ferreira, E. E. Sano
and H. Yoshioka; 11. Assessment of phenologic variability
in Amazon tropical rainforests using hyperspectral hype-
rion and MODIS satellite data A. R. Huete, Y. Kim, P.
Ratana, K. Didan, Y. E. Shimabukuro and T. Miura; 12.
Hyperspectral remote sensing of canopy chemistry, physi-
ology and biodiversity in tropical rainforests G. P. Asner;
13. Tropical Remote Sensing-Opportunities and challenges
- J. A. Gamon.

Aguiar, L. M., Pie, M. R. and Passos, F C. 2008. Wild
mixed groups of howler species (Alouatta caraya and Al-
ouatta clamitans) and new evidence for their hybridiza-
tion. Primates 49(2): 149-152.
Aldrich, B. C., Molleson, L. and Nekaris, K. A. I. 2008.
Vocalizations as a conservation tool: an auditory survey
of the Andean titi monkey Callicebus oenanthe Thomas,
1924 (Mammalia: Primates: Pitheciidae) at Tarangue,
northern Peru. Contrib. Zool. 77(1): 1-6.
Ange-van Heugten, K., Timmer, S., Jansen, W. L. and Ver-
stegen, M. W. A. 2008. Nutritional and health status of
woolly monkeys. Int. J. Prim. 29(1): 183-194.
Arroyo-Rodriguez, V., Asensio, N. and Cristobal-Azkarate,
J. 2008. Demography, life history and migrations in a

Neotropical Primates 15(1), January 2008
Mexican mantled howler group in a rainforest fragment.
Am. J. Primatol. 70(2): 114-118.
Arroyo-Rodriguez, V., Mandujano, S. and Benitez-Malvi-
do, J. 2008. Landscape attributes affecting patch occu-
pancy by howler monkeys (Alouattapalliata mexicana) at
Los Tuxtlas, Mexico. Am. J. Primatol. 70(1): 69-77
Babb, P. L., Gagneux, P., Fernandez-Duque, E. and Schurr,
T. G. 2008. Genetic variation and population structure
in the owl monkey, Aotus azarai. Am. J. Phys. Anthropol.
Supp146: 62.
Baker, M. E. 2008. Use of a mirror as a data collection aid.
Lab. Prim. Newsletter. 47(1): 9-10.
Bezanson, M., Garber, P. A., Murphy, J. T. and Premo, L.
S. 2008. Patterns of subgrouping and spatial affiliation
in a community of mantled howling monkeys (Alouatta
palliata). Am. J. Primatol. 70(3): 282-293.
Bezerra, B. M., Barnett, A. A., Souto, A. and Jones, G.
2008. Predation by the tayra on the common marmoset
and the pale-throated three-toed sloth. J Ethol: online
Caesar, C., Franco, E. S., Soares, G. C. N. and Young, R.
J. 2008. Observed case of maternal infanticide in a wild
group of black-fronted titi monkeys (Callicebus nigri-
frons). Primates. 49(2): 143-145.
Casar, C. and Young, R. J. 2008. A case of adoption in a
wild group of black-fronted titi monkeys (Callicebus nig-
rifrons). Primates. 49(2): 146-148.
Cecilia, V., Elena, B., Laura, D. and Paolo, A. 2008. Sys-
tematic revision of the neotropical primates in the prima-
tological collection of the Natural History Museum 'La
Specola', Florence. Folia. Primatol. 79(3): 153.
Chalk, J., Wright, B. W., Lucas, P. W., Verderane, M. P.,
Fragaszy, D., Visalberghi, E., Izar, P. and Ottoni, E. B.
2008. The mechanical properties of foods processed by
Cebus libidinosus at Boa Vista, Brazil. Am. J. Phys. Anthro-
pol. Suppl 46: 77.
Chollet, M. B. and Teaford, M. E 2008. Ecological stress
and linear enamel hypoplasia in Cebus. Am. J. Phys. An-
thropol. Suppl 46: 78-79.
Clarke, M. R. and Glander, K. E. 2008. Natal emigration
by both sexes in the La Pacifica population of mantled
howlers: when do some stay?. Am. J. Primatol. 70(2):
Covey, R. M. 2008. Prehensile tail use in white-faced capu-
chins Cebus capuchinus at La Suerte Biological Field Sta-
tion, Costa Rica. Am. J. Phys. Anthropol. Suppl 46: 82.
Derby, A. M. 2008. The interrelationship between popula-
tion structure, forest ecology and behavior in red howler
monkeys (Alouatta seniculus). Am. J. Phys. Anthropol.
Supp146: 89.
Di Fiore, A., Link, A. and Spehar, S. N. 2008. Multiple
males sire offspring in groups of wild spider monkeys
(Ateles belzebuth). Am. J. Phys. Anthropol. Suppl 46: 90.
Ehmke, E. and Boinski, S. 2008. Proximity patterns of
female brown capuchins in Suriname are inconsistent
with expectations of female-bonded primates. Am. J.
Phys. Anthropol. Supply 46: 94.

Fernandez-Duque, E., Juarez, C. P. and Di Fiore, A. 2008.
Adult male replacement and subsequent infant care by
male and siblings in socially monogamous owl monkeys
(Aotus azarai). Primates. 49(1): 81-84.
Field, M. Y. 2008. So hot you could fry a monkey: be-
havioral thermoregulation in human-commensal white-
fronted capuchins (Cebus ,lb.j'o~'e1. Am. J. Phys. Anthro-
pol. Suppl 46: 95-96.
Forgey, K., Scalzitti, J., Rizzo, K., Garza, J. and Stahler, A.
2008. Molecular genetic study of non-invasively collected
samples from Alouatta pigra at the Calakmul Biosphere
Reserve, Mexico. Am .J. Phys. Anthropol. Suppl 46: 97.
Garber, P. A., Gomes, D. E and Bicca-Marques, J. C. 2008.
Experimental field study of hand preference in wild
black-horned (Cebus nigritus) and white-faced (Cebus ca-
pucinus) capuchins: evidence for individual and species
differences. Anim. Cognition. (Advance online publica-
tion): online (1-11).
Garber, P. A., Gomes, D. E and Bicca-Marques, J. C. 2008.
Pull or push? Experimental field study of problem-solv-
ing using tools in wild capuchins (Cebus nigritus). Am. J.
Phys. Anthropol. Suppl 46: 101.
Gibson, K. N., Vick, L. G., Palma, A. C. and del Rocio
Carrasco, E M. 2008. Intra-community infanticide and
forced copulation in spider monkeys: a multi-site com-
parison between Cocha Cashu, Peru and Punta Laguna,
Mexico. Am. J. Phys. Anthropol. Suppl 46: 103.
Gomes, M. T., Baccarella, A. and Sineo, L. 2008. Conser-
vation activities for southern muriqui in Brazil's Atlantic
Forest. Folia Primatol. 79(3): 155.
Guillot, D. M. 2008. A video based study of wild Ate-
line positional behavior. Am. J. Phys. Anthropol. 46:
Gunst, N., Boinski, S. and Fragaszy, D. M. 2008. Acqui-
sition of foraging competence in wild brown capuchins
(Cebus apella), with special reference to conspecifics' for-
aging artefacts as an indirect social influence. Behaviour.
145(2): 195-229
Hanson, K. L. 2008. Juvenile-directed aggressive and ago-
nistic interactions in Alouatta palliata, mantled howler
monkeys. Am. J. Phys. Anthropol. Suppl 46: 112.
Jones, C. B., Milanov, V. and Hager, R. 2008. Predictors of
male residence patterns in groups of black howler mon-
keys. J. Zool. 275(1): 72-78.
Jost, C. A. 2008. Effects of habitat characteristics on the
behavioral ecology and habitat use patterns of mantled
howler monkeys, Alouatta palliata. Am. J. Phys. Anthro-
pol. Suppl 46: 127.
Kowalewski, M., Garber, P. A., Peker, S. M. and Zunino,
G. E. 2008. Reproductive competition and ambush be-
haviors in black and gold howler monkeys. Am. J. Phys.
Anthropol. Suppl 46: 134.
Martins, M. M. 2008. Fruit diet of Alouatta guariba and
Brachyteles arachnoides in Southeastern Brazil: comparison
of fruit type, color, and seed size. Primates. 49(1): 1-8.
Matsuda, I. and Izawa, K. 2008. Predation of wild spider
monkeys at La Macarena, Colombia. Primates. 49(1):

Matthews, L. J. 2008. Ranging behavior of white-fronted
capuchins (Cebus alb.f on3) in the Ecuadorian Amazon:
effects of resource use and intergroup interactions. Am. J.
Phys. Anthropol. Supply 46: 151.
McKinney, T. and Orozco Zamora, C. 2008. Crop-raiding
of mango fruits, Mangifera indica, by mantled howlers
(Alouatta palliata) in the Refugio de Vida Silvestre Curu,
Costa Rica. Am. J Phys. Anthropol. Supply 46: 153.
Melin, A. D., Fedigan, L. M., Hiramatsu, C. and Kawa-
mura, S. 2008. Polymorphic color vision in white-faced
capuchins (Cebus capucinus): Is there foraging niche
divergence among phenotypes?. Behav. Ecol. Sociobiol.
62(5): 659-670.
Muniz, L. and Vigilant, L. 2008. Isolation and charac-
terization of microsatellite markers in the white-faced
capuchin monkey (Cebus capucinus) and cross-species
amplification in other New World monkeys. Mol. Ecol.
Resources. 8(2): 402-405.
Nadjafzadeh, M. N. and Heymann, E. W. 2008. Prey for-
aging of red titi monkeys, Callicebus cupreus, in compari-
son to sympatric tamarins, Saguinus mystax and Saguinus
fuscicollis. Am. J Phys. Anthropol. 135(1): 56-63.
Notarnicola, J., Pinto, C. M. and Navone, G. T. 2008.
Host occurrence and geographical distribution of Dipeta-
lonema spp. (Nematoda: Onchocercidae) in neotropical
monkeys and the first record of Dipetalonema gracile in
Ecuador. Comparative Parasotol. 75(1): 61-68.
Perrin, E. M. and Kitchen, D. M. 2008. Using proximity
to understand association patterns among adult mantled
howler monkeys (Alouatta palliata). Am. J. Phys. Anthro-
pol. Supply 46: 170.
Perry, S. E., Muniz, L., Manson, J. H., Gros-Louis, J. and
Vigilant, L. 2008. Kin biased social behavior in wild
white-faced capuchin monkeys, Cebus capuchinus. Am. J.
Phys. Anthropol. Supply 46: 171.
Power, M. L., Verona, C. E., Ruiz-Miranda, C. and Ofted-
al, 0. T. 2008. The composition of milk from free-living
common marmosets (C .-.'. jacchus) in Brazil. Am. J.
Primatol. 70(1): 78-83.
Raguet-Schofield, M. 2008. The effects of human en-
croachment and seasonality on the risk of mantled
howler monkey (Alouatta palliata) predation by dogs
on Ometepe Island, Nicaragua. Am. J. Phys. Anthropol.
Suppl46: 176.
Rodrigues, M. A. 2008. The emergence of sex-segregated
association patterns in juvenile spider monkeys (Ateles
-. ....', Am. J. Phys. Anthropol. Supply 46: 181-182.
Rosales-Meda, M., Estrada, A. and L6pez, J. E. 2008.
Demographic survey of black howler monkey (Alouatta
pigra) in the Lachua Eco-region in Alta Verapaz, Guate-
mala. Am. J. Primatol. 70(3): 231-237.
Sabatini, V. and Ruiz-Miranda, C. R. 2008. Acoustical as-
pects of the propagation of long calls of wild Leontopith-
ecus rosalia. Int. J. Primatol. 29(1): 207-223.
Schrauf, C., Huber, L. and Visalberghi, E. 2008. Do capu-
chin monkeys use weight to select hammer tools?. Anim.
Cognition. (Advance online publication): online(i-10).
Spehar, S. N. and Di Fiore, A. 2008. The role of long-dis-
tance vocalizations in regulating association patterns and

Neotropical Primates 15(1), January 2008
social interactions in white-bellied spider monkeys (Ateles
belzebuth). Am. J. Phys. Anthropol. Supply 46: 199.
Stone, A. I. 2008. Seasonal effects on play behavior in im-
mature Saimiri sciureus in Eastern Amazonia. Int. J. Pri-
matol. 29(1): 195-205.
Tinney, A. and Bezanson, M. 2008. Effects of food exploi-
tation on activity and group size of mantled howlers (Al-
ouattapalliata). Am. J Phys. Anthropol. SuppI 46: 208.
Tolentino, K., Roper, J. J., Passos, E C. and Strier, K.
B. 2008. Mother-offspring associations in northern
muriquis, Brachyteles .'* .... .-:'.. .-4. J. Primatol. 70(3):
Van Belle, S., Strier, K. B. and Estrada, A. 2008. To fight
or to cooperate? Reproductive dilemmas in male black
howler monkeys. Am. J. Phys. Anthropol. Supply 46: 213.
Wallace, R. B. 2008. Towing the party line: territoriality,
risky boundaries and male group size in spider monkey
fission-fusion societies. Am. J. Primatol. 70(3): 271-281.
Webster, T. H. and Suarez, S. A. 2008. Transfer behavior
and association patterns of female spider monkeys (Ateles
belzebuth belzebuth) in Yasuni National Park, Ecuador.
Am. J Phys. Anthropol. Supply 46: 219.
Wheeler, B. C. 2008. Are the alarm calls of tufted capuchin
monkeys (Cebus apella) functionally referential?. Am. J.
Phys. Anthropol. Supply 46: 221.
Wolovich, C. K., Perea-Rodriguez, J. P. and Fernandez-
Duque, E. 2008. Food transfers to young and mates in
wild owl monkeys (Aotus azarai). Am. J. Primatol. 70(3):
Wright, B. W., Wright, K. A., Chalk, J., Verderane, M. P.,
Fragaszy, D., Visalberghi, E., Izar, P. and Ottoni, E. B.
2008. Fallback foraging as a way of life: dietary variability
and skeletal morphology in tufted capuchins. Am. J. Phys.
Anthropol. Supply 46: 225-226.
Young, H., Fedigan, L. M. and Addicott, J. F 2008. Look
before leaping: foraging selectivity of capuchin monkeys
on acacia trees in Costa Rica. Oecologia. 155(1): 85-92.


Selected abstracts from the 30th Meeting of the Ameri-
can Society of Primatologists. Salem, Wisconsin, June
20-23, 2007 http://www.asp.org/asp2007/showConf-

Bales, K. L., J. M. Dietz, A. J. Baker, W. A. Mason & S.
P. Mendoza. Perspectives on cooperative Breeding from
golden lion tamarins and coppery titi monkeys.
Bonnie, K. E. Considering the social in socially-mediated
learning: lessons from capuchins and chimpanzees.
Burrell, A. M. & J. A. French. Variation in early parental
care influences stress reactivity in juvenile Geoffroy's mar-
m osets (C .-' .. .. i
Caine, N. G. Is satiation a factor in time of retirement by
Callitrichid groups?
Caselli, C. B. & E. Z. Setz. Seasonality in long calls by Titi
monkeys (Callicebus nigrifrons) in the Atlantic forest of
southeast Brazil.

Neotropical Primates 15(1), January 2008
Clarke, M. R., R. Ford, D. Arden & K. E. Glander. Survey
of the howler population of la Pacifica, 2006.
Courtney, W., S. McGrath, D. Locke, V. Magrini, M. Hick-
enbotham, J. Armstrong, A. Quinlan, G. Marth & E. R.
Mardis. Use of next generation sequencing technologies
for generating non-human primate genomic resources.
Crast, J. & D. Fragaszy. How to account for the induc-
tion of traditions in captive groups of Capuchin monkeys
(Cebus apella).
Crockett, C. M., K. L. Bentson & R. U. Bellanca. Alo-
pecia and overgrooming in laboratory monkeys vary by
species but not sex, suggesting a different etiology than
Cronin, K. A. Cooperative problem solving by captive
Cotton-top tamarins (Saguinus oedipus): the effects of in-
equitable rewards
De la Torre S., C. T. Snowdon & P. Yepez. Precious vari-
ability: behavior, ecology and conservation of Pygmy
marmosets in Ecuador.
Depeine, C. D., D. E. Bernhards & J. D. Newman. Con-
sistency of vocal behavior during brief separations in
common marmosets.
Dindo, M., B. Thierry, E B. de Waal & A. Whiten. Social
transmission of experimental foraging techniques in Ca-
puchin monkeys (Cebus apella).
Drea, C. M. How to study socially biased learning in pri-
mates? Trends and future directions.
Ginther, A. J. & C. T. Snowdon. Like father, like son: social
strategies of cottontop tamarin family males (Saguinus o.
oedipus) in two life history stages.
Gunst, N., S. Boinski & D. M. Fragaszy. Studying how
social context aids acquisition of foraging skills in wild
brown Capuchins (Cebus apella) in Suriname.
Hankerson, S., M. Pitts, J. Dietz & B. Raboy. A test of
exploitative and interference competition in wild golden-
headed Lion tamarins (Leontopithecus chrysomelas)..
Hostetler, C. M., S. P. Mendoza, W. A. Mason & K. L.
Bales. Niii. .,d. .i-iri. .1. .;,- of alloparental care in Titi
monkeys (Callicebus cupreus).
Huntsberry, M. E., P. G. Roma, C. J. Christensen, A. M.
Ruggiero & S. J. Suomi. Token exchange and the selec-
tive-value effect in Capuchin monkeys (Cebus apella).
Jarcho, M. R., S. P. Mendoza, W. A. Mason, S. A. Blozis &
K. L. Bales. Neuroendocrinology of maternal behavior in
captive Titi monkeys (Callicebus cupreus).
Kurdziel, L. B. & P. G. Judge. Picture recognition of food
in brown Capuchin monkeys (Cebus apella).
Lilak, A. L. & K. A. Phillips. Consistency of hand prefer-
ence across unimanual and bimanual tasks in Capuchin
monkeys (Cebus apella).
Long, L. A., S. Gibson, L. Williams & C. Abee. Owl mon-
keys (Aotus sp.) utilize food directed and non-food di-
rected enrichment items.
Lukas, K. & C. Kuhar. An analysis of efforts to integrate
science and behavioral management of zoo primates.
Martin, A. L., M. A. Bloomsmith, A. W. Clay, M. E. Kelley,
M. J. Marr & T. L. Maple. The role of behavior analysis
in the behavioral management of nonhuman primates.

Matthews, S. A., J. E. Fite, M. Rukstalis, C. N. Ross, A. M.
Burrell & J. A. French. Early infant care, hypothalamic-
pituitary-adrenal (Hpa) axis activity, and pubertal timing
in a cooperatively breeding New World primate, Wied's
black tufted-ear marmoset (C .-( kuhlii).
Pappano, D. J., M. Ayers, D. Cairnes, M. C. May & J.
R. Kaplan. Unrelated females housed in unisexual groups
interfere in fights in a male-like pattern that is unassoci-
ated with affiliation and that tends to prolong aggressive
Phillips, K. A. & W. D. Hopkins. Cerebellar asymmetry
and handedness in chimpanzees (Pan troglodytes) and Ca-
puchins (Cebus apella).
Pickering, T., J. Pan, B. von Ammon, E. H. Kennedy & D.
Fragaszy. A Capuchin monkey masters mazes.
Pokorny, J. J. & E B. de Waal. Capuchin monkeys (Cebus
apella) demonstrate knowledge of social group members
via an oddity task.
Prudom, S. L., J. R. Gorst & T. E. Ziegler. The role of
prolactin and testosterone on male paternal care in the
common marmoset (C .-('. jacchus).
Rapaport, L. Socially mediated learning about food and
foraging in wild golden Lion tamarins (Leontopithecus
Rivera, J., C. K. Wolovich & S. Evans. Will work for food:
foraging behavior of owl monkeys (Aotus nancymaae).
Rukstali, M. Plasticity, development, and the social envi-
ronment: vocal communication in the marmoset.
Savage, A., T. E. Ziegler & J. A. French. Chuck Snowdon's
contributions to Callitrichid biology: communication,
reproduction, and conservation.
Siani, J. M. & J. M. Dietz. Costs of providing infant care in
wild golden Lion tamarins at Pogo Das Antas Biological
Reserve, Brazil.
Smith, A. S., K. R. Lane, A. K. Birnie & J. A. French.
Functional significance of sexually dimorphic acoustic
structures in Wied's black tufted-ear marmoset (Cal-
lithrix kuhlii).
Tapanes, E., C. K. Wolovich & S. Evans. Leaping into the
night: when owl monkeys (Aotus sp.) awake
Thomas, S. J., M. L. Schwandt, S. G. Lindell, S. J. Suomi,
C. S. Barr &J. D. Higley. Heritability of dominance rank
in laboratory-housed juvenile nonhuman primates.
Williams, S. H., C. J. Vinyard, K. E. Glander, M. F Tea-
ford, M. Deffenbaugh & C. L. Thompson. EMG telem-
etry in free-ranging primates: pilot data from howling
monkeys (Alouattapalliata) at La Pacifica, Costa Rica.
Wolovich, C. K., S. Evans & J. A. French. Dads don't pay
for sex but do buy the milk: food sharing and reproduc-
tion in Owl monkeys (Aotus nancymaae).
Ziegler, T. E. & C. T. Snowdon. Proximate mechanisms of
cooperative breeding in Cotton-Top tamarin and Mar-
moset monkeys by chemical communication.
Zucker, E. & M. Clarke. Proximities to specific adult males
by adult female howling monkeys (Alouatta palliata) in
Costa Rica.


31"s Meeting of the American Society of Primatologists.
June 18 21, West Palm Beach, Florida. For more informa-
tion visit the web site: http://www.asp.org/asp2008/index.
htm or contact Matthew Novak novakm@mail.nih.gov

II Congress Colombiano de Primatologia. Junio 26-28,
Universidad Nacional de Colombia, Bogoti D.C. El II
Congress Colombiano de Primatologia organizado por la
Asociaci6n Colombiana de Primatologia (ACP) tiene como
objetivos el intercambio de conocimientos y experiencias en
el area de la Primatologia, el establecimiento de redes de
trabajo (alianzas) en busca del desarrollo cientifico en los
pauses neotropicales y el mejoramiento de la calidad de vida,
no solamente de las species de primates sino tambien sus
ecosistemas. Durante el congress se abarcarin tres grandes
tematicas: conservaci6n in situ, conservaci6n ex situ y cien-
cias aplicadas, sociales y humans. La recepci6n de trabajos
finalizard el 30 de Mayo de 2008. Para mayores informes
visitar la pigina http://primatologiacolombia.org/index.
htm l o escribir a con,, .''. priiii u..l..1 ..l..i l-. i ..i_

22nd Annual Meeting of the Society of Conservation
Biology. July 13-17, the Chattanooga Convention Center,
Chattanooga, Tennessee, USA. The chair of the meeting
will be Dr. David A. Aborn, from the Department of Bio-
logical and Environmental Sciences, University of Tennes-
see at Chattanooga. As evidenced by several previous meet-
ing themes, there are connections among many aspects of
the environment and its conservation, and recognizing
those connections is critical for achieving the goals of con-
servation biology. To that end, the theme for the 2008
SCB annual meeting T.'ovw the mountains to the sea"will
examine several major ecosystems, both as separate compo-
nents and as a connected entity. For more information and
submission dates, go to: www.conbio.org/2008/

XXII1d IPS Congress. August 3 8, Edinburgh Inter-
national Conference Center, Edinburgh, Scotland. It
will be sponsored by the Primate Society of Great Britain.
Abstracts for oral and poster presentations must be submit-
ted by 30th November 2007. For information consult the
web page: http://www.ips2008.co.uk/index.html

XX"' International Congress of Zoology. August 26th
to 29th 2008, Jussieu Grand Campus, Paris, France. Held
every four years, the International Congress of Zoology
gives zoologists and professionals from all fields related to
zoology the chance to come together to discuss the cur-
rent status of zoology and to share ideas about future devel-
opment of all its disciplines. The International Society of
Zoological Sciences (ISZS) and the Societe Zoologique

Neotropical Primates 15(1), January 2008
de France invite you to the XX International Congress
of Zoology, that will be co-hosted by the Universities Pierre
et Marie Curie (PVI), Paris-Sud (PXI) and the Museum
National d'Histoire Naturelle. Registration opened on 1st
December 2007. For information about the Congress,
please visit: http://icz2008.snv.jussieu.fr or http://www.

6th International Conference on Methods and Techniques
in Behavioral Research. August 26 to 29, Maastricht, The
Netherlands. The theme of the 6th international confer-
ence on methods and techniques in behavioral research will
be Measuring Behavior. This conference is the premier inter-
disciplinary event for scientists and practitioners concerned
with the study of human or animal behavior. This unique
conference focuses on methods, techniques and tools in
behavioral research in the widest sense, from behavioral
ecology to neuroscience and from physiology to ergonom-
ics. In doing so, Measuring Behavior responds to a growing
need to cross disciplines and create bridges between them.
For more details go to www.noldus.webaxxs.net/mb2008/

XII Congreso de la Sociedad Mesoamericana para la
Biologia y la Conservaci6n. Noviembre 10-14, Hotel
Real Intercontinental, San Salvador, El Salvador. Es un
event particularmente atractivo para estudiantes y pro-
fesionales ya que es un Foro activo para la biologia de la
conservaci6n y desarrollo sostenible regionales. El envio
de restimenes de ponencias orales y carteles seri hasta el 30
de Junio de 2008. Para mayor informaci6n visitar la pigina


11"' Meeting of the German Society of Primatology.
February 24-26, University of Veterinary Medicine, Han-
nover, Germany. The abstract submission deadline is Sep-
tember 30', 2008. For more information go to http://

Neotropical Primate Husbandry, Research, and Conser-
vation Conference. October 13-15.
Brookfield Zoo is pleased to announce that it will host a
Neotropical Primate Husbandry, Research, and Conserva-
tion Conference. This conference will focus on a variety of
topics pertaining to Neotropical primates like husbandry,
conservation, and emergent issues pertaining to captive and
wild populations. The workshop will include three days of
presentations, a poster session, as well an icebreaker, silent
auction, and banquet. Additional information regarding
registration fees, travel information, and submission of
abstracts will be made available in late 2008. For questions
please contact vince.sodaro@czs.org

Conservation International Primate Publications,
Pocket Guides and Field Guides

Conservation International Order Form. For more information including full descriptions, author lists, and
ISBN numbers, please visit our website: http://www.conservation.org/xp/CIWEB/library/books
All prices include UPS Ground shipping within the continental United States.
For orders requiring faster service than UPS Ground, you will be responsible to pay all shipping costs.
Please email or call the phone number listed below for: overnight deliveries, wholesale orders, international
orders and books in Spanish.
Please complete the following form, print it out and mail or fax to:

Jill Lucena
Conservation International
2011 Crystal Drive, Suite 500
Arlington, VA 22202 USA
E-Mail: j.lucena@conservation.org
Phone: (703) 341-2536
Fax: (703) 553-4817

First Name Last Name

Company Name

Mailing Address City State Country Zip Code

Telephone Fax

E-Mail Address
Order Form should include credit card information or be sent along with check or money order, made payable
to Conservation International. Please allow 2-3 weeks for delivery.
Quantity Desired
Monkeys of the Guianas (x $7.95) = $ South Asian (x $7.95) = $
(Guyana, Suriname, Primates Pocket
French Guiana) Pocket Guide
Monkeys of the Atlantic (x $7.95) = $ Primates of (x $40) = $
Forest of Eastern Brazil Colombia Field
Pocket Guide Guide (English
or Spanish,
please specify
Primates of Amapa (x $7.95) = $ Marmosets and (x $7.95) = $
Pocket Guide Tamarins Pocket
Extended Family: Long (x$20.00) = $ Lemurs of (x $7.95) = $
Lost Cousins by Colin Madagascar
Groves Pocket Guide
Payment Method

Payment Enclosed (check or money order in US $ only) OR

Credit Card: VISA MasterCard
Name (as it appears on card)
Card Number Expiration Date /

A Climate for Life
Mail and Fax Order Form

A Climate for Life by Russell A. Mittermeier,
Michael Totten, Laura Ledwith Pennypacker,
Frederick Boltz, Cristina G. Mittermeier, Guy
Midgley, Carlos Manuel Rodriguez, Glenn
Prickett, Claude Gascon, Peter A. Seligmann,
and Olivier Langrand. 2008.
ISBN: 978-0-9818321-0-4.

Price: $75.00 (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 call the phone number listed below for:
overnight deliveries, wholesale orders, and
international orders.

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

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

First Name Last Name

Company Name

Mailing Address City State Zip Code

Telephone Fax

E-Mail Address

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

Quantity x $75.00 each Total:
Payment Enclosed (check or money order in US $ only)
Charge my credit card: VISA Mastercard
Name (as it appears on card)
Card Number
Expiration Date

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) 345-2852154,
e-mail: , and all Portuguese contribu-
tions to: Jdlio Cfsar Bicca-Marques, Departamento de Biodiversidade
e Ecologia, Pontificia Universidade Cat61lica do Rio Grande do Sul,
Av. Ipiranga, 6681 Pridio 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 compat-
ible text-editors (MS-Word, WordPerfect, Excel, and Access), and/
or e-mailed to (English, Spanish) or
(Portuguese). Hard copies should be supplied
for all figures (illustrations and maps) and tables. The full name and
address for each author should be included. Please avoid abbreviations
and acronyms without the name in full. Authors whose first language
is not English should please have their English manuscripts carefully
reviewed by a native English speaker.

Articles. Each issue of Neotropical Primates will include up to three
full articles, limited to the following topics: Taxonomy, 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.
(Resolution: 300 dpi. Column widths: one-column = 8-cm wide;
two-columns= 17-cm wide). Please keep these to a minimum. We
stress the importance of providing maps that are publishable.
Tables. Tables should be double-spaced, using font size 10, and
prepared with MS Word. Each table should have a brief title.
News items. Please send us information on projects, field sites,
courses, Thesis or Dissertations recently defended, recent 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 etal.,
In the list of references, the title of the article, name of the journal,
and editorial should be written in the same language as they were
published. All conjunctions and prepositions (i.e., "and", "In") should
be written in the same language as rest of the manuscript (i.e., "y"
or "e", "En" or "Em"). This also applies for other text in references
(such as "PhD thesis", "accessed" see below). Please refer to these
examples when listing references:
Journal article
Stallings, J. D. and Mittermeier, R. A. 1983. The black-tailed mar-
moset (Callithrix argentata melanura) recorded from Paraguay. Am.
J. Primatol. 4: 159-163.
Chapter in book
Brockelman, W Y. and Ali, R. 1987. Methods of surveying and
sampling forest primate populations. In: Primate Conservation in
the Tropical Rain Forest, C. W. Marsh and R. A. Mittermeier (eds.),
pp.23-62. Alan R. Liss, New York.
Napier, P. H. 1976. Catalogue of Primates in the British Museum
(Natural History). Part 1: Families Callitrichidae and Cebidae. British
Museum (Natural History), London.
Wallace, R. B. 1998. The behavioral ecology of black spider monkeys
in north-eastern Bolivia. Doctoral thesis, University of Liverpool,
Liverpool, UK.
Muckenhirn, N. A., Mortensen, B. K., Vessey, S., Fraser, C. E. 0. and
Singh, B. 1975. Report on a primate survey in Guyana. Unpublished
report, Pan American Health Organization, Washington, DC.
UNESCO. 2005. UNESCO Man and the Biosphere Programme.
United Nations Educational, Scientific, and Cultural Organisation
(UNESCO), Paris. Website: http://www.unesco.org/mab/index.
htm. Accessed 25 April 2005. ("Acessada em 25 de abril de 2005"
and "Consultado el 25 de abril de 2005" for articles in Portuguese
and Spanish respectively).
For references in Portuguese and Spanish:
"and" changes to "e" and "y" for articles in Portuguese and Spanish
"In" changes to "Em" and "En" for articles in Portuguese and Span-
ish respectively.
"Doctoral thesis" changes to "Tese de Doutoramento" and "Tesis de
Doctorado" for articles in Portuguese and Spanish respectively.
"MSc Thesis" changes to "Dissertaqa.o de Mestrado" and "Tesis de
Maestria" for articles in Portuguese and Spanish respectively.
"Unpublished report" changes to "Relat6rio Tfcnico" and "Reporte
no publicado" for articles in Portuguese and Spanish respectively.



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

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