Title: Neotropical primates
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Title: Neotropical primates a newsletter of the Neotropical Section of the IUCNSSC Primate Specialist Group
Abbreviated Title: Neotrop. primates
Physical Description: v. : ill. ; 27 cm.
Language: English
Creator: IUCN/SSC Primate Specialist Group -- Neotropical Section
IUCN/SSC Primate Specialist Group -- Neotropical Section
Conservation International
Center for Applied Biodiversity Science
Publisher: Conservation International
Place of Publication: Belo Horizonte Minas Gerais Brazil
Belo Horizonte Minas Gerais Brazil
Publication Date: August 2002
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Subject: Primates -- Periodicals -- Latin America   ( lcsh )
Primates -- Periodicals   ( lcsh )
Wildlife conservation -- Periodicals   ( lcsh )
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Dates or Sequential Designation: Vol. 1, no. 1 (Mar. 1993)-
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Neotropical Primates 10(2), August 2002


THE TRUE IDENTITY AND CHARACTERISTICS OF SIMIA ALBIFRONS HUMBOLDT, 1812:
DESCRIPTION OF NEOTYPE

Thomas R. Defler' and Jorge I. Hernandez-Camacho2t

SInstitutoAmazdnico deInvestigaciones (IMANI) de la UniversidadNacionalde Colombia, A.A. 215, Leticia (Amazonas), Colombia.
2 President, Fundacidn Biocolombia, Santa Fe de Bogotd, Colombia.


Abstract

The historic holotypic description of Alexander von Humboldt for the primate Simia j ... contains obvious errors
which have created various taxonomic problems, since it has been impossible to compare descriptions of subspecies with an
original holotype, which was never preserved. An historic taxonomic error was the recognition of Cebus j ... unicolor as a
different taxon from Cebus j. ... ',. ... which we correct in this paper by the recognition of Cebus ... unicolor as
a synonym for Cebus ... albifrons. We describe Cebus ... ',. ... for the first time, based on a neotype collected
by us close to the type locality. Additionally, confusions about the type locality are discussed and clarified. Maypures is
established as the correct type locality. General information on the geographic distribution and natural history of the species
and subspecies is also provided.

Key Words Primates, Cebidae, pale-fronted capuchin monkey, Cebus ... albifrons, Simia ... Humboldt, 1812
(type locality, characters, neotype), distribution, natural history.

Resumen

La descripci6n holotipica e hist6rica de Alexander von Humboldt para el primate Simia j ... contiene obvious errors
que han creado various problems taxon6micos, dado que ha sido impossible comparar descripciones de subespecies con un
holotipo original, el cual nunca se preserve. Un error hist6rico es el reconocimiento de Cebus .j ... unicolor como un
taxon distinto de Cebus ,. ... j ', .j ... el cual corregimos en este articulo por el reconocimiento de Cebus j ... unicolor
como sin6nimo para Cebus j ... albifrons. Describimos Cebus ... ', j ... adecuadamente en este articulo por la
primer vez, basado en un neotipo colectado por nosotros cerca a la localidad tipica. Adicionalmente, se discute y se clarifica
confusiones sobre la localidad tipica, indicando Maypures como la localidad tipica correct. Adicionalmente, se present
informaci6n general sobre la distribuci6n geografica y la historic natural de la especie y la subespecie nominal.

Palabras Claves Primates, Cebidae, capucino de frente blanca, Cebus j. ... ',. ] .. Simia j. ] ... Humboldt, 1812
(localidad tipica, caracteres, neotipo), distribuci6n, historic natural.



Introduction


During the years 1799-1804, on an extensive biological
and geographical reconnaissance of Colombia, Venezuela,
Ecuador, Peru, Cuba and Mexico, including the Orinoco
and Magdalena rivers and the Colombian and Ecuadorian
Andes, Baron Alexander von Humboldt and his
companion Aime Bonpland explored isolated and little-
known regions of South America, studying geographical,
climatological and geophysical aspects of the countryside,
local customs, archaeology and the innumerable plants and
animals native to those parts. As fruit of their explorations,
many organisms were described for the first time in the


thirty volumes which were published, complemented with
numerous articles in scientific journals. This was surely the
most extensive project of its kind ever published by one
individual, given that Humboldt's friend Aime Bonpland,
except for the botanical part, contributed very little to
the actual writing of the results of the voyage of these
two scientists. Even with the botanical work, Bonpland
prevaricated until Humboldt was forced to search for
other collaborators such as Karl Sigesmund Kunth in the
preparation of the seven volume Nova genera et species
plantarum (Botting, 1973: 205) [1].


1 Correspondence to: Thomas R. Defler, Instituto Amaz6nico de Investigaciones, Universidad Nacional de Colombia, A.A. 215, Leticia
(Amazonas), Colombia. E. mail: .
t It is with great sadness that I inform the primatology and conservation community of the untimely death of Professor Jorge Ignacio
Hernindez-Camacho on September 15, 2001. Professor Hernindez or "El Mono" was my friend, colleague and teacher for over 25 years.
My knowledge of Colombian primatology and other vertebrate fauna was greatly enriched by having known and worked with him. His
human qualities were considerable, and he will be missed by many. (TRD)





Neotropical Primates 10(2), August 2002


While in the Orinoco region, Humboldt (1812a: 305-
363) discovered and described five new taxa of primates:
Aotus trivirgatus [= Simia trivirgata Humboldt], ....
torquatus lugens [= Simia lugens Humboldt], Lagothrix
lagothricha [= Simia lagothricha Humboldt], Cacajao
melanocephalus [= Simia melanocephala Humboldt],
Chiropotes satanas chiropotes [= Simia chiropotes Humboldt]
and Cebus ,. ... [= Simia ,. ... Humboldt] [2].

This article discusses the characteristics of Simia ... ,
the species' taxonomy, its authentic type locality, the
fixation of a neotype for the taxon, the status of Cebus
gracilis Spix, 1823, the synonymy of Cebus j ... unicolor
with Cebus ,. ... albifrons, the geographic distribution
of C. albifrons, and aspects of the species' natural history.

Methods

Color terminology for pelage descriptions follows
Ridgway ("1912" = 1913). Measurements are expressed in
millimeters, unless otherwise noted. Head and body length
were obtained by subtracting the tail length from total
length. For this article we examined all specimens of Cebus
,.f ... ',.j ... and Cebus j. ... unicolor deposited in
Colombian collections (11 specimens), comparing them to
our knowledge of Venezuelan and Brazilian specimens of
( ... -j ... unicolor. Additionally we reviewed specimens
of other Cebus r j ... subspecies deposited in Colombia
(21 specimens), and we cursorily examined specimens
in the United States National Museum (68 specimens,
including holotypes for Cebus -j... cesarae and Cebus
,.f... pleei). There are no specimens of Colombian
Cebus .j... i=. '. f... <= C. a. unicolor) held in collections
outside of Colombia except for a series collected from the
right bank of the rfo Arauca and deposited in the Field
Museum of Natural History (FMNH), Chicago.

The following acronyms are used:

AMNH The American Museum of Natural History, New
York, USA.
FMNH Field Museum of Natural History, Chicago,
USA.
ICN Collection of mammals in the Instituto de Ciencias
Naturales, Museo de Historia Natural, Facultad de
Ciencias, Universidad Nacional de Colombia, Santa Fe de
Bogota, Colombia.
IVH Instituto de Recursos Biol6gicos Alexander von
Humboldt, Ministerio del Medio Ambiente, Villa de Lefva,
Boyaca, Colombia.
MNHNP Museum National d'Histoire Naturelle, Paris,
France.
UNIFEM mammal collection, Unidad de Investigaciones
Federic Medem, Institututo Nacional de Recursos Naturales
y Protecci6n al Medio Ambiente (INDERENA), Santa Fe
de Bogota, Colombia. This collection is now deposited in
IVH.
USNM Natural History Museum, Smithsonian
Institution, Washington, DC, USA.


Simia albifrons Humboldt, 1812

The original description of Simia albifrons. As von
Humboldt and Bonpland entered the world of the "Upper
Orinoco" [3] they began to see individuals of species of
"Matchfs" [4]. They were already acquainted with both
Cebus a,/1//,i, the "Sajou" [5], and the "saY" [6] from earlier
collections. Humboldt named this new monkey Simia
albifrons. "Face bluish gray with the exception of orbits and
forehead, which are pure white. The contrast of these two
colors distinguishes the "Ouavapavi", which I name Simia
albifrons, from the "Sal" and from the ordinary "Sajou"
(translation from the French) (Humboldt, 1812a: 324).

Humboldt (1812a: 324-325) described Simia .. as
follows:
"Le Matchi du Haut-Orenoque, que les Indiens
Guarekens [7] appellent Ouavapavi a Om,378 (14
pouces) de long du sommet de la tete a l'origine
de la queue: il a la face gris-bleuatre, a l'exception
des orbites et du front qui sont d'un blanc pur. Le
contrast de ces deux couleurs fait distinguer au
premier abord l'Ouavapavi, que je d&signe sous le
nom de Simia albifrons, du Sal et du Sajou ordinaire.
La tete est un ovale tres-alongd. Le pelage du corps
est grisatre, plus clair vers la poitrine et le venture, plus
obscur vers les extremites que son d'un brun-jaunatre.
Le sommet de la tte est d'un gris tirant sur le noir:
une strie cendrde se prolonge longitudinalemente de
la calotte par le milieu de front vers le nez: les sourcils
sont de meme d'un gris tres-obscur. Les yeux son
grands, bruns et tres-vifs. Les oreilles ont un rebord
et sont couvertes de poils. La queue est prenante,
mais toute couverte de poils, et par consequent sans
callosit&: elle est a peu pros de la longueur de corps,
cendrde par dessus, blanchatre par dessous, et d'un
brun-noir 1l'extremitd. Les ongles son tous arrondis
et tres-peu convexes. Une strie d'un gris foncd obscure
descend le long de dos.
SIMIA ALBIFRONS, imberbis, cauda prehensili, ex
.... cinerascens, vertice nigro, face coerulea, fronte et
orbitis niveis, cruribus et brachiisfuscescentibus."

Further on in the above publication, Humboldt lists Simia
f ... in his Tableau Synoptique des Singes de l'Amerique
as follows:
"19. Simia ... ex albo cinerascens, vertice nigro,
facie caerulea, fronte et orbitis niveis, cruribus et
brachiis fuscescentibus." (Humboldt, 1812b: 356).

Type specimen and type locality. Humboldt did not
specifically designate a type specimen nor a type locality,
although he added the following comment:
"Les Ouavapavissont tres-laids, mais extrememement
doux, agiles et moins criards que les Singes pleureurs
[Cebus nigrivitattus]. Ils habitent, par troupeaux, les
forces qui avoisinent les cataractes de l'Orenoque
[8] et la mission de Santa Barbara [9]. Nous en
avons trouv6 un individu a Maypures [10] qui, tous





Neotropical Primates 10(2), August 2002


les matins, saisissoit un cuchon sur lequel il resoit
mont6 toute la journde en parcourant la savane
qui environne les cabanes de Indiens [Maypures].
Nous l'avons meme vu souvent sur le dos d'un chat
qui avoit &td 6lev6 avec le Singe dans la maison du
missionnaire, et qui souffroit patiemment les effects
de la p&tulance de l'Ouavapavi."(Humboldt, 1812a:
324-325).

The original description was based exclusively on a captive
individual that Humboldt (1812a: 325) first reported he
saw at Maipures, on the left bank of the rfo Orinoco, but
added that he had observed wild individuals around the
Mission of La Esmeralda (Estado Amazonas, Venezuela),
on the upper rfo Orinoco (right bank) and in the forests
on both sides of the Orinoco between La Esmeralda and
Maipures. Later, Humboldt (1824a: 98-99, [1]) wrote in
his Relation Historique of the journey to the rio Orinoco
where he had supposedly observed Simia -j ... in the
Atures Mission [11], 50 km north of Maipures on the
opposite side of the river, directly contradicting his original
statement in Humboldt (1812a: 325).

This new account (Humboldt, 1824a: 98-99) [12] is
entirely based on the original one, including the Latin
diagnosis, but no mention of Maipures is made nor a
clarifying explanation. Under such circumstances, von
Humboldt certainly created a source of confusion twelve
years after the original description was published, since
there is no evidence of the presence of Cebus ..-
in the vicinity of Atures on the right bank (Venezuelan


bank) of the Orinoco, where Cebus nigrivittatus occurs
instead (see Bodini and Pdrez-Hernandez, 1987; Bodini,
1989) [13].

One of us (TRD) studied the distribution of Cebus ,....
and Cebus ,i'ic/ in the Maipures region and was not able
to confirm the presence of Cebus ,.j ... in the inmediate
location of Maipures, south of the rfo Tuparro (Defler,
1985). The site that Maipures formerly occupied is in an
extensive natural savanna with isolated forest patches that
contain some Cebus ,.'ic/l as well as Alouatta seniculus and
Aotus sp. (perhaps A. trivirgatus) [14]. However, C. ,....
is common (and replaces C. 7/''1,i immediately 3 km north
(on the northern bank of the lower rfio Tuparro) where there
are nearby extensive gallery forests (Fig. 1).

As such, this argues against the possibility that the specimen
upon which Humboldt based the original description of
Simia .j ... was obtained at the Atures mission or any
other locality in Venezuela and suggests that it was captured
in the neighborhood of Maipures mission, Vichada,
Colombia, where Cebus ,. ... (as currently understood)
is known to occur close by.

Possible doubts for the occurrence of Cebus .j ... on the
Colombian bank of the rfo Orinoco that could eventually
arise were definitively resolved with the findings of the above
survey and of the additional collection of the species in the
rfo Bita (Vita), further to the north of the survey area.


Ammended type locality
for Cebus albifrons
Rio Tomo


Rio Tuparro Rio

* = Cebus apella
o = Cebus albifrons


Figure 1. Map of Maipures region north to the rio Tomo showing the distribution of Cebus .. -I... Cebus apela, and including the
amended type locality for Cebus , .. (based on Defler, 1985).





Neotropical Primates 10(2), August 2002


Fixation of a neotype for Simia albifrons. The fixation of
a neotype is justified because: It is desirable to establish the
neotype to secure nomenclatural stability; the holotype was
not preserved; and it is convenient to clarify the type locality
of the species and suppress any ambiguity that could lead to
confusion or controversial interpretations.

The identity and the proper characterization of Simia
albifrons Humboldt. There can be no doubt the original
description of Simia j... based upon the specimen
kept in the mission at Maipures, since it was obviously
handled in order to take accurate measurements of head and
body length (taken from the crown). However, Humboldt
did not publish a figure of it, neither did he record its sex
or relative age, and unfortunately it was not preserved. The
specimen was possibly a juvenile, since it rode about on the
back of a pig, and piggy-back riding is typical of the young
(when pigs or other species are available).

Simia ,.j ... must unquestionably be considered as
belonging to the genus Cebus Erxleben, 1777, following
E. Geoffroy Saint-Hilaire (1812) and the current usage of
subsequent authors. The only species of non-tufted capuchin
approaching the original description is undoubtedly Cebus
albifrons, as interpreted by Hershkovitz (1949: 371).
There are discrepancies in the original description of the
coloration, as shown above, but there is no doubt that
the chromatic pattern of the species is the same in the
Department of Vichada population in Colombia as well
as the Amazonas state population in Venezuela. There
are some differences in the presence of eumelanin in the
pelage, which can be interpreted as individual variation.
The dark color of the hair of the tip of the tail described
by von Humboldt (1812a: 325) is a color pattern that the
authors have not detected in other individuals. This allows
us to hypothesize that the dark tail tip was an individual
variant and the general color of the "grayish" pelage may
suggest that the holotype was a young animal in which the
color was lighter than in adults, or that the captive animal
had been exposed to so much sunlight that depigmentation
(fading) had occurred and the yellowish-red tones had been
lost. We have noticed that captive specimens, when exposed
to too much sun tend to lose their natural color, such that
tonalities become duller towards gray.

However, as previously noted, no type or syntype of this
species was preserved. It has also been established here
that the type locality is fairly clearly the former site of the
Maipures mission, although only Cebus ,'iI//. exists in the
immediate vicinity of Maipures (south of the rfo Tuparro),
as established by Defler (1985). The species of Cebus can be
identified as Cebus .j -.. due to the white color of the
orbits and the forehead indicated by von Humboldt (1812a:
324-325) in the original description of Simia albifrons.

Hershkovitz (1949: 371) rightly indicated that the original
description (and Latin diagnosis) of Simia j ... "'refers
to a monkey with some characters that have never been
found [observed] until now in any of the specimens [in


other populations of the species]". Having now available
topotypes of Simia albifrons, in addition to a considerable
number of preserved and live specimens examined from
most of the geographic range of the species, we fully agree
with Hershkovitz' remarks.

The distinctive chromatic characters used by Humboldt to
define Simia .j ... are:
-Face bluish gray facieie caerulea') except for the orbital regions
and the forehead which are white ("niveis"). Comments: The
usual condition in this species for all specimens examined
is a depigmented facial skin, including the forehead, instead
of a pigmented facial skin, with unpigmented orbits and
forehead. However, a certain grade of facial pigmentation as
blotches ofeumelanin over a light pinkish skin color has been
observed in the populations of the rio Mataven (s. Vichada)
and in the rio Apaporis (s. Vaupes) region (T. R. Defler, pers.
obs.). It is conceivable that a general pale bluish gray color
could appear in the facial skin (excluding the orbits and the
forehead), due to small amounts of scattered eumelanin as an
individual variation.
-Eyebrows very dark gray ("gris tr&s-obscur'). Comment:
Von Humboldt undoubtedly referred to the presence of
very dark hairs (" .. ) in the eyebrows, as in Cebus
albifrons, and not to a dark superciliary stripe.
-Crown gray tending to black ("vertice nigro"), anteriorly
continued as a grayish narrow stripe ("strie cendrie") that
descends towards the nose. Comments: The basic contrasting
dark design of the crown occurs in all the various subspecies
of Cebus albifrons, but its color can change, varying from
a medium brown to a dark brown (almost blackish), but
never with gray tonalities. Possibly the dark gray almost
black color might be due to some degree of bleaching due
to excessive exposure to sunlight.
-Underparts grayish ("grisdtre"; ..... ... in the
Latin diagnosis), darker in the extremities ("cruribus et
brachiis") which are yellowish brown (" fuscescentibus") with
a darker middorsal stripe ("gris foncd ., ). Comments:
The "grisatre" coloration can be translated as "grayish"
or "grayish brown" that would be closer to the tonalities
present in the topotypes of Simia .j... (in any case,
not tending to ashy gray), and the middorsal stripe is not
darker gray. The color of the extremities, including hands
and feet, can be interpreted as yellowish brown, not duller
("fuscescentibus").
-Tail above ashy gray ("cendrie"), whitish underneath
(" blanchdtre par .. .. ) blackish brown (" br ... )
towards the tip. Comments: In the topotypes of Simia
f... the tail is more richly colored and not strikingly
darker (brownish black) towards the tip. In none of the
specimens of Cebus j ... does the tail tip approach a
very dark brown color; on the contrary, a tendency towards
a lighter tail tip is frequent.
-Breast and bll), whitish. Comment: The breast and belly
are brighter colored in the topotypes. Some individuals,
however, have a very white ventrum.

To summarize: the coloration described for the middorsal
area, the proximal dorsal surface of the tail and the sides of





Neotropical Primates 10(2), August 2002


the body in the type specimen of Simia ,.j ... is rather
duller and decidedly more grayish than in the topotypes,
and with a unique feature; the brownish black tip of the
tail. These discrepancies cannot be entirely attributed to
a process of bleaching or fading due to excessive exposure
to sunlight. It is well-known that black (eumelanin) hairs
bleach or fade to reddish brown instead of deep gray due to
excessive exposure to sunlight, unlike the case here. Under
these circumstances we can conclude that either the type of
Simia j ... was abnormally colored, or the published
chromatic description was affected by inaccuracies in the
terminology used, or the description was at least in part
subject to defective perception of the tonalities involved.
Otherwise we should expect a close similarity or even
identity in chromatic characters between the type and the
topotypes.

A strict interpretation of the original description of Simia
,.f... leads us to the conclusion that Cebus .f...
cannot be satisfactorily identifiable and should be replaced
by Cebus unicolor Von Spix, 1823. The obvious alternative
is to preserve the use of Simia .. based on the
characters of the topotypes now available and the acceptance
of Maipures as the type locality for this binomen. As this
procedure essentially is in accordance with the taxonomic
and nomenclatural treatment of Hershkovitz (1949) that
has generally been followed since then, and in order to
consolidate nomenclatural stability, in this article we fix
one of the specimens collected by us a few kilometers north
of Maipures as a neotype for Simia albifrons.

There are no arguments to suggest that the populations
subsequently observed by us in the wild nearby are not
of the same population as the holotype. Since the species
is polytypic with a wide geographic range, the taxonomic
identity needs to be consolidated according to Article 75
of the International Code of Zoological Nomenclature. It
is important to designate a neotype for the species with the
object of establishing a nomenclature and taxonomic base
for the species and genus.

Cebus albifrons albifrons (Humboldt, 1812)

Synonymic history

Simia *.f... Humboldt, 1812a: (original description and
Latin diagnosis).
- Humboldt, 1812b: 563 (Latin diagnosis; "Habite
les environs de Maypures et d'Atures, sur les bords de
I'Orenoque").
- Humboldt, 1824a: 98-99, footnote 1 (citation in the text
of the footnote; Latin diagnosis; characters; recorded among
the primates seen la mission de Atures", not Maypures!).

Cebus j... E. Geoffroy Saint-Hilaire, 1812: 111
(nomenclatorial transference to Cebus).
- Goeldi & Hagmann, 1904: 48 (including Cebus chrysopus
and C. gracilis in synonymy).


- Elliot, 1913: 88 (partim; synonymy including C. gracilis
and C. leucocephalus as synonyms; characters).
- Cruz Lima, 1945: 149 (characters based on the original
description; erroneous citation of type locality as "Santa
Barbara Mission, cataracts of rio Orinoco").
- Napier, P 1976: (partim; catalogue of specimens in the
British Museum [Natural History]).
- Handley, 1976: 42 (partim; Rio Mavaca, 108 km SSE
of Esmeralda, 140 km and Tamacama, Rio Orinoco, 135
m., T[erritorio] F[ederal] Amazonas [= Estado Amazonas],
Venezuela).
- Groves & Pulido, 1982: 228 (erroneous citation of type
locality as "Venezuela, Orinoco River").
- Cuervo-Dfaz, Hernandez-Camacho & Cadena [G6mez],
1986: (partim; actual distribution and synonymy with
C. a. unicolor).
- Bodini & P&rez-Hernandez, 1987: (partim; distribution
in Venezuela).
- Bodini, 1989: 105-106 (distribution of species in
Venezuela).
- Groves, 1993: 259 (erroneous citation of type locality as
"Venezuela, Orinoco River").
- Uribe Hurtado & Ortiz Von Halle, 1993: fig. s.n. (Cano
Lim6n, Department of Arauca, Colombia).
- Alberico, Cadena, Hernandez-Camacho & Munoz-
Saba, 2000: 58 (Department of Putumayo and Vichada,
Colombia).

C[ebus] c[apucinus] gracilis Pusch, 1941: 192 (partim:
including records from the Amazonian region as well as
"Cebus gracilis Hellsternig" described by Lbnnb erg [1939]).

C[ebus] c[apucinus] versicolor Pusch, 1941: 193 (partim:
including the records from the Amazonian region and
" Cebus flavus Geoffroy", non Cebus versicolor Pucheran,
1945 [= Cebus j.. versicolor]).

C[ebus] cuscinus cuscinus Pusch, 1941: 196 (partim: "Cebus
gracilis dunkelstirning" described by Ldnnberg [1939] and
a female from Chicosa, eastern Perd; non Cebus flavescens
cuscinus Thomas, 1901 [= Cebus j.. cuscinus]).

Cebus fiavus E. Geoffroy Saint-Hilaire, 1812: 111 (original
description; holotype MNHNP, no. 562 (type specimen
catalogue) and 458 (general collection); unsexed adult (?)
collected by Alexandre Rodrigues Ferreira in Brazil) [15].

Cebus unicolor Spix, 1823: 7, pl. 4 (original description;
holotype: Zoologische Staatssamlung Mtinchen, adult male,
skin and skull, collect by the expedition of Jean Baptist
Ritter von Spix and Carl Friedrich von Martius in Brazil).
Type locality: forests of Rio Tefe, near its junction with the
Amazon River near Ega [= Tefk], Amazonas, Brazil).

Cebus gracilis Spix, 1823:8, pl. 5 (original description;
holotype: Zoologische Staatssamlung, Miinchen, skin and
skull collected by the expedition ofJohann Baptist Ritter von
Spix and Carl Friedrich von Martius in Brazil; type locality:
Tefe, mouth of Tefe River on the Amazon River, Amazonas,





Neotropical Primates 10(2), August 2002


Brazil; distribution forests of the Solim6es from "la ville de
rio Negro [=Manaus, Amazonas, Brazil] vers le Peru").
- Cruz Lima, 1945: 149-150 (pl. xxiv (characters based on
the original description).

"Sajou a pieds doree au chrysope" F Cuvier, 1825: 2pp.,
pl. (description of a living menagerie specimen from
"I'Amerique septentrionale").

Cebus chrysopus Lesson, 1827: 55 (based on the original
description of the "sajou a pieds dories de F Cuvier in
1825; type not preserved).

"Machin (nuova vapari; [sic. = ouavapavi?]: Codazzi, 1841:
156 ("tiene pelo gris y cara azulada con las 6rbitas y la frente
como la nieve; Venezuela, without definite locality).

S[imia] .-f... Vergara & Velasco, 1902: 190 (cited in
the text; Colombia without precise locality; common name
"machfn").

C[ebus] ... Pittier & Tate, 1932: 278 ("Raudales del
Orinoco", Venezuela).

Cebus ..f... [albifrons] Defler, 1979a: 475, 487, 488
(ecological aspects of topotypical population).
- Defler, 1979b, 1979: 491, 501 (behavioral aspects of
topotypic population).

Cebus unicolor unicolor: Cruz Lima, 1945:150 (characters
based on the original description).

Cebus gracilis Spix (vel C. j ..- Humboldt ?): Lbnnberg,
1939: 17 et seq. q. (Codajaz, rio Solim6es, Amazonas,
Brazil; Irocanga, rio Tapaj6s, Para, Brazil; Jaburd, rio Purds,
Amazonas; Igarapd do Gordao, rio Jurua, Amazonas; Joao
Pessoa, rio Jurua, Amazonas; Lago Grande, rio Jurui,
Amazonas, Brazil; San Ant6nio, rio Eird, Amazonas,
Brazil).

Cebus 7 .. .. ... Hershkovitz, 1949: 370-372, fig.
54 taxonomicc revision; characters transcribed from the
original description).
- Cabrera, 1958: 160 (type locality: "selvas pr6ximas a los
raudales del Orinoco"; distribution "alto Orinoco").
- Hill, 1960: 450-451 (characters translated from the
original description; type locality and distribution after
Humboldt [1812b]).
- Pusch, 1941 (partim: original description only?, as
synonyms included belonging to the Cebus .,'i.// group
and geographical distribution is given as "Rio de Janeiro
and Sao Paulo", Brazil.
- Rylands, Mittermeier & Rodriguez-Luna, 1995: 120,
128, 137, 13 (Colombia; IUCN classification LR = Lower
Risk).

C[ebus] a[lbifrons] / ... Hernandez-Camacho &
Cooper, 1976: 58, fig. 10 (characters taken from Humboldt
compared with topotypical population; "eastern Vichada").


- Hernandez-Camacho & Defler, 1989: 91-92 (basic
characteristics; conservation status).
- Rylands, Schneider, Langguth, Mittermeier, Groves &
Rodrfguez-Luna, 2000:68 (tab. 5) 76, 78-79 (C. a. unicolor
included as a junior synonym of C. a. -] .* )

C[ebus] .ij/l.fjo'] unicolor: Hernandez-Camacho &
Cooper, 1976: 58, fig. 10 (possible junior synonym of
C. a. albifrons; range in Colombia: Vaupes and south of the
rio Caqueta (except for the interfluvium between the rio
Guamues and rio San Miguel or Sucumbfos, Department
of Putumayo, inhabited by C. a. yuracus, Hershkovitz,
1949).

C[ebus] a[lbifrons] [subsp.]: Hernandez-Camacho &
Cooper, 1976: 58, fig. 10 ("pale and dull colored population"
in western Arauca, northern Boyaca and southwestern
and southeastern Norte de Santander; somewhat similar to
C. a. adustus Hershkovitz, 1949).

Cebus .f... unicolor: Hershkovitz, 1949: 372-375,
fig. 54 (revision; characters; Marimonda, rio Orinoco,
Amazonas, Venezuela; Solano, rio Cassiquiare, Amazonas,
Venezuela; Yavanari, rio Negro, Amazonas, Brazil; Casas
Pereira Igarapd, rio Negro, Amazonas, Brazil; Puerto
Victoria, rio Pachitea, Huanuco, Perd; Tingo Maria,
Huanuco, Perd; no locality, Perd).
- Cabrera, 1958: 161-162 (distribution, including
southeastern Colombia).
- Hill, 1960: 451-453 (characters essentially based on
Hershkovitz [1949]; distribution).
- Rylands, Mittermeier & Rodrfguez-Luna, 1995:120, 128,
135, 137, Perd and Venezuela; IUCN classification LR =
Lower Risk).

Neotype. Young adult male in fresh pelage, skin and
skull (Table 1), UNIFEM (Unidad Investigativa Federico
Medem INDERENA) now deposited in the collection
of the Instituto de Investigaci6n de Recursos Biol6gicos
Alexander von Humboldt (IVH), Villa de Leyva, Boyaca,
Colombia, No. 2844, collected by T. R. Defler on 30
January, 1978 (Figs. 2 & 3).

Amended type locality. About 10 km north of Maypures,
200 m north of the Cerro Rocoso, El Tuparro National
Park, Department of Vichada, Colombia (5020'N,
67045'W) (Fig. 1).

Topotypes. Young adult male, skin and skull, UNIFEM,
No. 2843 by T. R. Defler on 30 January, 1978. Adult
female, skin and skull, UNIFEM, No. 2839 byT. R. Defler
on 30 January, 1978, all specimens collected from the same
group as the neotype and now deposited in the collection
of the Instituto de Investigaci6n de Recursos Biol6gicos
Alexander von Humboldt (IVH), Villa de Leyva, Boyaca,
Colombia.

Coloration of neotype. Alae nasi with dark brown
pigmentation with sparse light Cartridge Buff hairs over






Neotropical Primates 10(2), August 2002


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Neotropical Primates 10(2), August 2002


frontal region; dark blackish narrow line extends from nose
up, crossing supra-orbital band and ending in superciliary
hairs Hair Brown above center of forehead; supra-orbital
blackish band above and lateral to the orbits with Hair
Brown superciliary hairs; crown Sepia with bases of hairs
more pallid than Cartridge Brown; lightest parts of ears close
to Cartridge Buffbut slightly lighter and yellower; back at
mid-dorsal line rr :; ... - Pale Pinkish Buff; sides lighter
with less brown, lateral fringe slightly lighter than Cinnamon
Buff; no whitish patch in front of shoulders; scapular
region and shoulder Pinkish Buffx Pale Pinkish Buff; chest
and belly Ochraceous Buff x Ochraceous Orange; forearm
and thighs Ochraceous Buffx Zinc Orange contrasting with
dark lining of back; wrists and ankles Mikado Brown but
fingers with much less hair and contrasting with blackish
skin; feet dorsally same color as thighs; diminished hair
on digits shows blackish skin. Palmar and plantar surfaces
including digits dull pink; tail bicolored with broad dorsal
stripe Saccardo Umber with grizzled effect due to Ochraceous
Buffhair tips throughout to tail tip; distal dorsal part of
tail slightly lighter Xanthine Orange x Zinc Orange, with tail
ventrum lighter than Cartridge Buff.

Coloration of topotypes. UNIFEM No. 2843; similar
to the neotype but crown Warm Sepia with bases of hairs
more pallid than Cartridge Buff, back at middorsal line


Buckthorn Brown, anterior parts darkening to Russet over
hips; sides lighter with less brown; forearm and thighs
Ocraceous Buff x Ochraceous Orange; tail bicolored with
broad dorsal stripe Saccardo Umber proximal to the body,
lightening considerably over middle and distal parts to
Cinnamon Buff. UNIFEM No. 2839; essentially similar
to the neotype except for the color of the chest and belly
which appear slightly brighter Ochraceous Buff x Zinc
Orange.

Comparison with Cebus albifrons unicolor. Spix (1823:
7) described Cebus unicolor (= Cebusgracilis Spix, 1823: 8)
from near the mouth of Tef6 (formerly Ega), Amazonas,
Brazil. His animal has been identified throughout a
wide area of the middle and upper Amazon, including
Colombia, Peru and Bolivia, as well as part of the upper
Orinoco in southern Venezuela. Elliot (1913) included
Cebus unicolor Spix as a synonym of Cebus ,.j ..
and treated C. gracilis as a distinct species. Cebus gracilis
Spix was based on an adult female collected in the same
locality as the type male of C. unicolor Spix, and the
alleged differences between those nominal species fall in
the individual and sexual dichromatism known in other
populations of C. ,j ..- As a result, Hershkovitz (1949:
372-373) regarded unicolor and gracilis as synonyms.
The characters, as published by Hershkovitz (1949:


Figure 2. Color plate of neotype of Cebus j .. ', .. Humboldt, 1812. By Margarita Nieto.





Neotropical Primates 10(2), August 2002


372-375), based on the original description of material
and upon color plates of unicolor, are as follows: "Most
uniformly brightly colored race of albifrons, Cap Snuff
Brown to Bister, frontal region buffy to ochraceous; back
Ochraceous-Buff to Ochraceous-Orange or Tawny more
or less lined with dark brown; sides with less brown, lateral
fringe Ochraceous-Buff to Ochraceous-Orange; forearm
and foreleg Ochraceous-Buff to Tawny contrasting with
dark lining of black; hairs of belly Ochraceous-Buff to
Ochraceous-Orange, of chest like belly or white; whitish
patch obsolete on front of shoulder."

Cebus albifrons unicolor is a synonym of Cebus albifrons
albifrons. Hershkovitz' (1949: 372-374) description
of Cebus .f... unicolor seems very similar to the
population north of Maipures. One of us (JHC) examined
specimens from southern Venezuela (Amazonas State) in
the AMNH and the USNM also ascribed to C. a. unicolor,
and found them to be virtually identical to C. a. ....
as represented by its neotype. Specimens examined by us
from the area between the ri6s Amazonas and Vichada
(Amazonia, Colombia) usually ascribed to C. a. unicolor
also seem indistinguishable from C. a. ... described
here. This leads us to the conclusion that unicolor is a
synonym of albifrons.


Other specimens examined. UNIFEM 3022 adult male,
Miritf-Parana, Amazonas, Colombia; UNIFEM 3023 adult
male, Miritf-Parana, Amazonas, Colombia; UNIFEM
3029 juvenile male, Miritf-Parana, Amazonas, Colombia;
UNIFEM 2843 adult female; UNIFEM (uncatalogued)
adult female, Canro Brava, rfo Cotuh6, Amazonas,
Colombia; UNIFEM 0206, rfo Arauca, 65 km upriver
from the town of Arauca, Colombia; UNIFEM 1523,
San Jose de Ocun6 ("30 miles" [45 km] to the south),
Vichada, Colombia; UNIFEM 2667, rfo Peneya, Caqueti,
Colombia.

Distribution of Cebus albifrons albifrons as previously
recognized. The geographic range of this apparently isolated
population of C. a. *.f... i mapped in Defler (1985) and
is reproduced in Figure 1. The range includes the lower rfos
Tuparro (left bank), Tuparrito, Tomo, Bita and Meta (right
bank) in Vichada. On the upper parts of these rivers C.
. ', ..f ... is replaced by C. c//a, i (Hernandez Camacho and
Cooper, 1976; Defler, 1985). Another apparently isolated
population is found in Arauca, Colombia; though the limits
of this population are not clearly defined and possibly reach
into western Venezuela (Apure State). Because of the lack of
specimens in many areas, it is not possible to define securely
the limits of C. a. ,.j ... outside of Colombia.


Figure 3. Photograph of skull of neotype: a. Frontal view; b. Lateral view; c. Dorsal view; d. Inferior mandible.





Neotropical Primates 10(2), August 2002


Expanded geographic range of Cebus albifrons albifrons.
Accepting C. a. unicolor as a synonym for C. a. .j ...
extends the range to a huge area in the upper Amazon of
Brazil, Colombia, Peru and Bolivia from the right bank
of the Marafion River in the south, but also crossing the
Amazon and including much of southeast Colombia and
southern Venezuela (Fig. 4). A gazetteer of Colombian
specimens and observations by TRD of Cebus .. and
C. a. .-.. is given in Defler (in press). Limits for the
species are still imperfectly known.

Variation. Some specimens have an admixture more
yellowish than reddish on the arms and legs. One male
specimen collected at Puerto Rastrojo in the Miritif-Parana
of the Colombian Amazon weighed 4135 g (INDERENA
No. 3033) and another male from the same locality
(INDERENA No. 3022) weighed 3490 g. These specimens
have a darker, buffy-brown forehead as compared to the
neotype and topotypes, and the dorsal brown is darker.
In general these animals also have a more reddish cast
than the neotype, while the neotype and topotypes are
lighter and more buffy. A juvenile male specimen from


,. i



'
i

7


the same site (INDERENA No. 3029) is much lighter
than INDERENA No. 3033 and 3022, approximating the
neotype in most respects but without the grizzled effect on
the tail. A young male specimen (INDERENA No. 0206)
from Arauca Department (Colombia) was the lightest
specimen examined. Fur on the forehead and shoulders
is almost white, while the ventrum of the belly and tail
is very light buff. Another young specimen (INDERENA
No 1794) of about the same age as INDERENA No.
0206 is similar in coloration, although the forehead is
slightly darker buff and the dorsal coloring over the hips is
a darker brown. The most anomalous specimen examined
is INDERENA No. 2667 from the rfo Peneya, Caqueta
department (Colombia). This poorly preserved skin is the
darkest brown of all specimens examined although, like
the other specimens, the darker crown does contrast with
the brown back, and there are tonalities of chestnut red on
the arms, legs and hips. The forehead of this specimen is a
buffy-gray.

Natural history and ecology of Cebus albifrons. The
natural history and ecology of this species has been studied


,,^ >W
K K


400 W00 akm
-*irr*,


Figure 4. The geographic distribution of C ,,, .i/' The revised range of C. a. , .. indicated in dark grey. Sources: Aquino and
Encarnaci6n (1994), Bodini and PNrez-Hernindez (1987), Encarnaci6n et al. (1990), Hernindez-Camacho and Cooper (1976), Hersh-
kovitz (1949), Linares (1998).





Neotropical Primates 10(2), August 2002


in Colombia by Defler (1979a, 1979b, 1980, 1985) and
in two sites in Peru by Soini (1983) and Terborgh (1983).
Currently it is being studied in Trinidad by K. A. Phillips
(1998; in prog.). This review of the species' natural history
and ecology has been summarized from Defler (in press).

Habitat preferences. Defler (1985) showed that in the
region of the type locality the species prefers a slightly more
xeromorphic habitat than does C. ''.c//. Such habitats
include Bactris palm forests in seasonally dry riverbeds, and
rocky forests around the bases of inselbergs. C. *.f...
commonly crosses open tracts of rock and savanna from
forest patch to forest patch. In most areas studied it is also
commonly found in flooded forest, which C. a.c//.' tends to
avoid. Flooded forests, then, tend to be part of the habitat
used by C. .j ... where C. 7i.c//.r is sympatric on terra
firme. On the upper rfo Cahuinarf, Amazonas, Colombia,
there is a population of C .* j ... in forests growing on
white sand, where C. a,.c//.r is apparently absent. However,
both species are sympatric in very low numbers in white-
sand caatinga forest on the upper Guacayasi Creek in
Guainfa, Colombia. C. j .. is widely syntopic with
C. .ic//.i.

Group size. In eastern Vichada, Colombia, near the type
locality, C. a. ,. ... is found in very large groups of
around 35 animals. A study group at this site contained
10 adult females, four adult males, three subadult males,
five juvenile females, four juvenile males, one unclassified
juvenile, four infant females, three infant males and one
unclassified infant (Defler, 1979a). To the south, in the
closed rainforest of Peru, C. ...* groups are smaller,
with an average of about eight members at one site, and
15 per group at another (Soini, 1986; Terborgh, 1983),
perhaps because of competition from C. .'ic//.. The groups
are multi-male and multi-female. In Vichada the sex ratio is
2.5 females per male.

C. .j... has been seen to form large transient
congregations when seasonal conditions of food availability
are favorable. Hernandez-Camacho and Cooper (1976: 59)
reported an instance in August, 1956 where a congregation
of several hundred monkeys was observed in a few hectares
along a newly opened road through a secondary forest
association of"guamo" or "guamera" (Ingasp.) with a rather
dense 8-12 m canopy connected to two areas of primary
forest. This was on the road from El Centro to Quebrada
Lfsama (Antioquia). There were also, along a distance
of about 300 m on both sides of the road, considerable
numbers of parrots (Amazona amazonica, A. ochrocephala
and A. farinosa).

Density. At the type locality, the species is present in gallery
and isolated small forests surrounding local inselbergs, at
densities of about 30 individuals/km2 (Defler, 1979a).
Where C. ,*... is sympatric with C. j,.c//. it often
occurs in greatly reduced densities, making its detection
difficult. North of the lower rfo Apaporis in the Colombian
Department of Vaupes around the Estaci6n Biol6gica


Capard (105'33"S, 69030'48"W), for example, the density
is around 1 or less individuals/km2 (Defler, unpublished
data). In the Pacaya-Samiria National Park in Peru,
densities are 4.2-6.2 individuals/km2 (Soini, 1986) and in
the Manu National Park, Peru, densities are much higher,
around 35 individuals/km2 (Terborgh, 1983).

Home range. A group home range of more than 150 ha
was calculated in the Manu study (Terborgh, 1983), while
at least 120 ha were used in the study area in northern
Colombia (Defler, 1979a).

Day range. An average day range of 1820 m was calculated
in the Manu study (Terborgh, 1983).

Activity (time) budget. Terborgh (1983) estimated a
time budget for the species as follows: 18% resting, 21%
traveling, 22% feeding on plant material, and 38% foraging
and feeding on insect material (total feeding time 61%).

Diet. The diet of C. .j... includes fruits, small
invertebrates, their eggs and larvae, small mammals,
lizards, and bird's eggs. They especially like to eat ant and
wasp larvae, and are adept at robbing beehives for honey.
Defler (1979a) observed them hunting frogs (Hyla sp.) in
the interstices of the large fleshy plant Phenakospermum
guianense, which forms a water reservoir between the leaf
bases and the main stem. This water reserve shelters frogs,
and is also used by the monkeys for drinking when other
sources are scarce. The monkeys obtain these resources by
biting out chunks of the plant tissues that cover the water
reservoir.

During the dry season, when there are few fruits, they spend
more than half the day on the forest floor, searching for live
prey. Terborgh (1983) observed C. j ... exploiting
73 species of plants from 33 families, of which by far
the most important was Moraceae, with 17 species
included in the diet. Near the type locality, the palm
species Maximiliana (Attalea) regia is a key species, being
used more than any other plant resource. The palm nuts
provide a nutritious food during the dry season when little
other plant food is available. At the Manu National Park,
Astrocaryum and Scheelea palms are the most important
plants for these monkeys. Ants, ant eggs and small beetles
are searched for incessantly in rolled leaves and around the
leaf bases of palms.

Reproductive behavior. Copulation by the dominant male
lasts several minutes with the male mounting the female,
grasping her hind legs with his hind feet. The gestation time
is unknown, but it is probably similar to Cebus ,'i//ia, at
about 160 days.

Infant development. Usually one infant is born. During
the first 1-2 days it tries various positions for holding onto
the mother, including clasping the tail, hind leg, and arms,
until it discovers a position over the neck and shoulders
that is typical for the species. After some weeks of riding





Neotropical Primates 10(2), August 2002


oriented crossways over the shoulders the baby is able to
ride lengthwise on her back as do most primate babies. All
members of the group are interested in the newborn infant
and wish to be close to it. The genitals are of particular
interest to the other females in the group. As the infant
matures, various group members try to entice the baby
onto them for carrying, and, eventually, the mother allows
others, including adults and young monkeys, males and
females, to help carry the baby. Even the alpha male carries
infants.

Social behavior. Males are very friendly to each other and
seem very unassertive. Nevertheless, the sub-alpha males are
constantly aware of the position and activities of the alpha.
Males are very aggressive towards males of neighboring
groups, and aggressive chasing bouts occur where two
territories overlap. All members of the group, and especially
the alpha male, break branches over predators or over
perceived danger, causing noisy crashes through the forest
vegetation and much excitement. Near the type locality,
boas (Boa constrictor) and tayras (Eira barbara) have been
observed stalking the capuchins, but these predators were
usually detected and then ignored. Cebus .j ... showed
great fear towards a raptor, Spizaetus ornatus, which
attempted to attack them, causing all troop members to
drop to the ground and flee.

White-fronted capuchins are sometimes associated with
squirrel monkeys, Saimiri sciureus, brown capuchins,
Cebus ,jc//i, and woolly monkeys, Lagothrix lagothricha.
They can feed in trees occupied by red howling monkeys,
Alouatta seniculus (Defler, 1979a). The general condition of
sympatry between C. j ... and C. ai/,cll is evidenced
by the observation of both capuchin monkeys in 15
inventories of mammal species carried out in Amazonia
(Voss and Emmons, 1996: 103-114).

Ecological niche. Terborgh (1983) studied the strategy of
this species in closed-canopy forest when in sympatry with
C. j,//,7. Cebus ..j ... ri, I, widely, searching for patchy
resources such as Ficus, which it exploits exhaustively before
moving on. It seems probable that the strategy they use in
the gallery forests near the type locality may be different,
however, from the food-rich study site on the rio Manu in
southern Peru.

Acknowledgements

We are particularly grateful to the personnel of the
collections at the Instituto de Investigaci6n de Recursos
Biol6gicos Alexander von Humboldt (Fernando Gast and
Yaneth Mufioz), Instituto de Ciencias Naturales of the
Colombian National University (Alberto Cadena), and
the US National Museum (Richard Thorington III) for
permission to work with material in these collections.
We are grateful also to the American Museum of
Natural History for permission to work there. We thank
INDERENA, the American Peace Corps, the Universidad
Nacional de Colombia, Conservation International, and


the Margot Marsh Biodiversity Foundation for support for
various facets of this work, and Margarita Nieto for the
color plate in this article.

Notes

[1] Von Humboldt and Bonpland's itinerary has often
been described with the claim that they arrived for the first
time on Colombian territory on 2 April, 1801 at Puerto
del Zapote in the Bahfa of Cispata, C6rdova (formerly
C6rdoba) on the Caribbean coast of Colombia, The two
scientists stayed there for three days before continuing their
journey to Cartagena de Indias. However, the first day that
von Humboldt arrived on Colombian territory was not
the above date, but rather on 13 April, 1800, when they
landed on the Playa de Guaripo or Guaripa (ca. 556'N,
67030'W), above the Raudal de San Borja, Vichada
(Dugand,1954: 210).

[2] Hershkovitz (1987a: 54) published the following
pertinent remarks on von Humboldt's activities during
his trip to tropical America: "Monkeys, however, absorbed
more of Humboldt's attention than other animals. He
carried with him a number of live simians captured in the
upper Rio Orinoco region for shipment to the Jardin des
Plantes in Paris, via the Antillean island of Guadeloupe.
The newly discovered bearded saki (Chiropotes satanas
chiropotes Humboldt [= Simia chiropotes von Humboldt,
1812]...) died before transshipment, but its skin was saved
and arrived in Paris. The type specimen of a red howler
Simia ursina Humboldt (= Alouatta seniculus arctoides
Cabrera) survived the journey, whereas the first-known
douroucouli or night monkey (Aotus trivirgatus Humboldt
[Simia trivirgata von Humboldt, 1812] succumbed in
Guadeloupe". No mention is made by Hershkovitz of
Cebus ,. ... but in any case the type of this species did
not arrive in Paris, nor was it preserved.

[3] The "upper Orinoco" of von Humboldt is in effect the
Orinoco above the rapids of Rabipelado, San Borja and
Atures, which impede navigation and separate the upper
from the lower Orinoco.

[4] "Matchi" is a name derived from machine of Quechua
origin. "Machin" is a common name for Cebus capucinus
in the Departments of Bolivar and Sucre in northern
Colombia, and is also used in Colombia's middle
Magdalena valley for Cebus .j ... suggesting the early
influence of Jesuit missionaries in disseminating Quechua
names in various parts of Colombia.

[5] "Sajou" or "sapajou" (written with French phonetics) is
of Tupi-Guaranf origin, attesting to the influence of early
Portuguese explorers, who brought the pidgin Geral into
the Amazonian region, undoubtedly disseminated by early
Jesuit missionaries.

[6] "Sal" is a vernacular name of Tupi-Guarani origin that
was used by von Humboldt (1812a; 1812b; 1824a) for





Neotropical Primates 10(2), August 2002


the weeper capuchin monkey, identified by him as Simia
capucina [non Simia capucina Linnaeus (1758)].

[7] The Warekena Indians ("Guarekens" of Humboldt)
belong to the Maipure linguistic group of the Amazonian
Arawak family. They currently inhabit the banks of the
Cassiquiare Canal (Estado Amazonas, Venezuela), as well
as the upper rio Negro, adjacent to the Colombian and
Venezuelan borders (Estado Amazonas, Brazil) (Lizarralde,
1993). Ferreira (1974: 69-73) found what he called the
"Uerequena" in 1785 on the rios Icana and Xi&, northwest
Amazonas, Brazil.

[8] Humboldt alluded to both the Atures and Maypures
rapids.

[9] Santa Barbara is an extinct mission that was located at
the mouth of the rio Ventuari on the right bank of the rfo
Orinoco, Amazonas, Venezuela (Hershkovitz, 1949: 370).

[10] Maipures (5020'N, 67045'W) (from "maypuri"
meaning tapir [ Tapirus terrestris] in the Maipures language)
was a site originally established on the left bank (i.e. the
Colombian side) of the Orinoco alongside the Maipures
rapids, as a Jesuit cattle ranch, which was converted into
a Jesuit Mission in the early 1700s. The town was founded
by Don Josd Solano at the time of the expedition of the
boundaries in 1754. Dugand and Phelps (1945) describe
some of its history. "Maipures" was the name given by the
Jesuits to the rapids as well as to a tribe of extinct Indians
who spoke an Arawakan language which has left some
toponyms in the region: i.e., "Mataven" = "black river"
(Mataven River or Creek); "Amanav&n" = "crocodile river"
(Brazo Amanaven associated with the lower rfo Guaviare,
Colombia). According to Humboldt (1852: 235) the
"Maipures" Indians called the Maipures rapids "Quittuna"
and the Atures rapids "Mapara".

Von Humboldt and Bonpland arrived for the first time
with Father Zea on the night of 18 April, 1800, and left
on the afternoon of 31 April. On their return downstream
they arrived at Maypures on 29 May and stayed until
31 May (Dugand, 1956: 315). At that time the village
consisted of fewer than 60 people living in only 7-8
huts surrounding a small church built of palm logs, but
the village had a population of about 600 inhabitants
during the time of the Jesuits, including several white
families (von Humboldt, 1852: 297, 306). The location
has become well-known as a collecting site, since von
Humboldt and Bonpland collected several new species
of plants there as well as closely observing and describing
(although erroneously, so therefore requiring us to
establish a neotype) Cebus .j ... for the first time. The
two scientists ascended the Cerro de Manimi (near the El
Tuparro National Natural Park cabana, near the mouth of
the rfo Tuparro) various times during their stay, where they
enjoyed the magnificent landscape of the Maypures rapids,
and additionally collected plants on the Cerro, particularly
the type collection of Cyperus mainimi.


Many new taxa of birds have also been collected at
Maypures by Dugand, Cherrie, and others. Fortunately
Maypures rapids, and the savanna where the village stood,
along with the many granitic hills and huge boulders, are
now protected in El Tuparro National Park (Colombia),
which is gazetted as an International Biosphere Preserve.

[11] The Mission de San Juan Nepomuceno de los Atures
was founded in 1748 by the Jesuit missionary Padre
Francisco Gonzilez.

[12] "Parmi les singes que nous vimes a la mission d'Atures,
nous en trouvames une nouvelle espece de la tribu des
Sais et de Sajous, que les Espagnols-Am&ricaines appellent
vulgairment Machis. C'est l'Ouavapavi a pelage gris et a
face bleuatre. II a les orbites et la front blancs de neige; ce
quf le distingue, au premier abord, du Simia capucina, du
Simia. a.i//i,, du Simia trepida, et des autres singes pleureurs
si confusement decris jusqufa. Ce petit animal est aussi
doux qufl est laid. 11 saisissoit tous les jours, dans la cour du
missionaire, un cochon sur lequel il restoit montd, du matin
au soir, en parcourant les savanes. Nous l'avons vu aussi sur
le dos d'un gros chat qui avoit &td dlev6 avec lui dans la
maison de pere [Bernardo] Zea."

The text of the mentioned footnote 1 is as follows:
"Voyez ma monographie des singes de l'Orenoque, dans
le Red[ueil] d'obs[ervations de] Zool[ogie et d'Anatomi
comparee, Tom[e] I, p. 324 et 563 additionn] in -40).
L'Ouavapavi (mot de la Langue guareken) est mon Simia
albifrons, ex albo cinerascens, vertice nigro, facie caerulea,
front et orbitis niveis, cruribus et brachiis fuscescentibus."

[13] There is no record of the occurrence of Cebus
,.f... on the Venezuelan bank of the Orinoco below
San Fernando de Atabapo or the neighborhood of Atures.
Cebus nigrivitattus is known on the east bank by only one
specimen, collected 32 km south of Puerto Ayacucho at
135 m (Handley, 1976: 42; see also Bodini and Perez-
Hernandez, 1987; Bodini, 1989). Further research is needed
to clarify the Venezuelan distribution of the species.

[14] A karyotype of an Aotus captured on the banks of
the rio Orinoco showed a diploid number of 50, but the
chromosomes were organized in a manner that suggests
that it was not A. brumbacki. Hershkovitz (pers. comm.)
felt that the specimen might be A. trivirgatus (Defler and
Bueno, in prep.).

[15] In the original description of Cebus fiavus E. Geoffroy
Saint-Hilaire (1812: 111) Simia flavia Schreber (1774.
p.xxxi.-b) is mentioned as a validly proposed name based on
a color plate which illustrates a pale brown specimen with
an almost white coronal cap. The origin of that specimen is
unknown and it is not certainly identifiable as a Cebus. The
citation of Simia flavia Schreber in the account of Cebus
flavus by E. Geoffroy Saint-Hilaire implies that the latter
author identified C. flavus with S. flavia; thus the epithet
flavia was retained and emended to flavus when transferred





Neotropical Primates 10(2), August 2002


to the genus Cebus to fit the required grammatical
concordance. For this purpose the -ibefore the termination
-us was deleted, so that flavia became flavus. Both epithets
are homonyms under article 58.15 of the 4th edition of the
International Code of Zoological Nomenclature.

Simia flavia Schreber (1776: pl.) was regarded as
unidentifiable by Cabrera (1917a: 233; 1958: 170) and
Hershkovitz (1949: 336, 345), and Cercopithecus flavus
Goldfuss 1809 (non Cercopithecusfiavus Boddaert 1784) is
based on the Schreber color plate as indicated by Hershkovitz
(1949: 336) and thus is an absolute synonym of Simia
flavia. The fact that E. Geoffroy Saint-Hilaire (1812: 111)
mentioned Simiaflavia Schreber (1776) in his description
of Cebusfiavus implies that the latter could be regarded as a
nomenclatural amendment. However, according to Article
58 of the International Code of Zoological Nomenclature
(ICZN, 1999: 60-61) the amended form flavus would not
fall into homonymy of flavia (or flavius).

Cebusfiavus is, therefore, a valid name based on a mounted
specimen (with skull inside) said to be from Brazil and
designated by Rode (1938: 231) as a "type" preserved in
the MHNP (no. 362 of the types catalogue, and 458 of
the general collection), which according to Hershkovitz
(1949: 342) "is extremely faded with considerable portions
of hair of the underparts, head and face missing". The
specimen is part of the collection made by the Brazilian
naturalist Alexandre Rodrigues Ferreira (1974) during his
"Viagem Filos6fica", through the states of Amazonas, Mato
Grosso, Pard and Rond6nia, Brazil. The collection was in
the Museu Real d'Ajuda in Lisbon, until it was taken to
Paris as war booty by Napoleon's troops under the care of E.
Geoffroy Saint-Hilaire. Wagner (1855: 90) suggested that
Cebus flavus was identical to Cebus gracilis Spix (1823), a
possibility that Hershkovitz [1949: 341] accepted, writing:
"In any case, the question remains whether the specimen
determined as flavus by [E.] Geoffroy [Saint-Hilaire] is
to be regarded as a specimen referring to the amended
form of the name [Simia] flavia Schreber". In summary,
therefore, Cebus flavus E. Geoffroy Saint-Hilaire 1812
would be the earliest available name for the Amazonian
populations of Cebus j ... if our interpretation of the
nomenclatorial rules is correct. In order to preserve Cebus
flavus it would be necessary to reinstate the validity of the
former epithet. However, unicolor gained wide acceptance
after Hershkovitz' publication in 1949. Groves (2001: 148)
concurs that the type Cercopithecusfiavus Goldfuss is indeed
probably albifrons.

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Neotropical Primates 10(2), August 2002


PRIMATES OF THE JAU NATIONAL PARK, AMAZONAS,
BRAZIL


Adrian A. Barnett, Sirgio H. Borges
Carolina V de Castilho, Fernanda M. Neri
Rebecca L. Shapley


Introduction


The Jad National Park is some 220 km north of Manaus
(see Fig. 1) in the state of Amazonas, Brazil. At 2,272,000
ha it is one of the largest rainforest national parks in the
world (Borges et al. 2001). Bordered on the north by the
Rio Uninf and to the south by the Rio Carabinani, the
park comprises the complete drainage basin of the Rio Jad.
In addition to primary lowland tropical rainforest (70%),
it has the following natural habitat types: black-water
inundated forest (. -.' (12%), aningaland Mauritia palm
(buritizal) swamps (approx. 0.5%), and white-sand forest
(campinarana) and scrub (campina) (>0.1%) (FVA, 1998;
FVA-IBAMA, 1998; FVA, unpubl. data). Dwellings and
associated agricultural areas comprise a disturbed habitat
estimated to cover less than 0.5% of the park's area. The
vegetation types in the remainder of the area (some 18%)
have yet to be classified. About 800 people live in the park
(0.04 people/ha: FVA-IBAMA, 1998). This is a low density,
some 25% the average human population density for rural
Amazonia (Chapman and Peres, 2000).

Although primate research is a priority under the park's
management plan (FVA-IBAMA, 1998), there has been
little published work with the exception of on-going studies
of the golden-backed uacari, Cacajao melanocephalus
ouakary (see Barnett et al., 2000; Barnett et al., submitted).
A number of unpublished reports exist, but there is no
published summary of information of all the primates
known to occur within the park. In the hope of stimulating
further studies, we here bring together information from
the following documents relating to primates within the Jad
basin (the location of the study sites for each of the surveys
appears in Figure 2).

1. A brief survey by Anthony B. Rylands (17-21 April
1992) of the lower Rio Jad, including fieldwork and
interviews (Rylands, 1992)

2. A series of interviews conducted by S&rgio Borges and
Fernanda Neri on the hunting practices and the inhabitant's
knowledge of primates along the Rio Uninf (17 March-15
April 1998) (Neri and Borges, 1998). Fourteen long-term
park inhabitants in seven villages were interviewed. This
data was supplemented by short field surveys, walking trails
around villages where interviews were made.

3. Information collected during two short field surveys of
golden-backed uacaris (late August 1999, wet season, and


-4- 7/
i ---' <



Figure 1. Location of Jatd National Park in Central Amazonia.

20 October-7 November 2000, dry season) by Barnett
(1999) and Barnett and Castilho (2000) on the lower Rio
Jad in the region of Lake Miratucd (1999) and above the
village of Seringalzinho (2000). Data was collected by direct
observation and through interviews with nine people living
in the park.

4. Observations made by Yuri L. R. Leite, James L. Patton,
Maria Nazareth da Silva and Vera Vidigal during a small
mammal survey of Jad in May-June 1996 (see Silva and
Patton, 1996).

Nine primate species are known from the park. The
available information on them is summarized below.
Trinomial nomenclature follows Rylands et al. (2000).

Primates in the Jani National Park

Saguinus inustus, mottle-faced tamarin, soim
Reported as possibly present by Rylands (1992) on the basis
of interviews with local people on the lower Rio Jad, who,
however, considered it rare. Neri and Borges (1998) received
similar reports at four of the seven communities they visited
on the Rio Uninf. All the people along the Rios Jad and
Uninf indicated that it is restricted to the middle course
of the rivers to their headwaters. This being true, Saguinus
inustuswould be restricted to the western part of the park, a
pattern also detected for a number of bird species (Borges et
al., 2001). Confirmation would represent a range extension
to the east in the interfluvial basin between the Rios Negro
and Japuri-Solim6es (see Emmons and Feet, 1997). The
most easterly locality to date is the Lago Amana, north bank
of the Rio Japuri (A. B. Rylands, pers. obs.).

Aotus sp., night monkey, owl monkey, macaco-da-noite
A pair of Aotus was seen near the locality of Macaco by the
1996 Mammal Survey (Y. L. R. Leite, pers. comm.). Carlos
Durigan, Park Director, also saw a single individual one
evening in August 1999 in igapd near the park headquarters
at the mouth of the Rio Jad. According to data in Emmons
and Feet (1997), the species in the region should be
A. vociferans (sensu Hershkovitz, 1983). Based on interviews
with local inhabitants, Rylands (1992) had also reported





Neotropical Primates 10(2), August 2002


Figure 2. Primate survey sites in Jau National Park.


this species for the park, though he did not see it. It was
reported as present to Neri and Borges (1998), with
informants indicating group sizes of around six individuals
(see Kinzey, 1997).

Saimiri sciureus cassiquiarensis, squirrel monkey, macaco-
de-cheiro
The 1996 mammal survey team recorded the species near
Macaco (Y. L. R. Leite, pers. comm.). Neri and Borges
(1998) saw a 20-strong group in terra firme on the Rio
Uninf and reported that local inhabitants considered the
species common in both terra firm and igapd. In August
1999 a group of 30+ was seen in the trees among the ruins
of the abandoned town of Velho Airao, just outside the
park and another group was seen in still-flooded igapd in
the dry season of 2000. Squirrel monkeys are reported to
sometimes travel with groups of C. m. ouakary. Rylands
(1992) and Neri and Borges (1998) both reported that
squirrel monkeys are widely hunted, and locally-caught
animals are kept as pets by park inhabitants.

Cebus albifrons albifrons, white-fronted capuchin,
caiarara
In the dry season, C. j .. is reported to enter
unflooded igapd to eat the eggs of Podocnemis turtles,
raiding nests at nesting beaches. Raided nests were seen,
apparently excavated by small primate-like hands. But no
direct observations of obphagy have yet been made by us.
It may also eat the fruits of the palm Leopoldinia pulchra
at this time. Like C. ,'Ic//, C. a. ,-j ... is reported to
forage for the large earthworms that live in the fiber and
frass enclosed by the remnant frond bases on Leopoldinia
palm trunks when the igapd is inundated. Such earthworms
have been observed by one of us (C. de Castilho),
though their predation has not. C. j ... has not
been recorded from igapd in the flooded season. At this
time it is seen in campinarana and terra firme forest
(Barnett and Castilho, 2000). S&rgio Borges recorded
C. .j ... on six occasions in the Park in the last
two years; four of these were in campinarana forest.
White-fronted capuchins have not been reported to
be white-sand specialists (see Kinzey, 1997), but it is
suspected that, at Jad, it frequently enters campinarana,


S100 IN I


perhaps for specific food resources not exploited by
C. ,'ci//, though more field data is needed to confirm
this. Neri and Borges (1998) reported a locally-caught
animal being kept as a pet. Along with Saimiri sciureus,
C. a. ,.j ... was considered the second-most common
primate by inhabitants interviewed by Neri and Borges
(1998). It was reported as present by Rylands (1992).

Cebus apella apella, brown capuchin monkey, macaco-
prego
A group of four was seen in igapd near Seringalzinho,
and a locally caught juvenile was being kept as a pet on
a riverboat moored near Lago Miratucd. Local people
reported that it forages for the large earthworms that
live in the fiber and frass enclosed by the remnant frondbases
on Leopoldinia palm trunks when the igapd is inundated.
C. a. ,1,e//, eats the seeds from the woody fruits of
Couratari sp. (Lecythidaceae), banging the pyxidium on
a branch until the operculum comes free and the seeds
can be extracted, a behavior Peres (1991) has reported for
C. a,''//. elsewhere in Amazonia. In doing this, it may
well be in competition with C. m. ouakary, which opens
the fruits with its teeth. Neri and Borges' informants
considered C. a. ,,c//.f to be the park's most common
primate.

Callicebus torquatus lugens, yellow-handed titi, zogue-
zogue
Seen by Adrian Barnett and S&rgio Borges on the trail
near Seringalzinho on the morning of 26 August 1999.
A single individual was seen in a Mauritia palm in
campinarana close to a large squirrel. Though a clear view
was not obtained of the diagnostically-yellow hands, it
was considered to be this species based on colour, shape
and behaviour. This sighting confirmed the reports of
local inhabitants to Neri and Borges (1998). The record
is a slight westward range extension, within the Negro-
Solimoes/Japuri interfluvial basin (see Emmons and Feer,
1997). Local people informed Neri and Borges (1998) that
it is uncommon in the park.

Pithecia pithecia chrysocephala, golden-faced saki,
parauaci or macaco velho
Reported as present by Rylands (1992), based on interviews.
It was considered rare by all the people interviewed by Neri
and Borges (1998) and by Adrian Barnett in 2000. The
species was generally reported to occur only well away from
river margins, deep in the terra firme forest of interfluvial
basins. It was seen by Hilton Nascimento and Antenor
Anicacio in March 2000 (pers. comm.) in terra firme
forest, and by S&rgio Borges in May 1997 in campinarana
vegetation, and by Maria Nazareth Silva in June 1996.
All three sightings occurred at points adjacent to the Rio
Jad. Though the subspecies has long been known from
the region (see Hershkovitz, 1987), according to maps in
Emmons and Feer (1997), this record represents a slight
range extension, and the westernmost known population of
P. p. chrysocephala. Since species of Pithecia have never been
recorded in sympatry (see Kinzey, 1997), the presence of





Neotropical Primates 10(2), August 2002


P pithecia in Jad confirms that the range of the buffy saki
(P albicans) does not extend into the inter-fluvial area
between the northern bank of the Rio Japura and the
southern bank of the Rio Negro (see Hershkovitz, 1987).

Cacajao melanocephalus ouakary, golden-backed uacari,
bic6
Rylands (1992) reported seeing a band of approximately
10 individuals in April 1992 in both igapd and
campinarana. The group included a female with a 4-5
month-old offspring. A group of 15+ was seen by Neri
and Borges (1998) in igapd. Barnett and Castilho (2000)
observed bands of 5 to 100+, also in igapd as well as in
the terra firme, campina and campinarana. Observations,
supplemented by information from local informants,
indicate that C. m. ouakary eats parts of some 70+
plant species at Jad. These include the soft mesocarp of
Astrocaryumjauari and Oenocarpus bataua palms, the seeds
from hard shelled fruits such as Eschweilera tenuifolia and
Couroupita spp. (Lecythidaceae) and whole soft-shelled
fruits (e.g., Salacia sp. [Hippocrataceae]). In the dry
season, when little fruit is available (Ashton, 2001; Barnett
and Castilho, 2000), the diet is supplemented by leaves
(Mabea taquari [Euphorbiaceae], Buchenavia oxycarpa
[Combretaceae] and Eschweilera tenuifolia). They also raid
nests of Polistes wasps to eat the larvae.

Around half of the known fruits that C. m. ouakary eats are
soft-skinned, and most are also eaten by Cebus and Saimiri.
The woody fruits are also eaten by macaws (Ara chloroptera
and A. ararauna). Unlike populations of C. m. ouakary
on the upper Rio Negro, those at Jad are not reported to
raid the nests of freshwater turtles and eat the eggs (see
Barnett, in press). Reports indicate that they do descend to
the ground in the late dry season to eat beetle larvae and,
to a lesser extent, germinating sapotaceous seeds as has
been reported for C. c. calvus (see Ayres, 1986). The field
observations of Barnett and Castilho support interview-
based reports of Neri and Borges that C. m. ouakary spends
the wet season in inundated igapd, and migrates to terra
firme when it is dry and lacking fruit (see also Barnett and
da Cunha, 1991; da Cunha and Barnett 1990, for the upper
Rio Negro).

It is interesting to compare the observed pattern of habitat
use by C. melanocephalus ouakary at Jad, with those reported
for C. m. melanocephalus from the upper Rio Negro
(Boubli, 1999). Though C. m. ouakary has been observed
by us in white-sand vegetation (campinarana), it is much
more commonly seen in terra firme forest and flooded
igapd forest. This is in contrast with Boubli's studies, which
recorded intensive use of white sand soil vegetation, and
negligible use of igapd or terra firme.

Alouatta seniculus, red howler monkey, guariba vermelho
Two adults were seen on 24 August 1999 in igapd at Lake
Miratuc6. Howlers were also seen in terra firme at four sites
on the Rio Uninf and once in igapd by Neri and Borges
(1998). They are considered common by locals, who report


group sizes of up to 30. They were seen in terra firme forest
during the 1996 survey of Leite, Patton, Silva and Vidigal.
They were not seen by Rylands (1992), but reported as
present based on interviews.

Ateles belzebuth, white-bellied spider monkey, macaco
aranha, and Lagothrix lagothricha, woolly monkey,
macaco barrigudo?
The possibility remains that Ateles belzebuth and Lagothrix
lagothricha may occur in the park (see maps in Eisenberg
and Redford [1999] and in Kinzey [1997]), but none of
the short surveys summarized here were able to obtain any
evidence of this. The map in Fooden (1963), still the most
authoritative account for Lagothrix, extends the range to
the entire interfluvium of the Rios Japura and Negro,
but there are no collecting localities confirming this. The
easternmost localities are on the Rio Uaupes, some 600 km
north-west.

Information obtained by S&rgio Borges from a reliable
informant of Barcelos (upstream of Jad, see Fig. 1),
indicated that, south of the Rio Negro, A. belzebuth may
have its current eastern limit defined by a small river just
north of the town. Some palm species (e.g., Leopoldinia
piassaba and Barcela odora) also show this pattern of
limited eastern extension into the Negro-Solim6es/Japura
interfluvium (Henderson, 1995). This may reflect the
former distribution for A. belzebuth, so that it has never
occurred in the area covered by the current national park.
Queries to the managers of the mammal collections of the
American Museum of Natural History, Field Museum
(Chicago), Museu Goeldi (Belkm), Natural History
Museum (London), and the Smithsonian Institution
(Washington, DC) found no recorded specimens of either
Ateles or Lagothrix from the lower reaches of the Rio Negro,
nor from the Japura-Solim6es/Negro interfluvium.

However, the collection of the Museu Nacional Rio de
Janeiro (MNRJ) has eight specimens of Ateles belzebuth
(MNRJ-1702, 2491, 2456 to 59, 2499, 2500) from
Parana do Maiana, Amazonas. This locality is situated on
the Solim6es-Japura, a little north of Fonte Boa, close to
a village called 'Jacard'. Parana do Maiana lies between the
tributary rivers Auatf-ParanA and MamirauA and is close to
the headwaters of the Rio Jad.

The Museu de Zoologia da Universidade de Sao Paulo
(MZUSP) has the following specimens of Lagothrix 1.
lagothricha: MZUSP-19674 "AM, Rio Negro, 200 km
acima de Manaus", collected by A. Vertematti (no date);
"AM, Manaus" MZUSP-11232, 11233, collected by
A. Vertematti, August 1973; "AM, Manaus" MZUSP-
19676, collected by Jose Hidasi, January 1962.

These tantalizing records indicate that both Ateles and
Lagothrix might have occurred in the Jad region (or at least
on the lower Rio Negro) in the recent past. However, as
Jose de Sousa e Silva Junior (pers. comm.) has suggested,
it would seem likely that both these species may now be





Neotropical Primates 10(2), August 2002


extinct in the Jad region of the lower Rio Negro. Both these
large primates are favored by hunters Amazon-wide, and
both are extremely susceptible, having low reproductive
rates (Chapman and Peres, 2000). It is possible that
they were extirpated from the Jad river basin in the early
decades of this century when the human population of
the area was very much higher than today (see Leonardi,
1999; FVA-IBAMA, 1998). However, their existence in
the park is stoutly denied by all interviewees, even those of
considerable age. Rylands (1992) believes both A. belzebuth
and L. lagothricha may (still) occur in the far west of the
park where there are some regions uninhabited by people
(see Neri and Borges, 1998).

Other primates?
Both Neri and Borges (1998) and Adrian Barnett received
reports from well-informed local inhabitants of a small
black monkey with reddish markings on the face and
chest. This indicates the possibility that a titi monkey
besides ( .... torquatus (see above) may occur there.
The Negro-Solim6es/Japura interfluvium lies to the north
and east of the known ranges of the moloch group titi
monkeys (sensu Hershkovitz, 1988, 1990; Kinzey, 1997)
and visual confirmation is needed. Both Neri and Borges
(1998) and Adrian Barnett received reports of a second
form of Pithecia, the 'gog6-de-sola', described as similar
to P pithecia but with a naked throat. In both 1999 and
2000, Barnett received several reports of a rarely-seen
large completely black primate that fits no known taxon.
Further investigation is required to assess the meaning of
these reports. It is possible that the 'gog6-de-sola' may be
the mustelid Eira barbara or a hitherto undescribed form
of Pithecia.

Threats and Impacts

Monkeys are hunted in the Jad National Park (Neri
and Borges, 1998) as they are in most parts of the
tropics (Cowlishaw and Dunbar, 2000). We have little
information on the effects of such hunting practices on
the populations of monkeys in Jad. However, surveys
suggested that they are not a principal source of game.
Data on hunting and fishing practices in nine families in
Jad indicate that more than half of the meat in the diet of
these families was supplied by fishes and turtles, and the
principal sources of mammalian meat were ungulates and
caviomorph rodents (FVA, 1998). Neri and Borges (1998)
reported that monkey meat was generally not preferred.
Tapir and peccary were favoured. Nevertheless, one
household was personally observed (Rebecca Shapley) to
eat Cebus in 2000 and both Cebus species were reported as
being hunted on the Rio Uninf by Neri and Borges (1998).
Uacaris are hunted (Rylands, 1992), but according to Neri
and Borge (1998) are not preferred because they have little
meat, they move fast and, in flooded forests, they tend to
be lost after having been shot. Howler monkeys, widely
hunted elsewhere, are reported to taste and smell bad.
Pithecia are considered to move too fast and too high to
be worth hunting.


Compared to Asia and Africa (Cowlishaw and Dunbar,
2000), crop raiding by primates is an infrequent
phenomenon in the Neotropics (see Jiminez, 1970;
Warren et al., 1988); at Jad such incursions appear to be
minimal and primates are not hunted punitively. Lack
of financing prevents full policing of the park, and there
is no permanent conservation presence on two of the
rivers (Caribinani and Uninf). Hence, it is impossible
to assess the impact of hunting by day and weekend
trippers, who are known to make frequent excursions
to the area. There is no commercial logging in Jad,
and the extent of human-mediated habitat destruction
appears generally slight (FVA-IBAMA, 1998; Ferreira and
Prance, 1999). Regionally, Cebus and Saimiri are quite
commonly kept for pets and Cacajao rarely so. However,
such animals are often traded and it is currently unclear
how this effects the park's primates. Older inhabitants
favor the use of a suitably trimmed Cebus humerus as
a restraining wedge during the construction of fibre
baskets.

Primate Research in Jani

Nine species of primate are confirmed for Jad, with the
possibility of another five species (and two odd reports
requiring further investigation). This is a rich and
representative primate fauna for the middle Amazon (see
Mittermeier, 1987). Further research is clearly a priority.
Studies are underway on the ecology and behavior of
C. melanocephalus (Barnett et al., submitted), and on
primate densities, as part of a more general survey on the
impacts of hunting by Carlos Peres and Hilton Nascimento.
We suggest that the following studies need to be carried out
to obtain a better understanding of the primate populations
in the park.

1) Continued inventories of primates in Jad National Park,
especially in the headwaters and in remote areas such as the
uninhabited regions of the Rio Papagaio, Rio Uninf and
Rio Guariba (see Fig. 2).

2) Food and habitat preferences for all primate species in
the park.

3) Studies of seasonality of habitat use in primate species
other than C. melanocephalus.

4) Impacts of hunting on the primates of the Rio Carabinani
(Jal's third largest river).

5) The dynamics of egg predation by C. .j... and
nesting success of Podocnemis spp. and other Chelonia in
the Park.

Field survey work should also answer the following
questions: Are Ateles belzebuth and Lagothrix lagothricha
really absent from the park, are there any historical records
of their presence there or in the immediate region? Does a
member of the ( .... moloch group titi monkeys occur





Neotropical Primates 10(2), August 2002


in the park? If confirmed, this would be an eastward range
extension of an extent similar to those reported by Borges
and Carvalhaes (2000) and Borges et al. (2001) for several
bird species otherwise considered confined to the upper Rio
Negro. Do the reports of forms such as the 'gog6-de-sola'
represent new species or merely variants of existing species?
Given the large number of newly-discovered Amazonian
primates in recent times this possibility should certainly be
investigated.

Acknowledgments

Adrian Barnett, Carolina de Castilho and Rebecca
Shapley acknowledge the kindness, good guidance and
knowledge of Sr. Antenor Anicacio, who acted as guide
and informant during two visits to Jad. We thank S&rgio
Sa at IBAMA, Muriel Saragoussi, Executive Director of
the Fundagao Vit6ria Amazonica (FVA), and her staff,
especially Jose Luis Camargo and Osmar Silva. We also
thank staff of the INPA Herbarium for help with plant
identifications. Adrian Barnett thanks Ruediger Schmelz
for identifying the worms, and James L. Patton and Yuri
Leite for contributing their unpublished field observations.
We are also grateful to the following: Christopher Norris
(American Museum of Natural History), Bruce Patterson
(Field Museum, Chicago), Suely Marques (Museu Goeldi,
Belkm), Jean Philippe Boubli (Museu National, Rio de
Janeiro), Jose de Sousa e Silva Jdnior (Museu Goeldi),
Richard Harbord (Natural History Museum, London),
Linda Gordon (Smithsonian Institution) and Andrew
Collins (University of Wisconsin-Milwaukee). Sergio
Borges thanks Maria Nazareth da Silva for her primate
observations from Jad, and Marc G. M. van Roosmalen
for helpful discussion regarding Pithecia. Funding for
fieldwork by Barnett, Castilho and Shapley was provided
by Akodon Ecological Consulting, The Mammal Society
(UK), Margot Marsh Biodiversity Foundation, Percy
Sladen Memorial Fund, Primate Conservation Inc., the
University of Surrey, Roehampton, UK, and (in-kind)
by FVA.

Adrian A. Barnett, School of Life Sciences, University
of Surrey, Roehampton, London SW15 3SN, England,
UK, and Dept. of Anthropology, California Academy of
Sciences, Golden Gate Park, San Francisco, CA 94118,
USA (Research Associate), e-mail: ,
S&rgio H. Borges, Departamento de Zoologia, Museu
Paraense Emflio Goeldi, Av. Perimetral 1901/1907,
66077-530 Belkm, Brazil, Carolina Volkmar de Castilho,
Departamento de Botanica, Instituto de Nacional Pesquisas
da Amazonia, Caixa Postal 478, 69083-000 Manaus,
Amazonas, Brazil, Fernanda Maria Neri, Universidade
Federal de Sao Carlos (UFSCar) Programa de P6s-
graduacao em Ecologia e Recursos Naturais, 13565-905
Sao Carlos, Sao Paulo, Brazil, and Rebecca L. Shapley,
Akodon Ecological Consulting, Walnut Creek, CA 94518,
USA.


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Neotrop. Primates 8: 61-93.
Silva, M. N. da and Patton, J. L. 1996. Pequenos mamfferos
do Parque Nacional do Jad. Unpublished report,
Fundagao Vit6ria Amaz6nica, Manaus. 12pp.


CUIDADO BIPARENTAL EN EL MONO DE NOCHE
(AOTUS AZARAI) DE FORMOSA, ARGENTINA


Marcelo Rotundo
Eduardo Fernandez-Duque
Marina Gimenez


Introducci6n


Los monos de noche (Aotus spp.) viven usualmente
en grupos que incluyen de 2 a 5 individuos (Aquino y
Encarnaci6n, 1994; Fernandez-Duque et al., 2001). El
genero, que se encuentra distribufdo desde Panama al
noreste de Argentina, es el dnico que present habitos
nocturnos en el nuevo mundo (Wright, 1989). Los grupos
son aparentemente mon6gamos con una dnica hembra
reproductive que produce un infante por afio (Fernandez-
Duque et al., 2002). Algunos studios con animals en
cautiverio demostraron una gran participaci6n del macho
en el cuidado del infante recidn nacido (Dixson y Fleming,
1981; Wright, 1984).

A partir de esas observaciones de Aotus spp. en cautiverio y
de otras species de primates socialmente mon6gamas con
intensive cuidado paternal (Fragaszy et al., 1982; Hoffman
et al., 1995; Mendoza y Mason, 1986), se ha hipotetizado
que el cuidado intense del infante por parte de los machos
operarfa como una fuerza selective que podrfa favorecer
la evoluci6n de la monogamia. Los machos, al colaborar
con el cuidado del infante, obtendrian un mayor dxito
reproductive que si trataran de aparearse poligfnicamente
(Clutton-Brock, 1989).

Hasta el present no se disponfa de informaci6n sobre el
cuidado biparental en poblaciones silvestres de Aotus spp.
Esto probablemente se deba a que existen dos caracteristicas
del gdnero que limitan la obtenci6n de informaci6n sobre
el comportamiento social de machos y hembras. En primer
lugar, el mono de noche no present un dimorfismo
sexual detectable en el campo haciendo casi impossible la
identificaci6n de machos y hembras. A esto se suman los
habitos estrictamente nocturnos del gdnero en la mayor
parte de su distribuci6n geografica.

El objetivo de este trabajo fue describir el cuidado biparental
del infante de Aotus azarai a partir de observaciones de
individuos identificables realizadas durante el dfa. Esto
fue possible gracias a que, en el extreme austral de su
distribuci6n, el gdnero es catemeral (Tattersall, 1987),
presentando actividad tanto durante el dfa como la noche
(Arditi, 1992; Rotundo et al., 2000; Sloan y Fernandez-
Duque, 1999; Wright, 1989).

M6todos

Area y poblaci6n de studio
El studio se llev6 a cabo en la Estancia Guaycolec, al sudeste
de la provincia de Formosa en el Gran Chaco Argentino
(2554'S, 58013'O; Fig. 1). El primer studio sobre





Neotropical Primates 10(2), August 2002


PARAGUAY N
+


CHACO


0 = 10K


Figura 1. Area de studio.


A. azarai en dicha estancia se realize en 1977 (Rathbun
y Gache, 1980). Durante la siguiente decada, hubo otros
studios realizados por investigadores argentinos (Arditi,
1992; Arditi y Placci, 1990; Zunino et al., 1985).

En 1996, se dio inicio al Proyecto Mirikina (nombre
comin que recibe Aotus en dicha zona) con el objetivo
de llevar adelante studios a largo plazo sobre la ecologfa,
comportamiento y gen&tica de dicha especie (Fernandez-
Duque y Bravo, 1997; Fernandez-Duque et al., 2002;
Fernandez-Duque et al., 2001; Huntington y Fernandez-
Duque, 2001; Rotundo et al., 2000; Sloan y Fernandez-
Duque, 1999).

La poblaci6n de studio incluye 15 grupos sociales que
habitan las selvas en galerfa del Riacho Pilaga. Durante 1999,
se estudi6 el desarrollo de 9 infants hasta los seis meses de
vida (Rotundo y Fernandez-Duque, datos no publicados) y
el cuidado biparental de una de esas crias. Aquf se presentan
los resultados concernientes al cuidado biparental.

Observaciones
Los datos sobre cuidado biparental fueron tornados
de un grupo compuesto por un macho y una hembra
adulta, un juvenile del afio anterior y el infante nacido
durante el studio. En dicho grupo se habia identificado
inequivocamente al macho y a la hembra a partir de
observaciones previas de c6pula y amamantamiento. El
macho tenfa la cola significativamente mas corta que los
demas individuos en el grupo, lo que tambikn facility su
identificaci6n. Se pudo confirmar luego el sexo del individuo
cuando fue capturado, marcado y liberado con radio-collar.
El grupo, acostumbrado a la presencia de observadores, fue
monitoreado regularmente cada tres a cinco dfas a partir del
comienzo de octubre para establecer con exactitud la fecha
del nacimiento del infante. Se realizaron entire dos y tres
observaciones semanales durante las primeras 18 semanas
de vida del infante entire octubre de 1999 y marzo del 2000.
Se obtuvieron 47 horas de observaciones.


Durante dichas horas, se realizaron observaciones focales
del infante y del individuo mas cercano a este. Cada dos
minutes, al sonar de un indicador sonoro, se registry
si el infante se hallaba dependiente o independiente.
Se consider al infante como dependiente cuando el
s mismo tenfa dos o mas extremidades apoyadas sobre
otro individuo. A su vez, el infante estaba independiente
cuando tenfa una o ninguna extremidad en contact con
otro individuo. En caso de estar independiente se registry
la distancia que separaba al infante del individuo mas
cercano, asf como la identidad de este 6ltimo. Si el infante
studio estaba dependiente se registry la identidad del individuo
que lo llevaba a cuestas. Los perfodos de amamantamiento
y de compartir comida se registraron de manera continue.
Aunque los resultados son presentados como "tiempo",
M. en realidad se trata de ndmero de puntos muestrales, a
excepci6n del amamantamiento que fue medido en tiempo
absolute.


Resultados y Discusi6n

Los resultados indican un cuidado intenso del infante por
parte del macho. A excepci6n de la primer semana de vida,
durante el resto del tiempo el infante fue transportado
principalmente por el macho, quien lo transport en el
87% de las observaciones en las que el infante estuvo
dependiente (398 de 456 observaciones, Fig. 2). Durante la
primer semana de vida la hembra lo llev6 a cuestas la mayor
parte del tiempo transportandolo durante el 67% de las
observaciones (31 de 46). Nuestras observaciones coinciden
plenamente con los datos obtenidos en cautiverio (Dixson
y Fleming, 1981).

Cuando el infante comenz6 a independizarse y a
desplazarse por sf mismo, sigui6 prefiriendo mantenerse
pr6ximo al macho. En el 70% de las observaciones (145
de 207), el individuo mas cercano al infante fue el macho,
independientemente de cual fuera la distancia a dicho
individuo (Fig.3).

Observaciones cualitativas refuerzan la noci6n de un fuerte
vinculo entire el macho y la cria. En general el infante


60 M Hembra adulta
S0 lMacho adulto
o 40



1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18
Edad del infante (semanas)


Figura 2. Ndmero de puntos de muestreo ("observaciones") en
los que el infante fue observado transportado por el macho o la
hembra durante las primeras 18 semanas de vida.





Neotropical Primates 10(2), August 2002


80 i DMacho
g 60 U Hembra
0 40
20 -

E en 0-0.5 0.5-1 1-3 3-5
Z contact
Distancia al individuo mis cercano (mt)

Figura 3. Ndmero de puntos de muestreo ("observaciones") en los
que el macho y la hembra fueron los individuos mis cercanos al
infante para las diferentes distancias.

fue responsible de mantener una corta distancia con el
macho, ya que lo segufa en sus desplazamientos. Cuando
se agrupaban para dormir, el macho casi siempre llev6 al
infante en forma dorsal y el juvenile nacido en el afio anterior
se acomodaba en contact con este. Por el otro lado, la
hembra a veces descansaba separada en el mismo arbol o
en otro arbol cercano. El alejamiento de la hembra de a
moments es particularmente notable. En una oportunidad,
durante dos horas y media s6lo se pudo observer al macho
junto al infante y al juvenile del afio anterior pero en
ning6n moment se observ6 a la hembra. Los events
de amamantamiento (n = 9) fueron relativamente cortos
(media d.s. = 74 34 seg.) y el infante, luego de mamar,
por lo general volvi6 junto al macho.

El elevado porcentaje de tiempo que el infante pas6 en
cercanfa fisica con el macho adulto sugiere un important
rol del mismo en el desarrollo del comportamiento de
forrajeo del infante. Por ejemplo, en las cuatro ocasiones
que se observ6 al infante compartir comida lo hizo con el
macho; es possible que el macho sea el modelo a imitar.

La relativamente alta sobrevivencia de infants en esta
poblaci6n hastalos seis meses de vida (96%, n = 27 infants)
sugiere que el cuidado paternal puede ser una estrategia
exitosa para maximizar el dxito reproductive del macho
(Fernandez-Duque et al., 2002). Si bien los resultados aquf
presentados estan referidos a un dnico grupo, constitute
este el primer trabajo que evalda el cuidado biparental en un
grupo silvestre de Aotus y por lo tanto proporciona una base
para trabajos futures. Por ejemplo, es imperative confirmar
la eventual relaci6n gen&tica entire el macho y el infante,
para lo cual durante los pr6ximos afios se capturaran y
obtendran muestras de material gen&tico de todos los
individuos en la poblaci6n.

Agradecimientos

Agradecemos a todo el personal de la Estancia Guaycolec y
de Pilaga S.A. Ganadera, a las autoridades de la direcci6n
de fauna de la Provincia de Formosa y de la direcci6n de
fauna de Argentina por su apoyo. EFD agradece el apoyo
recibido de la Sociedad Zool6gica de San Diego y el
Conicet de Argentina (PIP 0051/98) que hicieron possible


el studio. Finalmente, agradecemos la colaboraci6n de los
muchos estudiantes argentinos que participaron del studio
y los comentarios de Claudia Valeggia para mejorar el
manuscrito.

Marcelo Rotundo, Fundaci6n ECO, Josd Maria
Uriburu 374, Formosa (3600), Argentina, e-mail:
, Eduardo Fernandez-Duque,
Sociedad Zool6gica de San Diego, California, USA, y
Centro de Ecologfa Aplicada del Litoral, Argentina, e-mail:
, y Marina Gimrnez, Fundaci6n
ECO, Josd Maria Uriburu 374, Formosa (3600), Argentina,
e-mail: .

Referencias

Aquino, R. y Encarnaci6n, F 1994. Owl monkey
populations in Latin America: Field work and
conservation. En: Aotus: The Owl Monkey, J. F Baer,
R. E. Weller and I. Kakoma (eds.), pp.59-95. Academic
Press, San Diego.
Arditi, S. I. 1992. Variaciones estacionales en la actividad
y dieta de Aotus azarae y Alouatta caraya en Formosa,
Argentina. Boletin Primatoldgico Latinoamericano 3:
11-30.
Arditi, S. I. y Placci, G. L. 1990. Habitat y densidad de
Aotus azarae y Alouatta caraya en Riacho Pilaga, Formosa.
Boletin Primatoldgico Latinoamericano 2: 29-47.
Clutton-Brock, T. H. 1989. Mammalian mating systems.
Proc. Roy. Soc. Lond. B. 236: 339-372.
Dixson, A. F y Fleming, D. 1981. Parental behaviour and
infant development in owl monkeys (Aotus trivirgatus
griseimembra). J. Zool., Lond. 194: 25-39.
Fernandez-Duque, E. y S. Bravo. 1997. Population genetics
and conservation of owl monkeys (Aotus azarai) in
Argentina: A promising field site. Neotrop. Primates 5:
48-50.
Fernandez-Duque, E., Rotundo, M. y Sloan, C. 2001.
Density and population structure of owl monkeys (Aotus
azarai) in the Argentinean Chaco. Am. J. Primatol. 53:
99-108.
Fernandez-Duque, E., Rotundo, M. y Ramirez-Llorens, P.
2002. Environmental determinants of birth seasonality in
owl monkeys (Aotus azarai) of the Argentinean Chaco.
Int. J. Primatol. 23: 639-656.
Fragaszy, D. M., Schwarz, S. y Shimosaka, D. 1982.
Longitudinal observations of care and development of
infant titi monkeys (C .... moloch). Am. J. Primatol.
2: 191-200.
Hoffman, K. A., Mendoza, S. P., Hennessy, M. B. y Mason,
W. A. 1995. Responses of infant titi monkeys, ( .-
moloch, to removal of one or both parents: Evidence for
paternal attachment. Develop. Psychobiol. 28: 399-407.
Huntington, C. y Fernandez-Duque, E. 2001. Natal
dispersal in the monogamous owl monkey (Aotus azarai)
of Formosa, Argentina. Am. J. Phys. Anthropol. 32(suppl.):
83-84.
Mendoza, S. P. y Mason, W. A. 1986. Parental division
of labour and differentiation of attachments in a





Neotropical Primates 10(2), August 2002


monogamous primate (C .... moloch). Anim. Behav.
34: 1336-1347.
Rathbun, G. B. y Gache, M. 1980. Ecological survey of
the night monkey, Aotus trivirgatus, in Formosa Province,
Argentina. Primates 21: 211-219.
Rotundo, M., Sloan, C. y Fernandez-Duque, E. 2000.
Cambios estacionales en el ritmo de actividad del mono
mirikina (Aotus azarai) en Formosa, Argentina. En:
Manejo de Fauna Silvestre en Amazonia y Latinoamerica,
E. Cabrera, C. Mercolli and R. Resquin (eds.), pp.413-
417. Asunci6n, Paraguay.
Sloan, C. y Fernandez-Duque, E. 1999. Cathemerality in
Argentinian owl monkeys. AnthroQuest 8: 1-3.
Tattersall, I. 1987. Cathemeral activity in primates:
A definition. Folia Primatol. 49: 200-202.
Wright, P C. 1984. Biparental care in Aotus trivirgatus and
( .... moloch. En: Female Primates: Studies by Women
Primatologists, M. Small (ed.), pp.59-75. Alan R. Liss,
New York.
Wright, P C. 1989. The nocturnal primate niche in the
New World. J. Hum. Evol. 18: 635-658.
Zunino, G. E., Galliari, C. A. y Colillas, 0. J. 1985.
Distribuci6n y conservaci6n del mirikina (Aotus azarae),
en Argentina: Resultados preliminares. En: A Primatologia
no Brasil-2, M. T. de Mello (ed.), pp.305-316. Sociedade
Brasileira de Primatologia, Brasilia.



BEHAVIOR OF SQUIRREL MONKEYS (SAIMIRI
SCIuREUS) 16 YEARS ON AN ISLAND IN FRENCH
GUIANA


Benoit de Thoisy, Olivier Louguet
Francoise Bayart, Hugues Contamin


Introduction


Squirrel monkeys (Saimiri spp.) are small frugivore-
insectivores widely distributed in the Amazon basin and
through the Guiana Shield. They occur in a number
of different habitats: primary and secondary forests,
mangroves, and remnant forests (Baldwin and Baldwin,
1981). The main features of their ecology are similar
throughout their range, although Boinski (1999) correlated
some differences in key aspects of the social organization
with biogeographic variations in fruit availability.

Since the late seventies, the Pasteur Institute of French
Guiana has used the squirrel monkey as an experimental
model for the study of human malaria. In addition to
the captive colony, the Institute manages an island
where 150 wild squirrel monkeys originating from
French Guiana and Suriname were introduced in 1981.
There were several reasons for the establishment of this
free-ranging population: (i) as a supply of animals for
experimental needs which could be easily trapped; (ii)
to accommodate older and post-experimental animals;
(iii) since a part of the area can be visited, this site is an
opportunity to educate people about primates and medical


research; and (iv) the population is isolated from major
infectious risks, allowing for a rapid re-establishment of
the captive colony in the case of an epidemic (de Thoisy
and Contamin, 1998).

Field work was conducted on the island in 1997 in order
to assess the status of the population 16 years after the first
release. Our goal was firstly to assess the potential of the
island in fulfilling the objectives outlined above, but also to
study the main eco-ethological patterns of these insularized
monkeys. A report on their feeding behavior will be
provided in a second paper.

Methods

The "Ilet-La-Mere" is a 56-ha, island offshore from
Cayenne (454'N, 5212'W), French Guiana (de Thoisy
and Contamin, 1998). The island is covered by dense
secondary forest with numerous lianas and epiphytes, and
the tree community is dominated by Spondias mombin
(Anacardiaceae), S, c~h,'f.i' morototoni (Araliaceae), Cecropia
obtusa (Cecropiaceae), Ficus spp. (Moraceae), and such
introduced species as Mangifera indica (Anacardiaceae) and
Carica papaya (Caricaceae).

The study was carried out during the rainy season, from
April to July 1997. Two free-ranging populations live on the
island: wild monkeys (throughout), and food-provisioned
monkeys in a 2-ha area around the camp. The wild
population was censused through ad libitum observations
and subsequently through direct observations of one focal
troop (Tl) and by radio-tracking two other troops (T2 and
T3). One adult female of each group was trapped and radio-
collared (Telonics, model 1A), and located three times a
day by triangulation (Harris et al., 1990). The fourth troop
(T4) was located and censused just once. The provisioned
population is about 90 monkeys, all in a single troop, but
the composition was not determined.

Troop T1 was studied for 101 hours spread equally during
the day, using the scan-sampling method (Altmann, 1974).
The locations of the individuals, both in terms of the
height in the forest and where they were in the home range,
and their behavior (foraging and feeding, locomotion,
resting, and social behavior (including agonistic and
affiliative interactions) were recorded every 10 minutes.
Vertical use of space and the different behaviors were
correlated using a Factors Correspondence Analysis.
Home range use was determined using the 50 x 50 m
grid-cell method. Daily travel distances were calculated by
measuring the distance between the centers of successive
grid-cells crossed. The home ranges of T2 and T3 were
determined using the convex polygon technique (White
and Garrott, 1990).

Results

The spatial distribution of the squirrel monkey population
is shown in Figure 1. The home range sizes ofT1, T2 andT3





Neotropical Primates 10(2), August 2002


were 12.3, 17.5, and 20 ha, respectively. TI was comprised
of two adult males, two subadult males, eight adult females,
five juveniles and six infants. T2 was comprised of 12
adults and subadults, and five infants; T3 of 27 adults and
subadults, and three infants; T4 of 15 adults and subadalts,
and five infants. The total population was 90, equivalent to
a density of 164 individuals/km2.

The core area ofT1 was 3 ha, that is 24.5% of its home
range with 60% of recorded locations (Fig. 1). Feeding and
foraging took up 63% of their daily activity, while 20.5%
was given over to locomotion, 12.5% to resting and 4%
to social behaviors. The daily travel distance was 4.5 km.
They spent most time in the lower levels of the forest,
with 41.7% of records during scans in the undergrowth,










N H HE M e
0000 **1^^^^^^^^
go.. ^ *]-1- ^^ ^
c IS^ :p 62:::::| ^


31.5% in the low canopy, and only 26.8% in the upper
levels (chi-square = 7.04, df = 2, p<0.05). They traveled
more in the lower canopy and fed more in the upper
levels. Resting and social behaviors were more frequent
in the undergrowth (Fig. 2, (p = 0.28, horizontal axis
contribution = 79%, vertical axis contribution = 21%).
In troop T1, males tended to be peripheral. Fifteen ago-
nistic behaviors were recorded, nine involving a male and
a female, one involving two males, and five involving fe-
males. Once, a coalition of females was observed pursuing
a male. Twenty-six affiliative coalitions were observed, 16
involving adult females. Five infants were born at the end
of the rainy season, and one later at the beginning of the
dry season. Male fatting and sexual behavior were not seen


Left side of island: Troop 1 Right side of island
0 < n < 6 commensal population


i5 < n < 16

_15 < n < 26

S25
*36
Score area


Troop 2

Troop 3


U Troop 4: ad hoc locations


n = number of troop locations in
each grid cell, during the study period


Figure 1. Spatial organization of the squirrel monkey population (Saimiri sciureus) on the Ilet-la-Mere, French Guiana.





Neotropical Primates 10(2), August 2002


0,5
0,4
0,3
0,2
0,1
0
-0,1
-0,2
-0,3
-0,4


during the study period. No interactions between troops
were recorded.

Discussion

Over the years, approximately 300 monkeys have been
released on the island, and the free-ranging population
today is close to 100 individuals. Newborns are numerous,
suggesting that fertility is certainly not a limiting factor.
There are no predators, and carrying capacity of the island
should therefore be the limiting factor to population size,
although infectious diseases and the survival rate of newly-
released animals are also important in controlling population
increase. Consequently, the initial objectives of Ilet-la-MWre
of both the capture of naive animals for experimentation
and the release of old animals as "retirement", probably
need to be reevaluated.

The habitat preferences, locomotory behavior, foraging
techniques and food preferences, are similar to those
observed for Saimiri oerstedii (Costa Rica), S. boliviensis
(Peru), and S. sciureus (Suriname) (Janson and Boinski,
1992; Boinski, 1999). The population density of 164
individuals/km2 in our study area is close to those observed
in other secondary forests in Peru (Neville et al., 1976),
Panama (Balwin and Baldwin, 1972) and Colombia
(Thorington, 1968). The small size of the troops can be
explained by the size of the island and the secondary forest
(Baldwin and Baldwin, 1971). Small home ranges have
been reported in other secondary forest areas (Thorington,
1968). The sleeping sites of the four troops were on the
southern coast of the island, which provided protection
against rain and wind. Activity patterns and the vertical
use of space observed in our study troop are comparable to
those of S. oerstediiin secondary forests (Boinski, 1987 and
1999). Group compositions are also similar to those reported
from other study areas (Baldwin and Baldwin, 1981); social
behavior traits, characterized by female coalitions and
female aggression, and males being peripheral to troops
of adult female troops, are typical also of S. boliviensis in
Peru (Mitchell et al., 1991). Interspecific differences in the
social behavior of squirrel monkeys, especially in female


Acknowledgment: The study was supported by the Institut
Pasteur de la Guyane.

Benoit de Thoisy, Association Kwata, BP 672, 97335
Cayenne cedex, French Guiana, France, e-mail:
, Olivier Louguet, Laboratoire
d'Ecologie Gnrn&ale, MNHN/CNRS-UMR 8571, 4
avenue du Petit Chateau, 91800 Brunoy, France, Fran9oise
Bayart, Laboratoire d'Ecologie Gnrn&ale, MNHN/CNRS-
UMR 8571, 4 avenue du Petit Chateau, 91800 Brunoy,
France, and Hugues Contamin, Centre de Primatologie,
Institute Pasteur de Guyane, BP 6010, 97306 Cayenne
cedex, French Guiana, France.

References

Altmann, J. 1974. Observational study of behavior:
Sampling methods. Behaviour 49: 237-265.
Baldwin, J. D. and Baldwin, J. I. 1971. Squirrel monkeys
(Saimiri) in natural habitats in Panama, Colombia, Brazil
and Peru. Primates 12: 45-61.
Baldwin, J. D and Baldwin, J. I. 1972. The ecology and
behavior of squirrel monkeys (Saimiri oerstedii) in a
natural forest in western Panama. Folia Primatol. 18:
161-184.
Baldwin, J. D. and Baldwin, J. I. 1981. The squirrel
monkey, genus Saimiri. In: Ecology and Behavior of
Neotropical Primates, Vol. 1, A. F Coimbra-Filho and R.
A. Mittermeier (eds.), pp.277-330. Academia Brasileira
de Ciencias, Rio de Janeiro.
Boinski, S. 1987. Habitat use by squirrel monkeys (Saimiri
oerstedi) in Costa Rica. Folia Primatol. 49: 151-167.
Boinski, S. 1999. The social organization of squirrel
monkeys: Implications for ecological models of social
evolution. Evol. Anthropol. 8: 101-112.
de Thoisy, B. and Contamin, H. 1998. The squirrel monkey
breeding colony of the Pasteur Institute, Cayenne, French
Guiana. Neotrop. Primates 6: 14-18.
Harris, S., Cresswell, W. J., Forde, P. J., Trewhella, J.,
Woollard, T. and Wray, S. 1990. Home-range analysis
using radio-tracking data: A review of problems and


bonding patterns, can be correlated with the abundance
and distribution of fruit resources, and essentially to food
loc.i.o competition (Boinski, 1999). In the Peruvian study area
L (S. boliviensis) fruit patches harvested were typically large
social and moderately dense, differing from the Surinamese
rest study site of S. sciureus, where smaller and more dispersed
feed patches were exploited (Boinski, 1999). On the Ilet-
la-Mere, fruit patches are also large and dense. Our
preliminary results suggest that, despite genetically-based
SHC variations of social organization in squirrel monkeys
-1 -0,5 0 0,5 1 1,5 (Boinski, 1999), habitat constraints can also modify social
organization, indicating considerable plasticity (Johns
and Skorupa, 1987). Saimiri has rarely been studied in
2. Vertical use of space by squirrel monkeys (Saimiri isolated habitats (Johns and Skorupa, 1987), and this
on the Ilet-la-Mere, French Guiana. UG = undergrowth;
wer canopy level; HC = higher canopy level, island population is of great interest for furthering our
understanding of their behavior, ecology and demographics
in these circumstances.


Figure
sciureus)
LC = lo





Neotropical Primates 10(2), August 2002


techniques particularly as applied to the study of
mammals. Mammal Rev. 20: 97-123.
Janson, C. H. and Boinski, S. 1992. Morphological and
behavioral adaptations for foraging in generalist primates:
The case of the cebines. Am. J. Phys. Anthropol. 88:
483-498.
Johns, A. D. and Skorupa, J. P 1987. Responses of rain-
forest primates to habitat disturbance: A review. Int. J.
Primatol. 8: 157-191.
Mitchell, C. L., Boinski, S. and Van Schaik, C. P 1991.
Competitive regimes and female bonding in two species
of squirrel monkeys (Saimiri oerstedi and S. sciureus).
Behav. Ecol. Sociobiol. 28: 55-60.
Neville, M., Castro, N., Marmol, A. and Revilla, J. 1976.
Censusing primate populations in the reserved area of
the Pacaya and Samiria rivers, Department Loreto, Peru.
Primates 17: 151-181.
Thorington, R. W. 1968. Observations of squirrel monkeys
in a Colombian forest. In: The Squirrel Monkey,
L. A. Rosenblum and R. W. Cooper (eds.), pp.69-85.
Academic Press, London.
White, G. C. and Garrott, R. A. 1990. Analysis of 11 j.
Radio-tracking Data. Academic Press, Harcourt Brace
Jovanovich, San Diego, CA.



ESTUDO PRELIMINARY DO MONITORAMENTO DO
BUGIO RUIvo, ALOUATTA GUARIBA CLAMITANS,
ATRAVES DA CONTAGEM DE BOLOS FECAIS NO
PARQUE ESTADUAL DE ITAPUA, RIO GRANDE DO
SUL, BRASIL

Gerson Buss
Helena P Romanowski
Introdufio

Existe uma necessidade imediata de incremento nos
estudos ecol6gicos e comportamentais de campo, em
busca de dados que nos fornegam uma base concrete para
um melhor conhecimento e que permit o manejo e a
conservacao de nossos primatas (Cullen Jr. e Valladares-
PAdua, 1997). 0 bugio-ruivo, Alouatta guariba, distribui-se
pela mata Atlantica, estendendo-se do sul da Bahia ate a
porcao mais ao sul desse bioma, atingindo Missiones, no
norte da Argentina (Fonseca et al., 1994, Printes et al.,
2001). Trata-se de uma esp&cie ameagada (Brasil, IBAMA,
1989) e a principal causa de seu desaparecimento tem sido
a destruicao do hAbitat (Neville et al., 1988).

A presenca de vestigios pode ser utilizada em estudos
de animals silvestres como indicador de sua ocorrencia
(Romanowski et al., 1998) e uso de hAbitat (Welch et al.,
1990). No caso das fezes, tambdm fornece importantes
informao6es sobre o estado de sadde do animal e dieta
(Prates et al., 1990; Stuart et al., 1998; Santos e Hartz,
2000). Adicionalmente, a detecqao de mudanca no ndmero
de bolos fecais pode ser satisfat6ria para fins de manejo
(Davis e Winstead, 1987; Mitchell et al., 1985; McIntosh


et al., 1995). Destaca-se como uma forma de coleta de
dados que nao provoca distdrbio aos animals, e que pode
ser utilizada corn especies de dificil visualizacao (Palomares
et al., 1991; Soldateli e Blacher, 1996). Apesar dos primatas
serem considerados animals de ficil visualizacao, pelo
habito diurno da grande maioria das esp&cies, certas
esp&cies sao mais dificeis de serem encontradas devido ao
seu comportamento. Esse d o caso de Alouatta, pois apesar
do ronco que facility a localizacao do grupo, sao animals
dificeis de serem localizados devido a existencia de perfodos
prolongados de inatividade diaria, comportamento tfpico
desse genero de primatas (Mendes, 1985; Marques, 1989;
Fortes, 1999).

No Parque Estadual de Itapua, o monitoramento, atraves
da contagem de bolos fecais e facilitado por esta ser a
dnica especie de primata nao-humano present na area e
devido a geomorfologia de Itapua, em geral, e do Morro
do Campista, em particular, que apresenta grande ndmero
de afloramentos graniticos no interior da mata, facilitando
a localizacao visual dos mesmos. Aldm disso, o aspect e o
odor caracteristico do bolo fecal do bugio-ruivo sao bastante
peculiares, o que facility sua identificaqao e localizacao.

O objetivo desse trabalho e verificar a eficacia da contagem
de bolos fecais para monitoramento das populao6es de
bugio-ruivo, Alouatta guariba clamitans Cabrera, 1940 no
Parque Estadual de Itapua, Viamro, Rio Grande do Sul,
bem como, trazer informao6es relatives ao uso do habitat.
Este trabalho e parte integrante do "Programa Macacos
Urbanos para Pesquisa e Conservacao do Bugio-ruivo (A. g.
clamitans) no Rio Grande do Sul".

M6todos

Area de Estudo
O Morro do Campista (30o23'S, 51o02'W), tambdm
conhecido como Ponta de Itapui, localiza-se no Parque
Estadual de Itapua, Viamao, Rio Grande do Sul (Fig. 1).
O "Campista" caracteriza-se como um complex orogenico,
granitico, cujo cume principal possui 182 m de altura, e
apresenta uma area aproximada de 300 ha. Estao presents
afloramentos rochosos no topo, enquanto suas encostas
encontram-se praticamente todas cobertas por mata.
0 clima local se classifica como Cfa pelo sistema de Kbppen,
descrito como subtropical 6mido, com mddia do m&s mais
quente superior a 220C (janeiro), mddia do mes mais frio
entire -3 e 180C (julho), sendo a temperature mddia annual
de 17,5C. A precipitagao mddia annual situa-se em torno de
1.300 mm (Brasil, Rio Grande do Sul, 1997).

A classificaqao fision6mica da vegetacao das unidades
amostrais foi realizada utilizando-se a classificacao proposta
por Brack et al. (1998), sendo a que segue:

(A) Mata higr6fila formaqao florestal que ocorre nos
funds dos vales e encosta sul dos morros, constituindo-
se algumas vezes em comunidades relictuais com forte
influencia da Floresta Pluvial Tropical Atlantica (Floresta





Neotropical Primates 10(2), August 2002


SUA




OCEANO


I Momao do Camp.sta


Legenda:
Ef Dunas
I11111 Campo
E-I Mata native
STrilhas

1.000 m
N
t


Figura 1. Localizacao do Morro do Campista (3023'S, 5102'W),
Parque Estadual de Itapua (1), Viamao, Rio Grande do Sul, Brasil,
apresentando as areas de mata e a rede de trilhas.


Ombr6fila Densa). As condi6ces de relevo, que permitem
uma maior umidade relative do ar, a maior profundidade
dos solos e a maior capacidade de armazenamento de Agua,
proporcionam condig6es mais seletivas para o crescimento
de uma vegetagao de grande porte e maior riqueza floristica
que as demais comunidades florestais. A mata higr6fila
contem especies que se destacam pela ampla superficie foliar
(latifoliadas). Em relagao a estrutura da floresta, verifica-se a
presenga de tries ou quatro estratos arb6reos.

(B) Mata mesohigr6fila constitufda por uma comunidade
florestal que ocupa a porgao mrdia ou baixa dos morros,
ou mesmo em terrenos mais ou menos pianos, onde as
condigoes ambientais nao sejam extremadas. Seus elements
florestais nao apresentam grande seletividade e term ampla
distribuigao no Estado, estando presents ainda na maior
parte das matas secundarias do municipio. A altura da mata
e de 10 a 15 m, sendo encontrados 2 a 3 estratos arb6reos.

(C) Mata subxer6fila matas baixas ou capes encontrados
nos topos ou encostas superiores dos morros, onde o solo
e muitas vezes raso (litossolo), sendo sua textura grosseira
corn feigoes pr6prias de solos corn baixa retencao hfdrica.
Com respeito ao mesoclima, estes locals de topo de morro
estao sujeitos a maior exposigao solar e ventos mais
intensos. A denominacao de mata subxer6fila e adotada
para caracterizar este tipo de vegetagao de ambientes
mais secos, onde morfologicamente a vegetagao tambem
evidencia tend&ncia de redugao da superficie foliar e
escleromorfismo. A altura mddia do dossel e de 6 a 12 m. A
estratificacao e mais simplificada do que a mata higr6fila,
corn presenga de 2 ou 3 andares arb6reos. Pode ocorrer


algumas vezes um estrato de individuos emergentes,
chegando a alcangar 15 m.

(D) Mata psam6fila tambem conhecida como mata
de resting, sendo uma mata caracteristica de terrenos
arenosos (paleodunas) entremeadas por banhados e outras
areas 6midas correspondents as margens de antigas
transgresses e regresses do Lago Guafba e Laguna dos
Patos. Tern uma altura que varia de 6 a 10 m, sendo que
as especies emergentes podem chegar a 15 m. Evidencia-se
alguma tend&ncia xerom6rfica nas folhas de muitas espucies
atraves da consistencia coriacea, do reduzido tamanho e
superficie lustrosa. Possui muitos elements floristicos que
sao comuns is matas subxer6filas.

Amostragem
A presenga de bolos fecais frescos de bugio-ruivo foi
registrada em trilhas previamente demarcadas no Morro
do Campista, Parque Estadual de Itapua videe Fig. 1). Um
bolo fecal foi definido como uma ou mais pelotas de fezes
agrupados, num raio de aproximadamente um metro, e
que estivessem sobre a trilha. As trilhas podem ser um local
preferencial para defecacao, considerando que sao livres de
vegetagao de sub-bosque (Gilbert, 1997).

Considerou-se cada trecho de 50 m de comprimento de
trilha como uma unidade amostral. Em cada unidade
amostral foi identificada a formanao florestal predominant.
Nos 5.350 m. de rede de trilhas, foram demarcadas
107 unidades amostrais, sendo 41 de mata mesohigr6fila,
37 de mata higr6fila, 18 de subxer6fila e 10 de psam6fila.
Os registros foram realizados corn um intervalo mfnimo de
5 dias. Em cada registro anotou-se o horario e a unidade
amostral em que foram encontrados. Em uma unidade
amostral era registrado no mAximo um bolo fecal por dia de
amostragem. Foram realizados 13 dias de amostragem, no
perfodo de novembro de 1999 r maio de 2000, totalizando
1.391 unidades amostrais vistoriadas em aproximadamente
69,5 km de trilhas percorridas.

Os dados foram analisados no program SPSS for Windows.
Para verificar a relagao entire as distintas formao6es florestais
e a presenga de bolos fecais foi utilizado o teste de associadao
atraves da analise de "maximum likelihood" (Sokal e Rohlf,
1981).

Resultados e Discussio

Em treze dias de amostragem foram registrados 48 bolos
fecais frescos em 34 unidades amostrais; destes, 24 (50%)
em mata higr6fila, 21 (43%) em mata mesohigr6fila e
3 (6,2%) em subxer6fila. Nao houve registro na mata
psam6fila (Fig. 2).

A presenga de bolos fecais de A. g. clamitans nas unidades
amostrais foi significativamente associada corn a formadao
florestal (G = 12,40; gl = 3; P = 0,006) (Tabela 1). Ao
considerarmos as visualizagoes de bugio-ruivo por unidade
amostral, de acordo corn os dados do censo realizado





Neotropical Primates 10(2), August 2002


Tabela 1. Ndmero de unidades amostrais por formayao florestal relacionado corn o registro de bolos fecais de Alouatta guariba clamitans,
Morro do Campista, Parque Estadual de Itapua (3023'S, 5102'W), Viamao, RS, Brasil, entire novembro de 1999 e maio de 2000.

Formacao Florestal
Bolos fecais Higr6fila Mesohigr6fila Subxer6fila Psam6fila Total
Presenga 15 16 3 0 34
Ausencia 22 25 16 10 73
Total 37 41 19 10 107


Tabela 2. Unidades amostrais com visualizacao de bugio-ruivo (A. g. clamitans) em relagao ao ndmero total de unidades amostrais nas
formagoes florestais do Morro do Campista, Parque Estadual de Itapua (3023'S, 502'W), Viamao, RS, Brasil, entire novembro de 1999
e maio de 2000 (adaptado de Buss, 2001).
Formacao Florestal
Visualizagao Higr6fila Mesohigr6fila Subxer6fila Psam6fila Total
Ausencia 19 26 17 10 72
Presenca 18 15 2 0 35
Total 37 41 19 10 107


por Buss (2001), estas tambem estao significativamente
associadas corn a formaqao florestal (G = 17,36; gl = 3; P =
0,0005) (Tabela 2).

Esses resultados indicam que a presenga de bolos fecais
mostrou ser um bom indicador de ocorrencia, podendo
fornecer importantes informagoes relatives ao uso do
habitat. Indicam tambem, um uso diferenciado das
formag6es florestais presents no Morro do Campista.
Considerando que essas formacoes apresentam diferengas
na estrutura e composigao de especies arb6reas (Brack
et al., 1998), bern como, nos aspects relacionados
a fenologia das especies arb6reas, sup6e-se que essas
caracteristicas resulted em diferengas no uso do habitat
pelo bugio-ruivo.

Este trabalho foi desenvolvido dentro de um estudo mais
abrangente sobre densidade e caracterizagao do habitat do
bugio-ruivo no Parque Estadual de Itapui, cujos resultados
estao sendo preparados para publicacao.

0 Parque Estadual de Itapua esteve fechado a visitagao
p6blica de 1990 a 2002. Dentro desse context, o


30-


| 20-
o15-
s-
10.
c 5.


Higr6fila Mesohigr6fila Subxertfila Psam6fila
formacao florestal


Figura 2. Ndmero de bolos fecais encontrados nas formagoes
florestais do Morro do Campista, Parque Estadual de Itapua
(30023'S;51002'W), Viamao, RS, Brasil, entire novembro de
1999 e maio de 2000.


monitoramento pela contagem de bolos fecais, devido
a sua facilidade de implementagao, pode colaborar no
control da situacao populacional do bugio-ruivo no
Parque, contribuindo, portanto, na avaliacao do impact
da visitagao sobre essas populagoes. Aldm disso, associado
com outros procedimentos, como por exemplo, a analise
de parasitas presents nas fezes, pode trazer valiosas
informagoes sobre a sadde dessas populagoes (Stuart et al.,
1998), e direcionar ages de manejo (Davis e Winstead,
1987) visando a conservagao dessa esp&cie.

Agradecimentos

A administracao do Parque Estadual de Itapua pela
autorizagao para realizacao do trabalho. Ao Prof. S&rgio
L. C. Leite (Departamento de BotInica, Universidade
Federal do Rio Grande do Sul) pelo apoio na caracterizacao
das formagoes florestais. Mauricio Peroni auxfliou nas
atividades de campo. Aos companheiros da Comissdo
de Luta pela Efetivagao do Parque Estadual de Itapua
(CLEPEI) pelo auxflio na manutencao da base de campo. A
Solange M. Kerpel pelos comentArios ao artigo e a CAPES
pelo suporte financeiro.

Gerson Buss, Programa de P6s-Graduacao em Ecologia,
Universidade Federal do Rio Grande do Sul (UFRGS),
Avenida Bento Gongalves 9500, 91540-000 Porto Alegre,
Rio Grande do Sul, Brasil, e-mail:
e Helena P. Romanowski, Departamento de Zoologia,
Universidade Federal do Rio Grande do Sul (UFRGS),
Avenida Bento Gongalves 9500, 91540-000 Porto Alegre,
Rio Grande do Sul, Brasil, e-mail: s.br>.

Referencias

Buss, G. 2001. Estudo da densidade populacional do
bugio-ruivo Alouatta guariba clamitans (Cabrera, 1940)
(Primates, Atelidae) nas formao6es florestais do Morro
do Campista, Parque Estadual de Itapua, Viamao, RS.





Neotropical Primates 10(2), August 2002


Dissertacao de Mestrado, Universidade Federal do Rio
Grande do Sul, Porto Alegre.
Brack, P., Rodrigues, R. S., Sobral, M. e Leite, S. L. C.
1998. Arvores e arbustos na vegetacao natural de Porto
Alegre, Rio Grande do Sul, Brasil. Iheringia, Ser. Bot. 51:
139-166.
Brasil, IBAMA. 1989. Portaria 1.522 de 19 de dezembro
de 1989. Didrio Oficial da Unido 22/12/89. Brasilia,
DF, pp.24156-24159. Institute Brasileiro do Meio
Ambiente e dos Recursos Naturais Renoviveis (IBAMA),
Brasilia.
Brasil, Rio Grande do Sul. 1997. Plano de Manejo: Parque
Estadual de Itapud. Departamento de Recursos Naturais
Renoviveis, SAA, Porto Alegre. 158 pp.
Cullen Jr., L. e Valladares-Padua, C. 1997. Metodos para
estudos de ecologia, manejo e conservacao de primatas na
natureza. Em: Manejo e Conservaado da Vida Silvestre no
Brasil, C. Valladares-Padua e R. Bodmer (eds.), pp.239-
269. MCT-CNPq e Sociedade Civil Mamiraud, Brasilia e
Tef6, Amazonas.
Davis, D. E. e Winstead, R. L. 1987. Estimaci6n de tama-
fos de poblaciones animals. In: Manual de Tecnicas de
Gestidn de Vida Silvestre, R. R. Tarres (ed.), pp.233-258.
The Wildlife Society Inc., Maryland.
Fonseca, G. A. B. da, Rylands, A. B., Costa, C. M. R.,
Machado, R. B. e Leite, Y. L. 1994. Livro L' !I'C/It dos
Mamiferos Brasileiros Ameacados de Extinado. Fundadao
Biodiversitas, Belo Horizonte.
Fortes, V. B. 1999. Dieta, atividades e uso do espago por
Alouatta fusca clamitans (Cabrera, 1940) (Primates:
Cebidae) na Depressao Central do Rio Grande do Sul.
Dissertacao de Mestrado, Universidade Federal do Rio
Grande do Sul, Porto Alegre.
Gilbert, K. A. 1997. Red howling monkey use of specific
defecation sites as a parasite avoidance strategy. Anim.
Behav. 54: 451-455.
McIntosh, R., Burlton, E W. E. e McReddie, G. 1995.
Monitoring the density of a roe deer Capreolus capreolus
population subjected to heavy hunting pressure. Forestry
Ecology and Management 79(1995): 99-106.
Marques, A. A. B. 1996. 0 bugio-ruivo Alouatta fusca
clamitans (Cabrera, 1940) (Primates, Cebidae) na
Estagao Ecol6gica de Aracuri, RS: Variaqoes sazonais
de forrageamento. Dissertacao de Mestrado. Pontificia
Universidade Cat61lica, Porto Alegre.
Mendes, S.L. 1985. Uso do espaco, padres de atividades
didrias e organizacao social de Alouatta fusca (Primates,
Cebidae) em Caratinga, MG. Dissertagao de Mestrado.
Universidade de Brasilia, Brasilia.
Mitchell, B., Rowe, J. J., Ratcliffe, P. e Hinge, M. 1985.
Defecation frequency in roe deer (Capreolus capreolus)
in relation to the accumulation rates of faecal deposits.
J. Zool., Lond. (A) 207: 1-7.
Neville, M. K., Glander, K. E., Braza, F. e Rylands, A. B.
1988. The howling monkeys, genus Alouatta. In: Ecol-
ogy and Behavior of Neotropical Primates. Vol. 2, R. A.
Mittermeier, A. B. Rylands, A. E Coimbra-Filho e G. A.
B. da Fonseca (eds.), pp.349-453. World Wildlife Fund,
Washington, DC.


Palomares, E, Rodriguez, A., Lafitte, R. e Delibes, M. 1991.
The status and distribution of the Iberian lynx Felis par-
dina (Temminck) in Coto Donana Area, SW Spain. Biol.
Conserv. 57: 159-169.
Prates, J. C., Gayer, S. M. P., Kunz Jr., L. F e Buss, G. 1990.
Feeding habits of the brown howler monkey Alouatta
fusca clamitans (Cabrera, 1940) (Cebidae, Alouattinae)
in the Itapua State Park: A preliminary report. Acta Biol.
Leopoldensia 12: 175-188.
Printes, R. C., Liesenfeld, M. V. A. and Jerusalinsky,
L. 2001. Alouatta guariba clamitans (Cabrera, 1940):
A new southern limit for the species and for Neotropical
primates. Neotrop. Primates 9: 118-121.
Romanowski, H. P., Dornelles, S. S., Buss, G., Brutto,
L. F G., Jardim, M. M. A., Printes, R. C. e Fialho, M. S.
1998. 0 bugio-ruivo Alouattafusca. In: Atlas Ambientalde
Porto Alegre, R. Menegat et al. (eds.), pp.63-64. Editora
Universidade, Prefeitura Municipal de Porto Alegre.
Santos, M. F M. e Hartz, S. M. 1999. The food habits
of Procyon cancrivorus (Carnivora, Procyonidae) in the
Lami Biological Reserve, Porto Alegre, southern Brazil.
Mammalia 63: 525-530.
Sokal, R. R. e Rohlf, F J. 1981. Biometry. 2nd ed. W. H.
Freeman and Company, New York.
Soldateli, M. e Blacher, C. 1996. Considerao6es preliminares
sobre o ndmero e distribuicao espaco/temporal de sinais de
Lutra longicaudis (Olfers, 1818) (Carnivora, Mustelidae)
nas lagoas da Conceicao e do Peri, Ilha de Santa Catarina,
SC, Brasil. Biotemas 9: 38-64.
Stuart, M., Pendergast, V., Rumfelt, S., Pierberg, S.,
Greenspan, L., Glander, K. e Clarke, M. 1998. Parasites
of wild howlers (Alouatta spp.). Int. J. Primatol. 19(3):
493-512.
Welch, D., Staines, B. W., Catt, D. C. e Scott, D. 1990.
Habitat usage by red (Cervus elaphus) and roe (Capreolus
capreolus) deer in a Scottish Sitka spruce plantation.
J. Zool., Lond. 221: 453-476.



BDGEOPRIM DATABASE OF GEO-REFERENCED
LOCALITIES OF NEOTROPICAL PRIMATES


Introduction


Andre Hirsch, Luiz Gustavo Dias
Livia de Oliveira Martins,
Renata Ferreira Campos
Elena Charlotte Landau,
Natdlia Almeida Teixeira Resende


One of the main problems for the conservation of the
Neotropical primates is that our understanding of their
geographical distributions is still poor. This is underlined
by the fact that many new forms are still being discovered:
31 species and subspecies since 1960 (three from the Atlan-
tic forest and the remainder from Amazonia), 13 of them
since 1990 (Rylands et al., 2001). Many of the Neotropical
primates are now threatened (Rylands et al., 1995, 1997),
and a database documenting their past and present distri-





Neotropical Primates 10(2), August 2002


butions is a vital tool for an understanding of their con-
servation status (degree of reduction in their range), for
priority-setting, and for planning conservation strategies,
allowing for information to be made available in a timely
fashion to field researchers, conservation organizations and
government institutions. In more dire situations, it is es-
sential to have access to the maximum information on the
past and present ranges of critically endangered species for
their management and the translocation of populations
from areas suffering strong human impacts to locations
where they can be guaranteed greater protection (see for ex-
ample, Garcia-Ordufia et al., 1987; Kierulff and Proc6pio
de Oliveira, 1994). Only recently was it possible to identify
the extent of the historic occurrence of the red-handed
howling monkey (Alouatta belzebul) in the north-east of
Brazil through some very few and obscure locality records;
a species now largely extinct in the region (Bonvicino et
al., 1984; Coimbra-Filho et al., 1995). The need for an
understanding of historic and recent distributions for the
conservation of primates in regions where forests have been
largely destroyed also became evident during surveys in the
Rio Doce basin in the state of Minas Gerais (the "Steel
Valley"), as well as when drawing up conservation priorities
and strategies for the Atlantic forest in the south of the state
of Bahia, Brazil (CI and IESB, 1997; Hirsch, in prep.).

The goal of the BDGEOPRIM, the Database of Geo-
referenced Localities for Neotropical Primates, is to organize
the scattered information available in gazetteers, in the
scientific literature (much of it grey), and from field studies,
of the locality records of all Neotropical primate species,
and to make it available for use in libraries and museums,
and by professionals in primatology, conservation,
biogeography, and taxonomy.

Although we have checked the entire database three times,
the users will undoubtedly find errors, and we would be
very grateful for comments and suggestions, as well as leads
regarding sources of information that we have missed. In
some parts of the Database and in the maps, the foreign
users will find some terms in Portuguese because we began
tabulation of the information in Brazil's native language. As
a next step, we will make the BDGEOPRIM available in
Portuguese, Spanish and English.

This database has not yet been published, but we
decided to put it on an Internet homepage (see //www.icb.ufmg.br/-primatas/home_bdgeoprim.htm>),
because there have been so many enquiries and requests
for information and analyses from numerous people and
institutions from Brazil (Rio Grande do Sul, Minas Gerais,
Paraiba, Conservation International, the Brazilian Institute
for the Environment IBAMA) and other countries such as
Argentina, Paraguay.

Our initial objectives were to a) tabulate all the localities
for Neotropical primates listed in the current literature;
b) arrange the information in a database format; c) geo-
reference all the tabulated localities; d) check the veracity


of the information by crossing all the data with maps
of primate geographical distributions, hydrography,
topography, vegetation biomess and ecosystems),
conservation units, and political divisions, and e) plot the
records by taxa on maps generated through a Geographic
Information System (GIS).

Methods

The taxonomy of Neotropical primates is still far from
definitive. Most especially the application of molecular
genetics and cytotaxonomy, along with the findings of
new species and subspecies, over the last two decades,
has resulted in numerous revisions and a far better
understanding of the true diversity of the Platyrrhini,
based increasingly on the Phylogenetic Species Concept
(see Groves, 2001). A number of genera (for example the
woolly monkeys, Lagothrix) and groups of species (for
example, the red howling monkeys, Alouatta seniculus)
are in need of a modern revision of their component
taxa, while some particularly tricky genera are still subject
to dispute (for example, the capuchin monkeys, Cebus,
the night monkeys, Aotus, and the squirrel monkeys,
Saimiri). For BDGEOPRIM, we adopted the taxonomic
arrangement proposed by Rylands et al. (2000).

To start, we made an extensive bibliographical review to
check the information already available. We first tabulated
all the records listed in the published gazetteers of such as
Hershkovitz (1977), Kinzey (1982), Torres de Assumpcao
(1983) and Oliver and Santos (1991). We then searched for
more recent scientific papers, including all those published
in Primate Conservation and Neotropical Primates. We
also checked some classic works from the 18th century,
such as Wied-Neuwied (1821) and von Spix and von
Martius (1981). To this, we added unpublished records of
primate localities from museum collections and from the
field, obtained by the authors of BDGEOPRIM and by
a number of researchers who spontaneously contributed
information from surveys.

The information associated with each record (locality) was
standardized, even if it was a type locality. This was necessary
for two reasons: 1) to sort and classify all the records in
alphabetical order, and 2) to avoid duplicating records from
the same place and/or the same taxon. When we confirmed
that information was erroneous, we assigned the correct data
and stored the original information in the "Observations"
field. When the information for a specific field was lacking
(a "missing value"), we completed it, when possible,
with the correct information. To give an example of the
standardization of references for the same "locality", "right
bank of Amazon River, Santar6m, PA, Brazil", "Amazon
River, right bank, Santarem, Brazil", and "Santarem, right
bank, Amazon River" were all recorded as:

Locality Municipality State Country


Amazon River, right bank Santarem


PA Brazil





Neotropical Primates 10(2), August 2002


Many abbreviations were used geographic names, Brazil-
ian states, categories of protected areas, IUCN categories of
threatened status, museum acronyms, and so on. For easy
identification, we drew up specific tables for each set.

All the information associated with each record (locality)
was tabulated in a Database with 58 fields (see Table 1).
In this way, it is possible to consult the Database using dif-
ferent combinations of information fields, and the output
report can be viewed either as a simple table (list) or as a
more complex matrix, crossing the fields one by one, or one
for several fields.

Almost all the geopolitical data which was not available in
the original scientific papers consulted, we obtained from
official publications, multimedia products and online
services released by government agencies, non-governmental
organizations and other institutions, including:


IBGE (),
SURAPA CD-Rom ( may99/msg00799.html>),
ESRI ArcData Online ( html>),
Expedia.com Maps Online ( pub/>),
USGS (),
UNEP/GRID (),
Garmin MapSource World Atlas ( cartography/>),
GEOMinas (), and other
printed world atlases.

After tabulating all the records, we linked the Database with
a Geographic Information System, using three information
fields: the record identification number (N_ID) and the
geographic coordinates (longitude and latitude). Geo-
referenced, it is possible to plot any field of information


Table 1. Database information fields and abbreviations associated with the locality records. Y = yes, N= no.


Field
1. Record Identification No. (# primary key)
2. Date
3. Operator
4. Family
5. Genus
6. Species
7. Subspecies
8. Description (Author)
9. Description (Year)
10. Common Name
11. Type Locality (Y or N)
12. Survey Area of Hirsch Ph.D. Thesis (Y or N)
13. IUCN (1996) Category
14. Present Population Status and Risk of Threat
15. Biome
16. Ecosystem or Habitat Type
17. Conservation Unit Category
18. Locality
19. Municipality or "City"
20. State, "Departamento" or "Provincia"
21. Country
22. Geog. Coord. (Latitude, dd)
23. Geog. Coord. (Latitude, mm)
24. Geog. Coord. (Latitude, ss)
25. Geog. Coord. (Longitude, ddd)
26. Geog. Coord. (Longitude, mm)
27. Geog. Coord. (Longitude, ss)
28. Geog. Coord. (Longitude, decimal format)
29. Geog. Coord. (Latitude, decimal format)
30. Altitude Minimum (m)
31. Altitude Maximum (m)


Abbreviation
N_ID
DATE
OPERATOR
FAMILY
GENUS
SPECIES
SUBSPECIES
DESCRIPTION
YEAR
COMMON_NAM
TYPE_LOCAL
THESIS_FRA
IUCN_1996
POPUL_STAT
BIOME
ECOSYSTEM
CU
LOCALITY
MUNICIPAL
STATE
COUNTRY
LAT_DD
LAT_MM
LAT_SS
LONG_DDD
LONG_MM
LONG_SS
LONGITUDE
LATITUDE
ALT_MIN
ALT MAX


32. Altitude Average (m)
33. Area (ha)
34. Year of Creation (ha), if it was a CU
35. Administration
36. Reference
37. Type of Record
38. Collector
39. Year of Collection
40. Museum
41. Number of Museum Collection
42. Original Record Number from Gazetteer
43. Change or Attributed of Genus (Y or N)
44. Change or Attributed of Species (Y or N))
45. Change or Attributed of Subspecies (Y or N)
46. Change or Attributed of Cons. Unit (Y or N)
47. Change or Attributed of Locality (Y or N)
48. Change or Attributed of Municipality (Y or N)
49. Change or Attributed of State (Y or N)
50. Change or Attributed of Country (Y or N)
51. Change or Attributed of Altitude (Y or N)
52. Change or Attributed of Area (Y or N)
53. Change or Attributed of GCs (Y or N)
54. Change or Attributed of GCs with ArcGIS
(Yor N)
55. Change or Attributed of GCs with Garmin
(Y or N)
56. Change or Attributed of GCs with Expedia
(Yor N)
57. Change or Attributed of GCs with SURAPA
(Yor N)
58. Observations


Abbreviation
ALT_AVG
AREA
YEAR_CREAT
ADMINISTRA
REFERENCE
TYPE_REC
COLLECTOR
COL_YEAR
MUSEUM
COL_ NUM
N_ORIG
ATB_GENUS
ATB_SP
ATB_SSP
ATBUC
ATB_LOCAL
ATB_MUNIC
ATB_STATE
ATB_COUNTR
ATB_ALT
ATB_AREA
ATB_CGS
ATB_ARCGIS

ATB_GARMIN

ATB_EXPED

ATB_SURAPA

OBS





Neotropical Primates 10(2), August 2002


on a projected map, showing the records in their actual
geographic position. Unfortunately, it was not possible
to locate 39 records that had no geographic coordinates,
so that we were unable to find their exact location. These
records were stored in the Database but are not visible on
the maps.

Because the distributions of the Neotropical primates
embrace South America (southern hemisphere) and
Central America (northern hemisphere), we used a World
Geographic Coordinate System and WGS84 Datum
(World Geographic System 1984). As such, we avoided
some problems with displacement and data matching, and
facilitated the combination of "overlays" from different sets
of data.

All the maps were generated using ArcGIS v. 8.1 (ESRI,
2001). Initially, we produced maps for all 18 genera,
showing the records (points) only for the species. The next
step will involve the production of species maps which
show the records for each subspecies.

At this stage, we decided not to trace lines delimiting the
distributions of the different species in each genus. This
is because for some the limits remain unclear, this kind of
delimitation is laborious and is, besides, often guesswork,
using inferences from natural boundaries, such as rivers,
mountain ridges, and vegetation types and, an often
inadequate, knowledge of historic changes in vegetation.


Summary Results

At the present stage, the BDGEOPRIM consists of 5,631
locality records, embracing all of the 18 Neotropical primate
genera, 110 species and 205 subspecies in 21 countries from
Central and South America (see Fig. 1).

* A total of 487 bibliographical references were reviewed,
naturally including the classic works (gazetteer) of
Hershkovitz (1977) with 807 records, Kinzey (1982)
with 679 records, and Oliver and Santos (1991) with
516 records. A further 45 references provided more than
50 records. Besides Hershkovitz' (1977) gazetteer for
callitrichids, a further 655 records were cited for the first
time and 472 records are exclusive citations.
* The map of localities, recorded in a 25 x 25 km grid,
shows that they are not uniformly distributed. The
highest density is concentrated in one continuous area
in the southeast region of Brazil, in the Atlantic forest.
In Amazonia, locality records are highly clumped,
distributed along the major rivers. The Cerrado has a
uniformly low density of records, while in Mesoamerica
the records show a patchy distribution as in Amazonia (see
Fig. 1).
* The genera with the highest numbers of records are:
1,166 for the howling monkeys (Alouatta); 894 for the
capuchin monkeys (Cebus); 665 for the marmosets (Mico
and ( .-1' ), 616 for tamarins (Saguinus); and 545 for
the titi monkeys (Callicebus).


Legend:
Records Density per Grid
High: 87
Low: 0
Hydrography Rivers
Hydrography Lakes
i[I Country Boundaries
BDGEOPRIM
Database of Georreferenced
Occurrence Localities of
Neotropical Primates
Cartographic Base:
IBGE (1998), ESRI (2001) and
GEOMINAS (2001)
Geographic Information System:
ArcGIS 8.1 (ESRI, 2001)
Geoprocessment:
UF/v\G Andr6 Hirsch
Primate Sector "
Department of Zoology
Biological Science Institut
Federal University of Minas Gerais
Belo Horizonte. August 2002.
Projection' Transverse of Mercator
Datum. WGS84
Scale: 1:37,500.000
0 500 1.000 1 .


Loalization
Map ,


iaorw w y ow


Figure 1. Distribution of Neotropical primate locality records. Density per 25 x 25 km grid.





Neotropical Primates 10(2), August 2002


* All the Central and Mesoamerican countries with
primates are included. Those with the highest numbers
of records are: Brazil with 3,680; Bolivia with 431;
Venezuela with 379; Peru with 299; and Colombia with
227.
* Regarding threatened species, 304 records are of Critically
Endangered (CR) primates, 632 records of Endangered
(EN), 1,078 of Vulnerable (VU), 2,922 records are of the
Low Risk (LR) category, and 20 records are from those
classified as Data Deficient (DD).
* Considering only the Brazilian biomes, 2,429 records
are from the Amazon, 1,843 from the Atlantic forest, 367
from the Cerrado, 84 from the Caatinga and 23 from the
Pantanal Matogrossense.
* A total of 1,746 records (31%) are from protected areas,
the majority National Parks, according to the base maps
provided by SURAPA (1999).
* Records from museum collections are not well-represented
in the database. A more comprehensive survey of the key
museums has still to be done. At this time, 1,003 records
are from museum specimens, representing 17.8% of the
total records.

Future Products

We hope that the BDGEOPRIM will be released in three
different languages (Portuguese, Spanish and English) over
the next year, as a CD-ROM, and/or in a form which will
allow for on-line interactive access, structured in such a
way that information stored in the Database will be easily
and quickly available. The BDGEOPRIM will eventually
include biological and ecological data on the Neotropical
primate species, with a picture of each.

A Dedication

The database is dedicated to Philip Hershkovitz 1909-
1997 (in memorial), Emeritus Curator of Mammals at the
Field Museum of Natural History, Chicago, and one of the
world's most distinguished mammalogists and prominent
primatologists of the Neotropical region. Over 50 years,
he described 75 new species and subspecies, and published
more than 160 scientific papers and 100 non-technical
publications. His book, Living New World Monkeys (1977),
along with numerous accompanying papers, put our
knowledge of platyrrhine systematics and distributions
years ahead of other primate groups.

Correct Reference Citation

Hirsch, A., Dias, L. G., Martins, L. de 0., Campos,
R. F, Landau, E. C. and Resende, N. A. T. In prep.
BDGEOPRIM Database of Geo-referenced Localities
for Neotropical Primates. Departamento de Zoologia,
Universidade Federal de Minas Gerais (UFMG),
Belo Horizonte. Web site: -primatas/home_bdgeoprim.htm>.


Acknowledgments

We are very grateful to the Margot Marsh Biodiversity
Foundation (Grant CP FY02/007) for financial support;
to Luiz Paulo de S. Pinto and Carlos A. Bouchardet from
Conservation International do Brasil, for their expertise
and help with the financial administration of the Database;
to the Universidade Federal de Minas Gerais (UFMG) for
logistical support; to Felipe B. C. de Sousa and Elizangela
M. dos Santos for their help with data tabulation; to
German A. Bohorquez Mahecha, Federal University of
Minas Gerais, for his comments and help with Colombian
administrative divisions and localities; to Anthony B.
Rylands and Thomas Brooks from the Center for Applied
Biodiversity Science (CABS) at Conservation International
(CI), for their incentive, comments and suggestions; to
Roberto Cavalcanti from CI-Brasil for his suggestions and
comments on the homepage; to Claudia M. Jacobi for
her patience in revising the English version; to Joao P. R.
Silva, Webmaster from ICBNet/UFMG, for his help with
the homepage management, and to numerous researchers
and colleagues who spontaneously contributed recent field
records of the occurrence of primates.

Andr6 Hirsch (Coordinator), Luiz Gustavo Dias, Livia
de Oliveira Martins, Renata Ferreira Campos, Setor de
Primatas, Elena Charlotte Landau, and Natalia Almeida
Teixeira Resende, Setor de Geoprocessamento, Laborat6rio
de Mastozoologia, Departamento de Zoologia, Instituto de
Ciencias Biol6gicas, Universidade Federal de Minas Gerais,
31270-901 Belo Horizonte, Minas Gerais, Brasil. E-mail:
.

References

Bonvicino, C. R., Langguth, A. and Mittermeier, R. A.
1984. A study of pelage color and geographic distribution
in Alouatta belzebul (Primates: Cebidae). Rev. Nordestina
Biol. 6(2): 139-148.
CI and IESB. 1997. The Economics of Biodiversity Conserva-
tion in the Brazilian Atlantic Forest/A Economia da Con-
servaado na Mata Atldntica Brasileira. CI Project Profile
1997. Conservation International (CI), Washington, DC,
Institute de Estudos Socio-Ambientais do Sul da Bahia
(IESB), Ilheus, Bahia, Brazil. 12pp.
Coimbra-Filho, A. E, Camara, I. de G. and Rylands, A. B.
1995. On the geographic distribution of the red-handed
howling monkey, Alouatta belzebul, in North-east Brazil.
Neotrop. Primates 3(4): 176-179.
ESRI. 2001. ArcView GIS v. 8.1. Environmental Systems
Research Institute (ESRI), Redlands, CA.
Garcia-Ordufia, E, Canales-Espinosa, D., Rodriguez-
Luna, E., Silva-Lopez, G., Jimenez-Huerta, J.,
Benitez-Rodrfguez, J. and Hermida-Lagunes, J. 1987.
Translocation program for the howler monkey (Alouatta
palliata): A report. Am. J. Primatol. 12: 363-364.
Groves, C. P. 2001. Primate Taxonomy. Smithsonian Institu-
tion Press, Washington, DC.





Neotropical Primates 10(2), August 2002


Hershkovitz, P 1977. Living New World Monkeys (PI'.ir, -
rhini) With an Introduction to Primates. Vol. 1. The Uni-
versity of Chicago Press, Chicago. 1117pp.
Hirsch, A. In prep. Fragmentagao do habitat e estrategias
de conservagao de primatas na bacia do Rio Doce, Minas
Gerais, utilizando um Sistema de Informacao Geografica.
Doctoral thesis, Universidade Federal de Minas Gerais
(UFMG), Belo Horizonte.
Kierulff, M. C. M. and Proc6pio de Oliveira, P. 1994.
Habitat preservation and the translocation of threatened
groups of golden lion tamarins, Leontopithecus rosalia.
Neotrop. Primates 3(suppl): 15-18.
Kinzey, W. G. 1982. Distribution of some Neotropical
primates and the model of Pleistocene forest refugia. In:
Biological Diversification in the Tropics, G. T. Prance (ed.),
pp.455-482 + gazetteer of 53pp. Columbia University
Press, New York.
Oliver, W. L. R. and Santos, I. B. 1991. Threatened en-
demic mammals of the Atlantic forest region of south-
east Brazil. Jersey -- Preserv. Trust, Special Sci. Rep. (4):
1-126.
Rylands, A .B., Mittermeier, R. A. and Rodrfguez-Luna, E.
1995. A species list for the New World primates (Platyr-
rhini): Distribution by country, endemism, and conserva-
tion status according to the Mace-Lande System. Neotrop.
Primates, 3(suppl.): 103-164.
Rylands, A. B., Mittermeier, R. A. and Rodrfguez-Luna, E.
1997. Conservation of Neotropical primates: Threatened
species and an analysis of primate diversity by country
and region. Folia Primatol. 68(3-5): 134-160.
Rylands, A. B., Schneider, H., Langguth, A., Mittermeier,
R. A., Groves, C. P and Rodrfguez-Luna, E. 2000. An as-
sessment of the diversity of New World primates. Neotrop.
Primates 8(2): 61-93.
Rylands, A. B., Mittermeier, R. A. and Konstant, W. R.
2001. Species and subspecies of primates described since
1990. Neotrop. Primates 9(2): 75-78.
SURAPA. 1999. Sistema de Areas Protegidas de la Amazonia:
Subred de Areas Protegidas del Amazonas. Food and Agri-
culture Organization / UN, Conservation International,
and Subred de Areas Protegidas del Amazonas, Bogoti.
CD-ROM.
Torres de Assumpcao, C. 1983. Conservation of primates
in Brazil: Atlantic Forest primates. In: Proceedings of the
Symposium on the Conservation of Primates and their Habi-
tats. 1. Primate Conservation in the Wild, D. Harper (ed.),
pp.34-48. University of Leicester, Leicester, UK.
von Spix, J. B. and von Martius, C. F. P. 1981. Viagem pelo
Brasil, 1817-1820. Vol. 1. 4a ed. EDUSP/Itatiaia, Sao
Paulo. 262pp.
Wied-Neuwied, M. P zu. 1821. Reise nach Brasilien in den
Jahren 1815 bis 1817. Vol. 2. 543pp.


BIRD PREDATION AND PREY-TRANSFER IN BROWN
CAPUCHIN MONKEYS (CEBUS APELLA)


Renata Ferreira, Briseida D. Resende
Massimo Mannu, Eduardo B. Ottoni,
Patricia Izar


Introduction


In the last decade capuchin monkeys, Cebus, have
received growing attention in the primatological literature
due to some striking convergences between them and
chimpanzees, Pan, such as: large brain size relative to body
size, long life span, tool use skills, and food-sharing among
group members (Fragaszy et al., 1990; Visalberghi and
McGrew, 1997). These similarities make capuchin monkeys
an attractive model for validating hypotheses about the
evolution of social life and cognition that are heavily based
upon Old World primates (Parr et al., 1997).

Food-sharing tendencies are of interest due to the view
that cooperative hunting with subsequent meat sharing
was a key factor in the adaptation and organisation of
early human societies (Butynski, 1982; Anderson, 1986;
McGrew and Feistner, 1992), and many studies have
focused on the cooperative hunting and meat sharing of
wild chimpanzees (Boesch, 1994). Some authors (e.g.,
Newcomer and De Farcy, 1985; Fragaszy, 1986) have
reported predation on vertebrates by capuchin monkeys in
different environmental conditions. However, the relation
between predation and prey sharing has only been analysed
for wild C. capucinus.

Rose (1997) reported predation on birds, coatis (Nasua
narica), and squirrels (Sciurus variegatoides) by two
groups of C. capucinus at Santa Rosa National Park,
Costa Rica. She concluded that, although predation
is a common event, food sharing is infrequent. Meat
is the most commonly shared food, and the only food
shared between adults (usually through falling scraps or
abandoned carcasses). Perry and Rose (1994) analysed the
sharing of captured coatis in three groups of C. capucinus
at two sites in Costa Rica. They concluded that: a) among
the species normally predated by capuchins, coatis are
riskier because adults are larger than adult capuchins and
normally defend their pups; and b) coati pups scream
while being eaten, so it is impossible for a monkey to
be rapid and furtive when eating them, giving plenty of
opportunities for other monkeys to beg from the carcass
owner.

The possible social value of food-sharing in captive groups
of C. api/vl has been emphasised by de Waal (1997; 2000;
de Waal et al., 1993) who observed that the occurrence of
this behavior is: a) related to affiliative relations and social
tolerance between pairs of individuals, b) linked to previous
events of food-sharing between the pair (even after a
delayed period) and, c) is more frequent when cooperation





Neotropical Primates 10(2), August 2002


is needed for predation or the acquisition of scraps from
other individuals.

Here we describe predation on birds and prey-sharing by
a semi-free group of brown capuchin monkeys (C. ,c/i//,).
Predation of this sort is rare, but when it does happen,
prey transfer is frequent. Due to the small number of
observations, the predation and particularly the prey-
transfer events are analysed only qualitatively, while
examining any relation between the occurrence of transfers
and the hierarchical and affiliative relationship between the
individuals involved.

Study Site, Group and Data Collection

The capuchin monkey group lives in an area of 18 ha in
the Tiete Ecological Park (Sao Paulo, Brazil). The area
was reforested and has two important features: 1) there
are no natural predators of Cebus, such as Harpia harpyja,
Felis pardalis, Boa constrictor or crocodiles (Freese and
Oppenheimer, 1981), and 2) the group is provisioned
daily with plentiful fruits and vegetables (see Ottoni and
Mannu, 2001). Besides the provisioning, the group forages
for other foods available in the area, taking up about 50%
of an individual's daily activities (RF, unpubl. data). The
wet season is from October to March, and the dry season
is from April to September (mean monthly rainfall is
178 mm and 69.3 mm, respectively) (Sao Paulo, DAEE,
2001). The group varied in size from 15 to 25 individuals:
five adult males (two castrated), five adult females, three
subadult males, one subadult female and 10 juveniles and
infants.

Our observations cover a period of five years, and were
collected on an ad libitum basis during the course of other
studies by MM (January 1996 to December 1999, see
Ottoni and Mannu, 2001), and BR and RF (2000/2001).
The total time of contact with the group was 2768 hours.

Results and Discussion

Table 1 summarises the occurrences of predation and
prey-transfers. The data are somewhat biased toward the
years 2000/2001 due to an increase in observation hours
per week relative to the previous years. For the purposes
of calculating the rates of prey sharing we consider two
types of predation data: a) the events when predation was
actually observed, and b) the predation event was not seen,
the animal merely being observed with a carcass.

Twenty-four predation events were recorded; a rate of 0.86
events per 100 observation hours. This is much lower than
the bird predation frequency described for C. capucinus in a
natural environment by Rose (1997); predatory behaviour,
however, did not constitute the focus of the studies in this
capuchin group as it did in Rose's study. Ten of the events
were observed in the first three years of observation (rate of
0.6 every 100 hours) and 14 in the remaining period when
the weekly hours of observation of the group were increased


(1.13 every 100 hours). This and the fact that predation,
and in many cases the consumption of the prey, is a very fast
and almost noiseless behaviour suggests that the frequency
is underestimated.

Predation frequency was the same between seasons: Dry
season 0.8 events every 100 hours, wet season 0.9 every
100 hours. Rose (1997) however, found a higher frequency
in the wet season (3.09 per 100 hours) when compared
to the dry season (2.04 per 100 hours). The lack of any
seasonal difference and the lower predation rates may be a
result of provisioning, but also to a reduced availability of
prey with our group ranging over a smaller, confined area
when compared to the C. capucinus of Rose (1997).

The 10 predation events recorded were all by males: six
by adults (four of these by the dominant male), three by
subadults and one by a juvenile. Of the 14 events in which
individuals were found with a carcass, the possessor was a
male (adult, subadult or juvenile) in 10 and an adult female
in three. In one case an adult male and an adult female eat
from the same carcass. Overall, the age/sex predation biases
are similar to those described for C. capucinus, where adult
males (especially the dominant) were the most efficient
predators (Perry and Rose, 1994; Rose, 1997).

Some sort of food transfer occurred in 18 of the 24
predation events. The transfers are classed as: co-feeding -
two individuals eat different prey near to each other (event
n 20); delayed scrounging- one individual eats the leftovers
of another (events no 4, 5, 6, 8, 9, 17, 19, 20, 22 and 23);
tolerated scrounging- the possessor allows another to come
near and retrieve dropped scraps (events no 4, 5, 7, 8, 16,
17 and 19); facilitated scrounging the possessor moves
towards an individual, drops food scraps and allows the
other to retrieve them (event no 10); passive food-sharing
- the possessor permits another to retrieve food items
from his/her hands or mouth (events n 3, 5, 13, 14, 17,
21 and 24); and theft one individual seizes the food
from another (event n 13). Note that different types
of food transfer can occur during the same predation
event, sometimes involving different individuals. (For a
discussion of terms and definitions see Ottoni et al., in
prep.) The proportion of prey sharing (in 18/24 predation
events) may be even greater if we consider that some of
the 'carcass' events may be the result of a previous non-
witnessed food-transfer. The predominant type of prey
transfer observed in this study was also the most common
type observed in C. capucinus by Rose (1997), that is, the
transfers were generally relaxed involving the collection of
leftovers or scraps.

Again, this high rate of prey-sharing that we observed may
be related to the food-abundance of the study site. A similar
phenomena was described in C. capucinus- higher rates of
prey-sharing were found in a rich environment (Lomas
Barbudal) than in an environment with marked seasonality
in food abundance (Santa Rosa, Costa Rica) (Perry and
Rose, 1994; Rose, 1997).





86 Neotropical Primates 10(2), August 2002


Table 1: Events and participants of predation and prey-transfer between individuals in a capuchin monkeys groups, Cebus apella, in the
Tiete Ecological Park, Sao Paulo.


Date Predation Prey-transfer Individuals
Observed Carcass
1. Sep/97 Juvenile male Jq or Qz


Juvenile male
Adult male and
sub-adult female
Adult male


5. Apr/99 Dominant
male


6. Oct 99 Dominant
male
7. Oct/99
8. Nov/99


9. Nov/99


Adult female
Dominant male


Dominant male


10. Dec/99 Adult male


11. Jun/00
12. Jul/00
13. Jul/00


Subadult male


Juvenile male


Adult male


14. Aug/00 Subadult male


15. Sep/00
16. Oct/00 Adult male


17. Nov/00


Adult female



Subadult male


18. Nov/ 00 Juvenile male
19. Jan 01 Subadult male


20. Feb 01 Dominant
male


Mar/01
Apr 01


Juvenile male
Adult female


Juvenile male


Dominant male


23. May/01


24. Jun/01


Passive sharing of a bird.


2. Sep/97
3. Jun/ 98


4. Jan/99


Med Jan


Mc -Ped


Bq Joao- Fis Man


Bq Frk


Fis Eli
Bq Med


Bq- Qz Med


Med-Jan


Frk- Edu


Subadult male collects scraps nearby and
then remains with the carcass
(Encaged bird). Adult male collects scraps
nearby. Adult female and infant eating the
carcass minutes later.
Juvenile eating the carcass minutes later.


Subadult male collects falling scraps nearby.
Adult male collecting scraps nearby after it
remains with the carcass.
Subadult male collected the discarded
carcass. Adult male collecting scraps nearby.
Adult male discarded carcass in front of
adult female.
Juvenile male interested.


Allows an infant but not a juvenile to take
some pieces of the carcass. Later the infant
remains with the carcass. Its mother steals
the carcass from him.
Avoids an adult male that follows him.
After 15 min the carcass' owner approaches
and permits a subadult female to take a
piece of the carcass.


Avoids a juvenile but allows an infant to eat
falling scraps nearby.
Subadult avoided adult male, who later
collected the carcass. Then subadult female
takes pieces of meat from the carcass, col-
lects scraps nearby and eats in contact with
adult male.
(Leaves the bird uneaten).
Allows a juvenile to eat falling scraps. Juve-
nile collects abandoned carcass.
(Predation on nestling birds). Adult male
cofeeding. Juvenile collects abandoned
carcass.
Infant takes pieces of meat from the carcass.
Dominant male collects the discarded
carcass.
Dominant male collects the discarded
carcass.
Adult female and juvenile taking pieces of
the carcass.


Qz- Kk-Jq


Sus Lob Dw


Qz-Jq- Kk


Frk
Ped Frk


Bq Med Joa


Man Dw
Fis Bq


Edu- Bq


Bq Fis Man


Med Joa Lob Jan





Neotropical Primates 10(2), August 2002


Prey transfers occurred 12 times between adults or
subadults: five from a male to a female (5, 10, 14, 17, 24);
two from a female to a male (7, 22); six between males (4,
5, 8, 9, 17, 20) and none between females. In seven events
the transfer was from an adult or subadult to a juvenile or
infant (5, 6, 13, 16, 19, 20, 24), and in two events in the
opposite direction (13, 23). One food transfer event was
between juveniles (21). As such, the frequency of sharing
between adults is greater than that between adults and
youngsters, and sharing occurs mainly between males or
from males to females. This contrasts with the observations
for C. capucinus, in which prey transfer was rarely observed
between adults and occurred mainly from mother to
infants or between immatures (Perry and Rose, 1994; Rose,
1997).

In 15 events the transfer was from a high to a low ranking
individual. In C. capucinus the rank of the possessor was
either unrelated to the direction of sharing or merely
facilitated the theft of the subordinate's prizes by the more
dominant individuals. Dominance relationships were
inferred by aggression, chasing, cowering, and avoidance,
and affiliative relationships were inferred by spatial
proximity and grooming (RF and PI, in prep.).

It is noteworthy that in 10 of 18 food transfers there was
a close affiliative relationship between the individuals
involved: in events 3 and 10, the female and male adults
were preferential partners in grooming, sleeping and
allocare (see Izar [1997] for descriptions on preferential
partnerships in C. ,,//la); in events 5, 22 and 24 the
transfers were between dominant male and female and their
offspring; in event 17 between an adult male and subadult
female that belonged to a small subgroup; in event 19,
transfer was between subadult and juvenile males which
were preferential partners in play. As affiliated individuals
stay close to each other, spatial proximity may be the factor
influencing the occurrence of transfers in these 10 events.
However, in another three events (described in greater
detail below) spatial proximity could not have been the only
factor, as there were at least three individuals close by, and
the possessor shared the prey with only one of them.

(Event 13) 00': Medeiros, an adult castrated male, is seen
eating a bird carcass. 10': Joana, an 11-month old infant
often carried and groomed by Medeiros, watches him,
collecting some scraps nearby. Lobato, a 3-year old juvenile
approaches, Medeiros chases him away. Joana bites pieces of
the carcass from Medeiros' hands. Medeiros leaves, Joana
remains with the carcass. 22': Janete (Joana's mother) steals
the carcass from Joana, who then suckles. [Medeiros and
Janete are the preferential partners already described in the
events 3 and 10].

(Event 16) 00': Noises indicating a fight are heard, and
Suspeito (a castrated adult male) leaves the area carrying
a bird in its mouth. 02': Suspeito eats the bird in a tree.
Lobato, a juvenile, approaches. Suspeito turns his back on
Lobato. Lobato approaches Suspeito again. Suspeito pushes


Lobato's head away from the carcass. 04': Lobato is nearby,
making some attempts to approach Suspeito. Suspeito
repeatedly turns his back or avoids Lobato. 07': Darwin
approaches Lobato and tries to play with him. Darwin sees
Suspeito. 07'30": Darwin approaches Suspeito and collects
some scraps. 09': Darwin takes a small piece of the carcass
and eats it. Suspeito moves higher in the tree. 11': Darwin
approaches Suspeito, takes another piece of the carcass, and
eats it. 12': Darwin starts playing with Lobato, Suspeito
remains with the carcass. 25': Suspeito leaves the carcass.
[In this event, there is social affinity between Suspeito and
Meire (Darwin's mother), similar to that observed between
Medeiros and Janete, that is, Suspeito and Meire are
preferential partners for sleeping and grooming, although
Suspeito does not allocare Darwin as much as Medeiros
allocares Joana.]

(Event 14) 00': Quinzinho, a subadult male, catches a bird.
Joaquim, an adult male, witnesses the predation. 01': After
eating the head of the bird, Quinzinho walks carrying the
prey in his mouth. Joaquim follows him for about 50 m.
Kika, a subadult female, also begins to follow him. 03':
Quinzinho stops in a tree and eats the bird for about 15
minutes. After some failed attempts to approach Quinzinho,
Joaquim leaves the area. Meanwhile, Kika remains foraging
about 20 m from Quinzinho. 18': Quinzinho, still holding
the carcass, approaches to 1 m from Kika. She approaches
him, makes an aggressive display (not towards him) and
then takes a big piece of the carcass. They both eat in close
proximity for another 5 minutes. 23': they leave. [Again,
the social relationships data show that Quinzinho and
Kika are "preferential partners". In contrast, Quinzinho
and Joaquim were seen fighting several times (Joaquim is
dominant over Quinzinho)].

In the first two events the carcass owner clearly tolerated
the approach and begging of an individual with which it is
affiliated but not from another with which it is less affiliated.
In the third event, the possessor avoided the approach
attempts of one individual and actively approached another,
with which it is affiliated, and shared the meat.

We are not sure whether the observed differences between
C. j,,i//,. and C. capucinus in predation rates and prey
transfer rates and directions are due to the type of prey, to
the study site or to the species under study. Predation on
birds differs from predation on coatis because birds may
be easily caught and eaten secretively by the individuals.
Robinson (1986) reported that capuchins successfully
foraging on nestling birds were discrete in finding a nest,
and frequently moved away from the rest of the group.
The particular characteristics of this study site make the
results difficult to generalise. However, other studies have
shown differences in territorial behaviours and hierarchical
rigidity of C. q,',,//. and other capuchins (C. q,',,//, is a non-
territorial and more despotic species) (Janson, 1986; Perry,
1998) which suggests the possibility of specific differences
in the dynamics involving social relationships and food
sharing.





Neotropical Primates 10(2), August 2002


Recently, Mitani and Watts (2001) compared three
hypotheses about the hunting and sharing of meat in wild
chimpanzees. Their data did not support the ecological
(i.e., in periods of food shortage) or hunting-for-sex
hypothesis, but did support the hypothesis that the sharing
of meat is used as a social tool to enhance bonding between
adult males.

Although the bird predation events described here did not
involve cooperative hunts by the group members, sharing
does seem to be influenced by the affiliative relationships
in the group. There are indications that individuals of
C. ,'I//. are capable of distinguishing and behaving
differentially towards other group members. Janson (1984)
described non-tolerance by the dominant males towards
another males' offspring in feeding trees. The work of de
Waal (1997, 2000; de Waal et al., 1993) also suggests this
capacity. Overall, the analysis of prey transfer described
here, and most especially in three events, suggest that, in
C. ,''//., highly valuable food items are preferentially
shared with more affiliated individuals.

The drawbacks in data collection and analysis and the
many possible proximate variables interfering in these
events of meat sharing (for example, recent fights between
the individuals involved or how hungry the carcass
owner is), do not allow us to be conclusive about
the dynamics involving affinity and food sharing.
Nevertheless, the apparent refusal to share with some
individuals and tolerance towards others in three events
raises two questions: to what extent are these tripartite
events of food transfer indicative of the social complexity
and social knowledge of the capuchin monkeys? Likewise,
is preferential prey sharing a tool for improving and
maintaining valuable relationships within the C. ,i///.
groups? Experiments on food transfer in situations
involving three individuals, and further observations of
other tripartite relations, such as coalitions, could help to
answer these questions.

Acknowledgments

The authors thank the staff of the Tiete Ecological Park,
especially to Priscilla T. Oliveira and Liliane Milanelo. This
work was funded by CAPES grants to RF (no 2172/97-6)
and MM, and FAPESP grants no 99/11573-2, 97/14443-
7, 00/14590-4 to BR, EO and PI respectively.

Renata Ferreira, Department of Biological Anthropology,
University of Cambridge, Downing Street, Cambridge
CB2 3DZ, England, UK, Briseida Dogo de Resende,
Massimo Mannu, Eduardo B. Ottoni, Departamento
de Psicologia Experimental, Instituto de Psicologia,
Universidade de Sao Paulo, Avenida Professor Mello
Moraes 1721, 05508-900 Sao Paulo, Brazil, and Patricia
Izar, Departmento de Ecologia Geral, Instituto de
Biociencias, Universidade de Sao Paulo, 005508-900 Sao
Paulo, Brazil.


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the brown capuchin monkey Cebus ,aj'//I. (Primates:
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Janson, C. 1986. The mating system as a determinant
of social evolution in capuchin monkeys (Cebus). In:
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C. Lee (eds.), pp.169-179. Cambridge University Press,
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Int. J. Primatol. 18: 677-681.



THE BLACK HOWLER MONKEY (ALOUATTA PIGRA)
AND SPIDER MONKEY (ATELES GEOFFROYI) IN THE
MAYAN SITE OF YAXCHILAN, CHIAPAS, MEXICO: A
PRELIMINARY SURVEY

Alejandro Estrada, LeAndra Lluecke, Sarie Van Belle
Kirk French, David Muioz, Yasminda Garcia
Lucia Castellanos, Adridn Mendoza
Introduction

The black howler monkey of Mesoamerica, Alouatta pigra,
has a restricted geographic distribution in Belize, Guatemala
and Mexico. The majority of its range (c. 80%) is in Mexico
in parts of the states of Tabasco and Chiapas, and it is the
only Alouatta species present in the Yucatin peninsula
(Smith, 1970; Horwich and Johnson, 1986; Watts and
Rico-Gray, 1987). Spider monkeys (Ateles .. -r.., coexist
with A. pigra in many areas, but because of hunting for
food and to obtain pet infants, and the destruction and
fragmentation of their forests, they are among the most
endangered primates in Mesoamerica (Kinzey, 1998).

Information on population parameters and conservation
status for A. pigra are available from only a few localities,
namely, two sites in Belize in the Bermuda Landing and
Cockscomb Wildlife Reserve (Horwich, 1998; Silver et
al., 1998; Ostro et al., 1999, 2000), in Tikal, Guatemala
(Coelho et al., 1976), in the Muchukux forest in Quintana
Roo, Mexico (Gonzailez-Kirchner, 1998) and in Palenque,
Chiapas, Mexico (Estrada et al., 2002). A similar situation
prevails in the case of A. ..-rr.. . with information
available only from few localities in Mexico, namely Los
Tuxtlas, Veracruz (Estrada and Coates-Estrada, 1996; Silva-
L6pez and Jfmenez-Huerta, 2000) and the Muchukux and
Naji Tucha forests in Quintana Roo, Mexico (Gonzilez-
Kirchner, 1999). Some information is available on
populations of A..rn .. 'from Tikal, Guatemala (Coelho
etal., 1976).


Such paucity of information and the rapid fragmentation
and conversion of the natural habitat of A. pigra and
A. ,..rr .., to pasture lands and agricultural fields in
northern Mesoamerica, coupled with intensive hunting
pressure and trafficking of infants as pets, makes the task
of protecting these primate species particularly difficult
(Estrada and Coates-Estrada, 1988; Rylands et al., 1995).
Data on group size, density, and age and sex composition
for populations of A. pigra and A., ..rr .., in large forest
tracts and in landscapes modified by man may provide
information on the variability of population parameters,
and may also improve our understanding of their tolerance
of habitat loss and fragmentation (Estrada and Coates-
Estrada, 1996; Estrada et al., 1994; Crockett, 1998;
Cuar6n, 2000).

In this paper we provide preliminary data on group
size, population density and demographic structure for
populations of A. pigra and A. ..rr .., in the protected
forest surrounding the ruins of the Mayan site of Yaxchilin,
Chiapas, Mexico. The data we present are part of a series
of surveys of primate populations inhabiting the protected
forests surrounding major Mayan archeological sites in
southern Mexico (Estrada et al., 2002; in prep.).

Methods

Study area and sites
The study was carried out at the Mayan site of Yaxchilin,
Chiapas, Mexico (1653'N, 9057'W, 250 m above sea
level), near the Rio Usumacinta, that marks the international
boundary between Mexico and Guatemala (Fig. 1). There is
a protected forest of about 2700 ha surrounding the Mayan
site, of which 1100 ha are contained within an omega-
shaped area by the river, while the rest extends inland
(Fig. 1). This forest is connected to 35,000 ha of protected
rain forest in the Community Reserve "La Cojolita". The
climate is hot and humid, and average annual precipitation
is 1951 mm, with a dry season from December to April
(average monthly rainfall = 42.4 +12.7) and a wetter period
from May to November (average monthly rainfall = 256.0
+100.1 mm). Mean annual temperature is 25.5 +2.2C
(range 21-28C).

Tall evergreen rain forest (tree heights between
15-45 m) is the dominant vegetation at the study site
(). Abundant trees in this forest
are Brosimum alicastrum, B. costaricanum, Poulsenia armata,
Ficus glabrata (Moraceae), Manilkara zapota, Pouteria sapota
(Sapotaceae), Bursera simaruba (Burseraceae), Lonchocarpus
sp. (Fabaceae), and Spondias spp. (Anacardiaceae) (Meave,
1990).

The Mayan site dates back to about 500 AD (Coe, 1998).
Only about 5% of the ruins of the site have been excavated,
the rest are covered by rain forest vegetation, and vestiges
of buildings can be easily observed amidst the vegetation or
roots of trees. Several of the Mayan structures were built at
the top of the many hills, while the majority of the largest





Neotropical Primates 10(2), August 2002


buildings and plazas are found close to the edge of the Rio
Usumacinta (Figs. 1, 2).

Primate surveys
Primate surveys were conducted in November 2001 and
in April 2002 in a 100 ha area around the Mayan ruins
of Yaxchilan. We triangulated early morning (0500 hrs)
choruses to determine the presence and location of howler
monkey troops. Vocalizations emitted by spider monkeys
were also recorded for the same purpose. An existing system
of trails was used to gain access to different parts of the
100 ha area. To triangulate monkey vocalizations in the
early morning, we climbed to the top of the tallest (50 m)



/I
1KM

B N

CH I
GUATEMALA /





Figure 1. Location of Yaxchilin by the Rio Usumacinta
demarcating the international boundary between Mexico and
Guatemala. The omega-shaped area comprises the protected
forest of the Mayan site, about 1100 ha, with an additional 1600
ha inland. The black dot is the area where the ruins are located.
The dotted line around the omega shape shows the route followed
during the river survey of howler and spider monkeys. CH =
Chiapas, B = Belize.


Figure 2. Study area (100 ha) where the Mayan ruins of Yaxchilin
are located. The shaded area in the upper right corner is the Rio
Usumacinta. Codes refer to howler monkey troops detected. The
three dots without code are three troops that were heard howling,
but could not be located. Grid cells are 200 x 200 m.


Mayan structures (Temples 39, 40 and 41) from which we
could monitor vocal exchanges between troops in the 100
ha area. The direction from which vocalizations were heard
were determined with a compass and plotted on a detailed
map of the site.

Two teams searched for the monkeys. An average of 8.0
+2.0 hrs/day was spent exploring different sections of the
study area by walking slowly (1.0 km/hr) through the
forest or along the existing system of trails. When a troop
of howler monkeys or a subgroup of spider monkeys was
sighted we noted its location on a detailed map of the
archeological buildings. A GPS (Garmin GPS III, Kansas,
USA) was used to obtain precise georeference points. We
completed 200 man/hours and walked 62.3 km surveying
howler and spider monkeys in the study area.

Contacted howler and spider monkey groups were followed
for several hours and repeatedly counted by each team
to confirm identification and age and sex composition.
Individuals were classified as adults, juveniles and dependant
infants. Accurate identification of the sex was only possible for
the adults and juveniles. All groups detected were found and
followed on consecutive days, further aiding in confirmation
of their size and composition and identity. We carefully
examined consistency in the age and sex composition of
each group, their location in relation to the trail system
and topographical and archaeological features of the terrain,
and their relative location with other troops. Trees in which
howler and spider monkeys were sighted were measured
(height and diameter at breast height dbh). Average weights
of A. pigra and A ... rr .., available in the literature (Coelho
et al., 1976) were used to estimate the biomass (kg/ha)
represented by the monkeys in the study area.

Two additional surveys of howler and spider monkeys
were conducted from a boat by navigating 13.7 km along
the Rio Usumacinta, following the contour of the omega-
shaped area where the site of Yaxchilan is located (Fig. 1).
They started at 0530 hrs and were conducted down river
(S-N direction) with the outboard motor off, allowing
for a gentle and silent flow of the boat. Vocalizations and
sightings were located using the GPS. A GPS reading was
taken every 500 m to obtain an estimate of the length and
width of the omega-shaped study area.

Results

Howler monkeys
Vocalization surveys resulted in the detection of 11 troops of
howler monkeys in the 100 ha area surrounding the Mayan
buildings. Eight were in the forest and repeatedly counted
on consecutive days, yielding a total 54 howler monkeys
and one solitary male. The other three troops could not be
found, but were heard howling on other days in the same
general location (W-SW of the ruins) (Fig. 2).

Forty-one percent of the individuals counted were adult
males, 30% were adult females, 8% were juvenile males,





Neotropical Primates 10(2), August 2002


6% were juvenile females and 15% were infants (Table 1).
Average troop size was 6.6 +2.1 individuals, ranging from
4 to 10. Troops had an average of 2.8 +1.6 adult males,
2.0 +0.5 adult females, 0.8 + 0.4 juvenile males, 1.0 +0.0
juvenile females and 1.3 +0.5 infants (Table 1).

Adult male to adult female ratio was 1:0.73, and in juveniles
the sex ratio was 1:0.75. Adult to non-adult ratio was 1:
0.40, and adult female to immature ratio was 1:0.97 (Table
1). Using the average troop size calculated for the eight
counted, and the total number of troops detected, howler
monkey density in the 100 ha study area was estimated at
0.72 ind/ha or 72.6 ind/km2. Total biomass represented by
the eight troops and the solitary male was estimated at 367
kg, and mean biomass per troop was 44.8 +15.0 kg. Using
this last figure, we estimated howler monkey biomass at
492.8 kg/km2 or 4.9 kg/ha.

Spider monkeys
We confirmed the existence of three subgroups of spider
monkeys in the 100 ha study area with a total of 17
individuals. They were observed several times in different
locations, and sometimes in the same trees as howler
monkeys. Their identity was confirmed by the size of the
subgroup and by its age and sex composition. Adult males
accounted for 35.3% of individuals counted, adult females


Table 1. Results of the survey of howler and spider monkeys in a 100
at the edge of the Rio Usumacinta.


for 29.4%, juvenile males for 5.9%, juvenile females for
11.8% and infants for 17.6% (Table 1).

Mean subgroup size was 5.67 +3.06 individuals, and mean
sex and age composition of these subgroups was 2.00 +1.00
adult males, 1.67 +1.15 adult females, 1.50 +0.71 juveniles
and 1.50 +0.71 infants. The adult male to adult female sex
ratio was 1:0.83 and in juveniles it was 1:2.0; the adult
female to immature ratio was 1:1.20. Density was estimated
at 0.17 ind/ha or 17 ind/km2 and spider monkey biomass at
106.45 kg/km2 or 1.06 kg/ha (Table 1).

Vegetation types used by howler and spider monkeys
All sightings of howler and spider monkeys were in tall
evergreen rain forest. The mean height and dbh of trees
used by howler monkeys were 11.07 +6.9 m (range 4-30
m) and 63.4 +28.5 cm (range 45-120 cm), respectively. In
the case of spider monkeys, mean height and dbh of trees
used were 19.6 +7.3 m (range 4-30 m) and 78.7 +28.3 cm
(range 45-120 cm), respectively. Spider monkeys preferred
taller trees than howler monkeys (U test, P<0.01) (Fig. 2).

River survey
The survey down the Rio Usumacinta along the contour
of the omega-shaped study area resulted in the auditory
detection of 17 troops of howler monkeys and one subgroup


ha area around the Mayan site ofYaxchilin, Chiapas, Mexico, located


Adult Juvenile
--------- -------- -------- ------------------T o t a l
Males Females Males Females Infants
Alouatta pigra
T25 5 2 0 1 2 10
T33 3 2 1 1 2 9
T30 2 2 1 1 6
R1 1 2 1 4
R2 5 2 1 8
PA 2 1 1 1 5
T41 1 3 1 5
LSTRIP 3 2 1 6
Total 22 16 4 3 8 53
Mean 2.8 2.0 0.8 1.0 1.3 6.6
+ sd 1.6 0.5 0.4 0.0 0.5 2.1
Solitary males 1 1
Total howler monkeys 54
Ateles geoffroyi
Subgroup
1 1 1 1 3
2 3 1 1 5
3 2 3 2 2 9
Total spider monkeys 6 5 1 2 3 17
Mean 2.00 1.67 1.00 2.00 1.50 5.67
+ sd 1.00 1.15 0.00 0.00 0.71 3.06





Neotropical Primates 10(2), August 2002


of spider monkeys, along a stretch of 13.7 km. Sixty-five
percent of the howler monkey troops (n = 11) and the single
spider monkey subgroup were detected on the Mexican side
of the river. Howler monkey troops were detected at a rate
of 0.80 troops/km surveyed on the Mexican side and 0.48
troops/km surveyed on the Guatemalan side.

Discussion

The results of the primate surveys presented here should
be viewed as preliminary. Further field work will provide
information on the consistency and variability of the
demographic traits we have observed for A. pigra and A.
..-rr .. at Yaxchilan. Our surveys showed that howler
troops and Ateles subgroups were detected at a rate of 0.18
troops/km and 0.048 subgroups/km surveyed, respectively,
confirming that A. pigra is more numerous than A. .. r ..,
at Yaxchilan. The 13.7 km river survey along the edges of
the omega shape area in which Yaxchilan is located, also
showed a predominance of howler monkeys, with spider
monkeys present in lower numbers.

Howler monkeys
The density of 72.6 individuals/km2 we estimated for
A. pigra in Yaxchilan is significantly higher than those
reported for this species in other large rain forest tracts in
Mexico, such as Muchukux, Quintana Roo (15.1 ind/km2;
Gonzalez-Kirchner, 1998) and Calakmul and Palenque,
Chiapas (15.2 ind/km2 and 23 ind/km2, respectively;
Estrada et al., 2002, in prep.). Coelho et al. (1976) and
Schlichte (1978) reported a density of 5-9 individuals/km2
(1978) atTikal, Guatemala.

High densities of A. pigra have been reported from Belize,
ranging from 47-178 individuals/km2 in fragmented strips
of riparian vegetation and small forest patches, which


26-30

21-25
E
I-
2 16-20
ui
11-15

6-10


J

JM


MAteles N = 28
DAIouatta N = 50


authors have indicated may be due to overcrowding (Silver
etal., 1999; Ostro etal., 1999, 2000; Horwich etal., 2001).
However, the high densities found in Yaxchilan and in other
large tracts of rain forest such as Calakmul, Campeche
(Estrada et al., in prep.), Palenque, Chiapas (Estrada et
al., 2002), and in Muchukux, Quintana Roo (Gonzalez-
Kirchner, 1998), seem to contradict such an assumption.

Mean troop size in Yaxchilan (6.6 +2.1 individuals)
compares to troop sizes reported for A. pigra in Calakmul,
Campeche (7.5 +2.3 individuals; Estrada et al., in prep.)
and Palenque, Chiapas (7.0 +2.8 individuals; Estrada et al.,
2002), but they are higher than those reported in Belize and
Guatemala, where mean troop size varies from 4.4 to 6.3
individuals (Coelho etal., 1978; Bolin, 1981; Horwich and
Gebhard, 1983; Ostro et al., 1999), and the small troops
averaging 3.16 individuals reported for A. pigra in central
Quintana Roo, Mexico (Gonzalez-Kirchner, 1998).

Seventy-five percent of the troops detected in Yaxchilan
had more than one adult male, as was found in Palenque,
Chiapas, and in Calakmul, Campeche, where 75% and
60% of the troops, respectively, were multimale (Estrada et
al., 2002, in prep.). However, at Tikal, Guatemala, troop
surveys by different authors consistently reported unimale
troops (Coelho et al., 1976; Schlichte, 1978; Horwich and
Johnson, 1986). In Yaxchilan, Calakmul, and in Palenque
the overall adult sex ratio was 1:0.73 to 1:0.90 (Estrada et
al., 2002, in prep.). Data from Belize showed most recorded
troops to be unimale, and the adult sex ratio was 1:1 to
1:1.63 (Bolin, 1981; Ostro et al., 1999; Horwich et al.,
2001).

Spider monkeys
The density (17 ind/km2) we report for A. ..-rr .. in
Yaxchilan falls within the range reported for the species in
other extensive tracts of rain forest in Quintana Roo, Mexico,
such as Najil Tucha (14.5 ind/km2) and Muchukux (27.1
ind/km2) (Gonzalez-Kirchner, 1999), and in Calakmul (25
ind/km2), Campeche, Mexico (Estrada et al., in prep.). In
Tikal, Guatemala, densities for A. ,... ..i were found to
range from 26 to 45 ind/km2 (Coelho et al., 1976; Cant,
1978), while in Costa Rica population densities ranged
from 6-9 ind/km2 (Freese, 1976; Chapman, 1988). In
fragmented landscapes in Los Tuxtlas, Mexico, A. ..-rr ..,
is found at densities of 0.22 ind/km2 (Estrada and Coates-
Estrada, 1996), but in more extensive forest in the same
region, the density was reported at 0.66 ind/km2 (Silva-
L6pez and Jfmenez-Huerta, 2000).


Spider monkey subgroup size (5.6 +3.06 individuals) in
Yaxchilan is similar to that reported for A. ..-rr .., in
0 10 20 30 40 50 Calakmul, Campeche (6.6 individuals) (Estrada et al., in
% USED prep.), in Chiapas, Mexico (5.0 individuals) (Eisenberg,
1966) and in Los Tuxtlas, Veracruz at 0.66-6.2 individuals
Figure 3. Distribution of heights of trees used by spider and (Silva-L6pez et al., 1988, 2000). It is higher than that
howler monkeys at the site of Yaxchilin. Note the preference by reported for the Muchukux and Najil Tucha forests in
spider monkeys for tall trees (>10 m), whereas howler monkeys Quintana Roo, where subgroup sizes averaged 4.5 and
preferred trees >4 and < 25 m). 3.8 individuals, respectively (Gonzalez-Kirchner, 1999), as





Neotropical Primates 10(2), August 2002


well as in Belize and Guatemala (4.5 and 2.6 individuals,
respectively) (Coelho et al., 1976; Cant, 1978, 1990),

The adult sex ratio detected in Yaxchilan (1:0.83) strongly
favoring adult males, contrasts with the ratio reported
for the same species in Calakmul (1:1.96) (Estrada et
al., in prep.) and in Muchukux and Najil Tucha forests
in Quintana Roo (1:1.26) (Gonzalez-Kirchner, 1999). A
sex ratio of 1:1.56 was reported for spider monkeys in
disturbed forest areas in Los Tuxtlas, Veracruz (Silva-L6pez
et al., 1988), while a ratio of 1:3.25 was reported for a
population of spider monkeys in an undisturbed forest
site in the same region (Silva-L6pez and Jfmenez-Huerta,
2000). The adult sex ratio for A. geoffroyi reported in
Tikal, Guatemala was 1:2.23 (Coelho et al., 1976). The 1:
1.20 adult female to immature ratio in Yaxchilan suggests
a population with a capacity to sustain itself and grow
(Clarke et al., 2002).

Spider monkeys in Yaxchilan preferred the tall trees of the
upper canopy (70% used were 16- > 30 m in height), as has
been noted in other localities in Mexico, such as Quintana
Roo (Gonzalez-Kirchner, 1999) and Calakmul (Estrada
et al., in prep.), besides other Neotropical sites (Van
Roosmalen and Klein, 1988; Symington, 1988; Yoneda,
1990). They can, however, be seen at all levels of the forest
when traveling and will often forage in low trees bearing
ripe fruit. The howler monkeys in Yaxchilan preferred lower
strata than spider monkeys, spending much more time in
the middle and lower canopy. This was similar to our
observations in Calakmul (Estrada et al., in prep.).

General comments
The differences in population parameters for A. pigra and
A. ...-r..-, r. r ., Yaxchilan and other sites, may be within
the natural variation in their populations, due to hunting
or to the lack of data on both species in Mexico, Belize
and Guatemala. Clearly, more sites need to be surveyed to
document the range of variation in density, group size and
other demographics for A. pigra and A. -rr .., within the
range of their current geographic distribution in northern
Mesoamerica.

In this vein of thinking, it has been indicated that A. pigra
is typically found in riparian forests at elevations below 400
m, and that the population in Tikal, Guatemala (Ostro et
al., 2000) is exceptional. However, our survey in Yaxchilan
showed no concentration of howler troops along the Rio
Usumacinta; the majority of the troops detected were
distributed inland. In Palenque, Chiapas and Calakmul,
Campeche in Mexico, A. pigra populations are common
in the non-riparian habitats dominating these sites, and
in Palenque they occur in forests at 500 m above sea level
(Estrada et al., 2002; Estrada et al., in prep.).

While discriminating separate howler monkey groups is
relatively easy, it is more difficult for spider monkeys. The
members of relatively large groups or communities travel in
small temporary subgroups of unstable composition (Van


Roosmalen and Klein, 1988; Kinzey, 1996). Because of
the fission-fusion nature of their social organization it is
rare to see all members of the community together, and
group sizes are difficult to estimate (Coelho et al., 1976;
Klein and Klein, 1977). The surveys conducted along
the Rio Usumacinta in Yaxchilan detected more howler
and spider monkeys on the Mexican side than on the
Guatemalan side of the river. During our surveys we noted
much deforestation (slash and burn), as well as hunting
(rifle shots heard several times) on the Guatemalan side.
Although preliminary, these observations suggest the need
for further surveys to better assess and monitor the status
of A. pigra and A. .. rr ..i along the international border
formed by the Rio Usumacinta.

The presence of the important Mayan ruins at Yaxchilan
has resulted in the permanent protection of the surrounding
rain forest, and its populations of A. pigra and A. ..nrr ,
Yaxchilan. This is also true of sites such as Palenque (Estrada
et al., 2002) and Calakmul (Estrada et al., in prep.), and
together they constitute important foci for the conservation
of A. pigra and A. .. r.. in this area of Mesoamerica.

Acknowledgments

We are grateful to the Lincoln Zoo Scott Neotropic Fund
for support to conduct this study. We thank the Instituto
Nacional de Antropologia e Historia through Lic. Juan
Antonio Ferrer, Director of the archeological site of
Yaxchilan, for permission to carry out the primate surveys
and for providing invaluable logistical support.

Alejandro Estrada, Laboratory of Primatology, Estaci6n
de Biologfa Los Tuxtlas, Instituto de Biologfa, Universidad
Nacional Autonoma de M6xico, LeAndra Lluecke,
Department of Anthropology, University of Texas, Austin,
Texas, USA, Sarie Van Belle, Kirk French, Department of
Anthropology, University of Cincinnati, Cincinnati, Ohio
45221, USA, David Mufioz, Yasminda Garcia, Lucia
Castellanos andAdrin Mendoza, Laboratory ofPrimatology,
Estaci6n de Biologfa Los Tuxtlas, Instituto de Biologfa,
Universidad Nacional Autonoma de M6xico, Apartado
Postoal 176, San Andres Tuxtla, Veracruz, Mexico 95700.
E-mail: Alejandro Estrada .

References

Bolin, I. 1981. Male parental behavior in black howler
monkeys (Alouatta palliata pigra) in Belize and Guate-
mala. Primates 22: 349-360.
Cant, J. G. H. 1978. Population survey of the spider
monkey (Ateles .. rr .., I at Tikal, Guatemala. Primates
19: 525-535.
Cant, J. G. H. 1990. Feeding ecology of spider mon-
keys, Ateles ..n .., at Tikal, Guatemala. Hum. Evol. 5:
269-281.
Chapman, C. A. 1988. Patterns of foraging and range use
by three species of neotropical primates. Primates 29:
177-194.





Neotropical Primates 10(2), August 2002


Clarke, M. R., Crockett, C. M. and Zucker E. L. 2002.
Mantled howler population of Hacienda La Pacifica,
Costa Rica, between 1991 and 1998: Effects of deforesta-
tion. Am.J. Primatol. 56: 155-163.
Coe, A. 1998. Archeological Mexico. Moon Publications,
Inc., Chico, California.
Coelho, A. M. Jr., Coelho, L., Bramblett, C., Bramblett, S.
and Quick, L. B. 1976. Ecology, population characteris-
tics, and sympatric association in primates: A socio-bio-
energetic analysis of howler and spider monkeys in Tikal,
Guatemala. Yearb. Phys. Anthropol. 20: 96-135.
Crockett, C. 1998. Conservation biology of the genus
Alouatta. Int. J. Primatol. 19: 549-578.
Crockett, C. and Eisenberg, J. E 1987. Howlers: Variation
in group size and demography. In: Primate Societies, B. B.,
Smuts, D. L. Cheney, R. M. Seyfarth, R. W. Wrangham
and T. T. Struhsaker (eds.), pp.56-68. The University of
Chicago Press, Chicago.
Cuar6n, A. D. 2000. Effects of land-cover changes on
mammals in a Neotropical region: A modeling approach.
Conserv. Biol. 14: 1676-1692.
Estrada, A. and Coates-Estrada, R. 1988. Tropical rain
forest conversion and perspectives in the conservation
of wild primates (Alouatta and Ateles) in Mexico. Am. J.
Primatol. 14: 315-327.
Estrada, A, and Coates-Estrada, R. 1996. Tropical rain
forest fragmentation and wild populations of primates at
Los Tuxtlas. Int. J. Primatol. 5: 759-783.
Estrada, A. R., Coates-Estrada, R., and Meritt, D., Jr.
1994. Non flying mammals and landscape changes in
the tropical rain forest region of Los Tuxtlas, Mexico.
Ecography 17: 229-241.
Estrada, A., Castellanos, L., Garcia, Y., Franco, B., Munoz,
D., Ibarra, A., Rivera, A., Fuentes, E. and Jimenez, C.
2002. Survey of the black howler monkey, Alouatta pigra,
population at the Mayan site of Palenque, Chiapas,
Mexico. Primates 44: 51-58.
Freese, C. H. 1976. Censusing Alouatta palliata, Atelesgeof-
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In: Neotropical Primates: Field Studies and Conservation,
R. W. Thorington Jr. and P. G. Heltne (eds.), pp.4-9.
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Gonzilez-Kirchner, J. P. 1998. Group size and population
density of the black howler monkey (Alouatta pigra) in
Muchukux forest, Quintana Roo, Mexico. Folia Primatol.
69: 260-265.
Gonzilez-Kirchner, J. P. 1999. Habitat use, population
density and subgrouping pattern of the Yucatin spider
monkey (Ateles .. .. yucatanensis) in, Quintana Roo,
Mexico. Folia Primatol. 70: 55.60.
Horwich, R. H. 1998. Effective solutions for howler con-
servation. Int. J. Primatol. 19: 579-598.
Horwich, R. H. and Gebhard, K. 1983. Roaring rhythms
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E. 1995. A species list for the New World primates
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Schlichte, H. 1978. A preliminary report on the habitat
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pp.551-561. Smithsonian Institution Press, Washington,
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spider monkeys (Ateles .. r . vellerosus) in the forest
of the crater of Santa Marta, Veracruz, Mexico. Neotrop.
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1988. The status of Ateles geojfroyi and Alouatta palliatain
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R. 1988. Feeding ecology of the black howler monkey
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263-279.
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45-67.
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Neotropical Primates 10(2), August 2002


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New WorldMonkeys, La Macarena, Colombia 3: 13-18.



A POSSIBLE EXAMPLE OF COERCIVE MATING
IN MANTLED HOWLING MONKEYS (ALOUATTA
PALLIATA) RELATED TO SPERM COMPETITION

Clara B. Jones
Introduction

If females with internal fertilization mate more than once
during a reproductive cycle, the potential exists for sperm
from more than one male to compete for access to eggs
inside her reproductive tract. This phenomenon, termed
"sperm competition," has received increasing attention
in the mammalian, including primate, literature in recent
years (e.g., Harcourt, 1997; Gomendio etal., 1998; Dixson,
1998). Several studies have demonstrated that coercive
mating ("coercive copulations," "forced copulations,"
"rape") may be associated with multiple mating by female
insects (Moller, 1998), and Smith (1984; also see Thornhill
and Palmer, 2000) suggested that human rape might have
evolved as an adaptation to sperm competition, although
there is no necessary or sufficient relationship between rape
and sperm competition. The present short communication
describes an apparent case of coercive mating in the context
of sperm competition for mantled howling monkeys
(Alouatta palliata), a species in which coercive mating has
not been reported. Attempts to force copulations by male
mantled howlers are generally rebuffed successfully by
females with a bared-teeth, open-mouth display, sometimes
accompanied by vocalizations (Jones, 1985). Such behavior
may implicate coercive mating in the evolutionary history
of this species. A reanalysis of the raw data for the 1985
report revealed one case of apparently coercive mating in
association with multiple mating by a female.

Methods

The study was conducted in 1976 and 1977 at Hacienda
La Pacifica, Cafias, Guanacaste, Costa Rica (10028'N,
85007'W). Details on the research, including group
composition (Group 5), methods (focal), habitat (riparian),
social organization (multimale-multifemale), sexual
behavior, and life history can be found elsewhere (Jones,
2000 and references therein).


Results

On 5 March 1977 (late dry season), the focal animal was
male R, the third and lowest-ranking male in the study
group. He was observed to lie and rest in a tree along the
Rio Corobici with female PY who demonstrated evidence
of estrus -2 (a few days subsequent to "peak" estrus [see
Jones, 1985]). A sub-adult/young adult male who had
not yet joined the male hierarchy rested approximately 50
feet downriver, and several adult females and young were
nearby. No other adult males were sighted. Male Rvocalized
continuously with high intensity guttural barks to female
PY, the young male, and/or anotherr individualss. While
the functions of vocalizations in mantled howlers have not
been investigated, these barks have been interpreted to
communicate motivation (Jones, 2000).

At 1:04 pm, male R initiated the stereotyped lingual gesture
(tongue moving rapidly in and out of mouth, a sexual signal
characteristic of the genus Alouatta [Carpenter, 1934]) with
female PY, who responded in kind. Reciprocal lingual
gesturing continued for 3 min. Male R subsequently
lay rear-present to female PY, a posture that has been
interpreted as submissive (Jones, 2000). At 1:20 pm, male
R sat up and mounted female PY, thrusting 40 times in 37
sec with an ejaculatory pause. Subsequent to copulation, the
pair rested. During the resting period, male R occasionally
emitted high guttural barks.

At 2:21 pm, female PY initiated lingual gesturing with male
R. The male, lying on a branch with the female, failed to
reciprocate the female's lingual gestures and shifted posture
in apparent vigilance. At 2:30 p.m., male R initiated lingual
gestures with female PY and subsequently sat up, looking
downriver in the direction of the young male. At 2:33 pm,
female PY moved 30 ft upriver past male R, sitting rear-
present to the male. Male R moved towards the female and
sat 10 ft behind her. Female PY continued to move upriver.
Male R continuously vocalized at low intensity.

At 2:37 pm., male G, the second-ranked male of Group
5, moved rapidly upriver past male R, mounted female
PY without preliminaries to copulation, and trusted
approximately 37 times with an ejaculatory pause. The
female did not resist intromission, did not exhibit the open-
mouth bared-teeth display, or emit vocalizations. Male R
moved downriver (away from male G and female PY),
continuously emitting low intensity vocalizations, began
feeding at 2:42 pm ("sham feeding"? [Carpenter, 1934]),
and continued to feed, sit, and mingle with other group
members (including sexual inspection of several adult
females) until 4:50 pm when this day's record ended.

Discussion

Multiple mating by mantled howler females has been
previously documented (Carpenter, 1934; Jones and
Cortes-Ortiz, 1998). The case reported here, however,
provides evidence that sperm competition may occur





Neotropical Primates 10(2), August 2002


in mantled howlers, combined with apparently coercive
mating by the second male to copulate, and may exemplify
cases of coercive mating in primate species in which females
mate multiply (e.g., Pongo: Rodman and Mitani, 1987;
Dixson, 1998). The above copulation by male G was
judged to be coercive because there were no preliminaries,
because this male appeared to intercept female PY from the
male guarding her (male R), and because male G's mount
appeared to be executed hastily and with some degree of
force since intromission occurred in association with rapid
movement. In this situation, it might be expected that
there would be potential for an escalated conflict situation
between the two males, possibly explaining female PY's
failure to resist male G's advances, as well as explaining
male R's vocalizations. The present report, however,
cannot completely exclude the possibility that some visual
or auditory signal was exchanged between female PY and
male G which might have communicated receptivity by
this female to the male. Nonetheless, it may provide direct
evidence for Smith's (1984; also see Thornhill and Palmer,
2000) idea that coercive mating may operate in relation to
sperm competition in primates. Future studies of primate
reproductive behavior should consider the likelihood that
coercive mating is beneficial to males, and possibly to
females (Moller, 1998, p. 72), in some ecological and social
conditions.

Acknowledgments. I thank W. G. Daniel and three
anonymous reviewers for constructive comments on a
previous version of this note and A. B. Rylands for editorial
advice.

Clara B. Jones, Department of Psychology, School
of Liberal Arts, Livingstone College, Salisbury, North
Carolina 28144, USA. E-mail: ,
.

References

Carpenter, C. R. 1934. A field study of the behavior and
social relations of howling monkeys. Comp. Psychol.
Monog. 10: 1-167.
Dixson, A. F 1998. Primate Sexuality. Oxford University
Press, Oxford.
Gomendio, M., Harcourt, A. H. and Roldan, E. R. S. 1998.
Sperm competition in mammals. In: Sperm Competition
and Sexual Selection, T. R. Birkhead and A. P. Moller
(eds.), pp.667-756. Academic Press, San Diego.
Harcourt, A. H. 1998. Sperm competition in primates. Am.
Nat. 189: 189-194.
Jones, C. B. 1985. Reproductive patterns in mantled
howler monkeys: Estrus, mate choice, and copulation.
Primates 26: 130-142.
Jones, C. B. 2000. Alouatta palliata politics: Empirical and
theoretical aspects of power. Prim. Rep. 56: 3-21.
Jones, C. B. and Cortis-Ortiz, L. 1998. Facultative
polyandry in the howling monkey (Alouatta palliata):
Carpenter was correct. Bol. Primatol. Latinoamer. 7: 1-7.


Moller, A. P. 1998. Sperm competition and sexual selection.
In: Sperm Competition and Sexual Selection, T. R. Birkhead
and A. P. Moller (eds.), pp. 55-90. Academic Press, San
Diego.
Rodman, P. S. and Mitani, J. C. 1987. Orangutans: Sexual
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Smith, R. L. 1984. Human sperm competition. In: Sperm
Competition and the Evolution of Animal Mating Systems,
R. L. Smith (ed.), pp.601-659. Academic Press, San
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Thornhill, R. and Palmer, C. T. 2000. A Natural History of
Rape: Biological Bases of Sexual Coercion. The MIT Press,
Cambridge.







PRIMATE OBSERVATIONS IN SURINAME

Brownsberg Nature Park
I visited Brownsberg Nature Park (4o57'06.8"N,
5510'45.5"W) from 2-7 February, 2002 to assist in
expanding a camera-trapping monitoring program. Located
in the state of Broko-Pondo, north-central Suriname, the
park is a 21/2 hour drive from the capital city of Paramaribo
and hence is Suriname's most heavily visited protected
area. It is a long and narrow, flat-topped plateau (504 m
elevation) with heavily forested, steep slopes and gullies
on all sides. The forest is strongly dominated by Hevea
guianensis, Terminalia dichotoma, Cecropia spp. and several
guava species (Family Myrtaceae).

I observed primates on a number of occasions while setting
up camera traps to record large mammals and terrestrial
birds. All observations were within 6 km of the visitor's
center and were made from or near established park trails. I
observed red howler monkeys (Alouatta seniculus) each day
and heard their loud and long (30 seconds or more) pre-
dawn chorus beginning at 5:45 am. Black spider monkeys
(Atelespaniscus) were heard daily, their calls emanating from
the forest below the plateau. A single, male white-faced saki
(Pitheciapithecia) was observed for several minutes near the
visitor center on the plateau. Golden-handed tamarins (Sa-
guinus midas) were seen from a trail at 12:20 pm. Bearded
saki monkeys (Chiropotes satanas) were observed at 2:20 pm
in a tree nearby feeding red howler monkeys. Common
squirrel monkeys (Saimiri sciureus), wedge-capped capu-
chins (Cebus olivaceus), and brown capuchins (C. ,,'//I)
were not observed but are listed in the management plan as
being present in the park.

Central Suriname Nature Reserve
I also visited the Central Suriname Nature Reserve from
8-16 February, 2002, to establish a camera trapping





Neotropical Primates 10(2), August 2002


monitoring program. The reserve headquarters is located
at Raleighvallen (443'39.4"N, 5612'34.3"W) where a
permanent grass airstrip is maintained. During my daily
excursions to place camera traps between the headquarters
and the Voltzberg, 6 km distant, I observed red howler
monkeys (Alouatta seniculus), brown tufted capuchins
(Cebus q'c//i), black spider monkeys (Ateles paniscus),
bearded saki monkeys (Chiropotes satanas), and common
squirrel monkeys (Saimiri sciureus) with young nearly
every day during my stay. Less frequently, I also observed
wedge-capped capuchins (Cebus olivaceus), golden-
handed tamarins (Saguinus midas) and, on three occasions,
separate single groups of one to three white-faced saki
monkeys (Pithecia pithecia). Red howler monkeys and
bearded saki monkeys were observed once in the same
fruiting tree.

Acknowledgments: I thank STINASU, the Foundation for
Nature Conservation in Suriname, for my research permit.
I also thank Bart De Dijn, Sutrisno Mitro, and Raymon
Clemens (STINASU), Iwan Dervold, Semmie Andre,
Kupias Tawadi, and Kamajna Panashekung (Conservation
International) for their assistance.

Jim Sanderson, Center for Applied Biodiversity Science,
Conservation International, 1919 M Street NW, Suite
600, Washington, DC 20036, USA, e-mail: conservation.org>.



MURIQUIS NO PARQUE NATIONAL DA SERRA Dos
ORGAOS

Um estudo sobre os muriquis no Parque Nacional da
Serra dos Orgaos, Teres6polis, Rio de Janeiro (Programa
de Conservagao do Muriqui), esta sendo realizado desde
j aneiro de 2002 corn a finalidade de localizar os grupos ainda
existentes e determinar o ndmero de individuos observados,
assim como a composigao sexo etaria dos mesmos.

O Parque Nacional da Serra dos Orgaos possui cerca de
11.800 ha, apresentando um relevo bastante acidentado
com escarpas ingremes que se elevam acima da floresta
ombr6fila densa da Serra do Mar. Quatro tipos de vegetagao
sao presents dentro da Area do parque: floresta ombr6fila
densa submontana, floresta ombr6fila montana, floresta
ombr6fila alto montana e campos de altitude. Alim do
muriqui, ocorrem tries outras esp&cies de primatas no
parque: ( .-' aurita, Cebus nigritus e Alouatta guariba.

Ate o moment foram realizadas tries expedigoes, sendo
que os muriquis foram encontrados em duas delas. Ao
todo, encontramos dois grupos e contamos 17 animals
(sete individuos num grupo e 10 em outro). Acreditamos
que sejam Brachyteles arachnoides, a forma sulina do
muriqui, pois alguns individuos que foram observados
durante mais tempo tinha a face bem negra. Os grupos
estavam compostos por femeas e machos adults e alguns


individuos jovens. Informao6es sobre as caracteristicas da
vegetagao na qual os muriquis estao sendo encontrados,
bem como coleta de amostras de plants das quais foram
observados comendo, e fezes de individuos que por ventura
defecarem, tambem estao sendo registrados e coletadas. As
fezes estao sendo armazenadas a fim de serem estudadas
as caracteristicas gen&ticas da populagao do muriqui no
parque (estudo a ser realizado pela Dr. Val&ria Antunes da
Universidade Federal do Espirito Santo) e tambem para
serem feitas anAlises parasitol6gicas. Sementes encontradas
nestas fezes serao colocadas para germinar, para que no
future possam ser plantadas em Areas que precisem ser
recuperadas dentro do parque.

A reacao dos animals com relagao a presenga dos
pesquisadores foi bastante hostile, o que prejudicou uma
melhor observagao e contagem mais correta de todos os
individuos. Estressados, os animals fugiram rapidamente e,
devido as dificuldades de se andar nesta Area nao foi possivel
acompanha-los por muito tempo. Mas, quando os animals
ficam nervosos, geralmente eles defecam e neste moment
aproveitamos a oportunidade para coletar as suas fezes.

Um outro registro interessante que nos foi passado por
um excursionista foi o fato de um muriqui ter sido visto
e fotografado numa Area do parque, cuja a vegetagao
predominante e de campos de altitude a mais de 2.000 m
de altitude. Era apenas um indivfduo que passou correndo
pelo chao e depois se embrenhou num mosaico arbustivo de
altura baixa. Isto eleva um pouco mais o limited altitudinal
da esp&cie de 1.800 metros registrada por Aguirre (1971)
para 2.000 metros. As raz6es pelas quais este indivfduo
tenha usado este tipo de ambiente podem estar baseadas
em tries suposigoes: ou ele estava fugindo das matas que se
encontram nas altitudes mais baixas de algum predador
ou cagador; ou ele estava a procura de outras fontes de
alimento nao encontradas nas matas de altitude mais baixa;
ou estava apenas atravessando de uma mata para outra
usando o campo de altitude.

Paralelamente a este estudo, um trabalho de educadao
ambiental esta sendo realizado no entorno do parque com a
finalidade de conscientizar e aumentar o conhecimento das
pessoas que vivem nesta regiao sobre o pr6prio muriqui e a
importancia de preservar a ele e seu ambiente.

Agradecimentos- Este estudo esta sendo financiado por um
convenio celebrado entire o IBAMA e a Organizagao Nao-
governamental TEREVIVA (No. 45/01) para o Programa
de Conservagao do Muriqui no Parque Nacional da Serra
dos Orgaos.

Vania Luciane Alves Garcia, Programa de Conservagao
Muriqui, Parque Nacional da Serra dos Orgaos, Av.
Rotariana s/no., 25960-602 Alto-Teres6polis, Rio de
Janeiro, Brasil, e-mail: ,
e Jovelino Muniz de Andrade Filho, Diretor, Parque
Nacional da Serra dos Orgaos, Av. Rotariana s/no., 25960-
602 Alto-Teres6polis, Rio de Janeiro, Brasil.





Neotropical Primates 10(2), August 2002


FUNCTIONAL ANATOMY OF THE CALLITRICHID
FORELIMB AND LONG BONES

Lesa C. Davis defended her Ph.D. dissertation entitled
"Functional Morphology of the Forelimb and Long
Bones in the Callitrichidae (Platyrrhini, Primates)" at the
Department of Anthropology, Southern Illinois University
at Carbondale on 3 May, 2002. Her doctoral advisor was
Dr. Susan M. Ford. Financial support for this study was
provided by the National Science Foundation (DBS 92-
03884), a Smithsonian Institution Short-Term Research
Grant, and a Sigma Xi Scientific Research Society Grant-
In-Aid of Research. The following is an abstract of the
dissertation.

The small-bodied callitrichids use a diversity of positional
behaviors in negotiating their habitat and procuring food.
The primary goal of this study was to identify species-level
anatomical correlates to locomotor, postural, and foraging
behaviors in the forelimb and long bones in callitrichids.
Nine callitrichid species, representing five of the six
recognized genera, were included in the study: (...
pygmaea, ( .-.' .... '. C. penicillata, r ..
S. midas, S. ...rr .., S. mystax, Leontopithecus rosalia, and
Callimico goeldii. An extensive comparative outgroup of
select strepsirhine and non-callitrichid platyrrhine species
was used to help determine which morphological attributes
of callitrichids may be 'universal' consequences of specific
activities, which are the result of heritage, and which are
uniquely callitrichid. Quantitative osteological data from
the scapula, humerus, radius, ulna, hand, and hindlimb
long bones were analyzed using univariate and multivariate
statistical methods. In addition, these data were tested
for significant correlations with positional behavioral
frequencies for each species.

Results indicate that callitrichids are distinguished from
the outgroup on the basis of scapular morphology. Within
the Callitrichidae, Leontopithecus rosalia has a uniquely
elongated and gracile forelimb, built, at least in part,
for manipulative foraging. Tamarins, Saguinus midas,
S. .. rr .., and S. mystax share a suite of elbow and forearm
adaptations for quadrupedalism while their close relative
S. fuscicollis is more similar to L. rosalia in many features.
Callimico exhibits several significant adaptations for vertical
clinging, some of which are exclusive of another platyrrhine
vertical clinger, Cebuella. Morphofunctional analysis of
( .-.'. jacchus and C. penicillata was limited due to
the lack of locomotor and postural data available for these
taxa, but a few features in the scapula and radius unite the
marmosets. This study supports growing evidence that body
size and behavior do not form a predictable relationship
and that standard indices, such as the intermembral index,
are limited in their usefulness as predictive tools. These, and
additional morphofunctional correlations, are crucial for
understanding the evolutionary radiation of this primate
family, and become even more critical in importance as new


species, both extant and extinct, continue to be discovered
in the wild.

Lesa C. Davis, Assistant Professor of Anthropology,
Northeastern Illinois University, Department of
Anthropology, 5500 N. St. Louis Ave., Chicago, IL 60625,
USA. E-mail: .

Reference

Davis, L. C. 2002. Functional Morphology of the Forelimb
and Long Bones in the Callitrichidae (Platyrrhini,
Primates). Doctoral dissertation, Department of
Anthropology, Southern Illinois University, Carbondale.



DIET, ACTIVITY AND HOME RANGE OF ALOUATTA
BELZEBUL DISCOLOR (PRIMATES, ATELIDAE) IN
PARANAfTA, NORTHERN MATO GROSSO

On 28 January, 2002, Liliam Patricia Pinto defended her
master's thesis entitled "Dieta, padrao de atividades e area
de vida de Alouatta belzebul discolor (Primates Atelidae) em
Paranafta, norte do Mato Grosso" for the Graduate Program
in Ecology, Department of Zoology, Universidade Estadual
de Campinas (UNICAMP), Sao Paulo, Brazil. Her supervisor
was Eleonore Z. F. Setz, and the study was supported by the
Fundacao de Amparo a Pesquisa do Estado de Sao Paulo
(FAPESP). The following is an abstract of the thesis.

This study examined the feeding ecology of seven to nine
red-handed howlers (Alouatta belzebul discolor) in primary
forest in the north of the state of Mato Grosso, southern
Amazonia. The red howler, Alouatta seniculus, also occurred
in the area, and the interactions between the species are
described. Diet, activity patterns, day range, and home
range in A. b. discolor were recorded by instantaneous scan
sampling over 10 months, between October 1999 and Oc-
tober 2000. During 45 complete days, howlers fed on 67
plant species (n = 2039 feeding records) from 24 families.
Dialium guianense (Leguminosae, Caesalpinioideae) was
the species with the most records. Fruits were predominant
in the diet (55.6% of records), followed by new leaves
(19.8%) and flowers (5.7%). Mature leaves formed only a
small part of the diet (5.0%). Bark, live wood, dead wood,
and woody branches together comprised 10.2% of the diet.
During the study period, the group spent 58.7% of the day
resting and sleeping, 20% feeding, 14.2% traveling, 4.0%
moving around within trees, and only 2.1% in social inter-
actions. Although there were no significant differences in
time budget between the rainy and the dry seasons, there
were slight changes in activity patterns. In the rainy season
there were fewer activity peaks and they retired to their
sleeping trees earlier than in the dry season. Home range,
calculated by superimposing a grid of quadrats, was 50.1
ha. The group used all of the three habitat types in their
range; 23 ha terra firme forest, 4 ha of acaf (Euterpe) palm
forest, and 23.1 ha of seasonally flooded forest ( ..'i The





Neotropical Primates 10(2), August 2002


convex polygon method gave a home range of 63.2 ha.
The home range was larger than has been reported for any
other species of Alouatta except in Central America. Aver-
age day range length was 761 m (n = 45) and did not differ
significantly between seasons. Dialium guianense fruits were
abundant during the dry season, and their intensive use
contributed to the absence of significant seasonal variation
in fruit consumption, activity patterns and day range.


Reference

Pinto, L. P. 2002. Dieta, padrao de atividades e Area de
vida de Alouatta belzebul discolor (Primates, Atelidae)
em Paranaita, norte do Mato Grosso. Master's thesis,
Departamento de Zoologia, Universidade Estadual de
Campinas (UNICAMP), Sao Paulo.


DIETA, PADRAO DE ATIVIDADES E AREA DE VIDA
DE ALOUATTA BELZEBUL DISCOLOR (PRIMATES, LISTA DAS SPECIES DA FAUNA AMEACADAS DE
ATELIDAE) EM PARANAfTA, NORTE DO MATO EXTINCAO NO RIO GRANDE DO SUL, BRASIL
GROSSO


Este estudo aborda aspects da ecologia de um grupo de
bugios-de-maos-vermelhas (Alouatta belzebul discolor)
composto por sete a nove individuos, em mata primAria
no norte do estado do Mato Grosso, sul da Amazonia. E
relatada a presenca de mais uma esp&cie de bugio na area,
Alouatta seniculus, com descrihao das suas interaqoes. A
dieta, o padrao de atividades diarias e a area de vida de
A. b. discolor foram investigadas durante dez meses, entire
outubro de 1999 e outubro de 2000, atraves do m&todo
de varredura instantInea. Durante 45 dias completos, os
bugios se alimentaram de 67 especies de plants (n = 2039
registros de alimentacao) de 24 families. Dialium guianense
(Leguminosae, Caesalpinioideae) foi a especie mais utilizada
na alimentacao. Os frutos foram os itens mais consumidos
(55,6%), seguidos de folhas novas (19,8%) e flores (5,7%).
A participacao de folhas maduros na dieta foi pequena
(5,0%). Casca de Arvores, lenho de tronco vivo, madeira em
decomposicao e ramos lenhosos, juntos perfizeram 10,2%
da dieta. Durante todo o perfodo, o grupo gastou 58,7%
do tempo em repouso, 20,0% em almentacao, 14,2% em
viagem, 4,0% em movimentacao, e somente 2,1% em
interao6es sociais. Nao houve diferencas significativas entire
as estac6es chuvosa e seca no tempo dedicado as principals
atividades. No entanto, a distribuiiio das atividades ao
long do dia foi diferente nas duas estaq6es. Na estadao
chuvosa, o recolhimento para a Arvore de dormir se deu
mais cedo e houve um ndmero menor de picos de atividades
do que na estacao seca. Os bugios utilizaram tries tipos de
ambiente: terra firme, igap6, e acaizal (Euterpe). A Area de
vida, calculada pelo m&todo de quadriculas, foi de 50,1 ha,
sendo 23,0 de floresta de terra firme, 4,0 ha de acaizal, e
23,1 ha de igap6. Corn o m&todo de polfgono convexo,
obteve se 63,2 ha. Areas de vida maiores que a observada
neste estudo s6 tem sido encontrados em bugios na Am&rica
Central. 0 percurso diArio mddio do grupo foi de 761 m
(n = 45) e nao various significativamente entire as estaq6es.
Frutos de Dialium guianense foram abundantes durante a
estacao seca. A utilizacao intensive deste item contribuiu
para a ausencia de variaqao sazonal significativa no consume
de frutos, nos padres de atividade e de uso de espaco.

Liliam Patricia Pinto, Programa de P6s-graduaqao em
Ecologia, Departamento de Zoologia, Instituto de Biologia,
Universidade Estadual de Campinas, Caixa Postal 6109,
13083-970 Campinas, Sao Paulo, Brazil.


A iniciativa de elaborar a primeira lista das esp&cies da
fauna ameacadas de extincao no Rio Grande do Sul surgiu
de forma independent em dois grupos de pesquisadores
gadchos. No final de 1999, a associaqao ambientalista
PANGEA, com o apoio da Fundacao Biodiversitas,
estabeleceu contatos com a Secretaria Estadual do Meio
Ambiente (SEMA) e com alguns pesquisadores gadchos no
sentido de dar infcio ao process de elaboraqao de uma lista
a ser sancionada por decreto governmental.

Paralelamente, em agosto do mesmo ano, teve infcio o
projeto Livro Vermelho da Fauna Ameagada de Extindao
no Rio Grande do Sul, do Museu de Ciencias e Tecnologia
da Pontificia Universidade Cat61lica do Rio Grande do Sul
(MCT-PUCRS), corn a proposta mais ampla de elaborar,
alum de uma lista, tambem um livro sobre as especies
ameacadas no Estado, gerenciando as informaqoes atraves
de uma base de dados permanentemente atualizAvel.

Para evitar a duplicidade de esforcos e elaborar uma lista
dnica, as equipes foram unidas sob coordenaqao geral do
"Projeto Livro Vermelho", contando com apoio da SEMA
atraves da Fundacao ZoobotInica do Rio Grande do Sul. A
lista aqui apresentada result do trabalho desenvolvido desde
entno. Ela foi elaborada a partir do esforgo conjunto de 43
zo6logos diretamente vinculados ao projeto, representando
18 instituio6es de pesquisa, e 128 colaboradores.

Corn o prop6sito de formalizar o apoio da Secretaria
Estadual do Meio Ambiente ao Projeto Livro Vermelho
e encaminhar o process de homologacao da lista pelo
Governo do Estado, um convenio entire o MCT-PUCRS e a
SEMA foi firmado em agosto de 2001. Posteriormente, em
5 de dezembro do mesmo ano, os resultados de dois anos
de trabalho do projeto foram sinteticamente apresentados
a sociedade gadcha em uma audiencia p6blica organizada
pela SEMA. Participaram representantes do poder pdblico
estadual e federal, organizadores da lista, zo6logos vinculados
ao projeto, colaboradores, pesquisadores, t&cnicos e
representantes de organizacoes nao governamentais, alum
da comunidade.

Ao final, 261 especies foram classificadas como
efetivamente ameacadas de extincao no Rio Grande do Sul,
enquadrando-se nas categories de ameaga descritas no texto
do decreto (Tabela 1).





Neotropical Primates 10(2), August 2002


Tabela 1. Nrumero de species ameagadas no Rio Grande do Sul
por grupo e categoria de ameaga. As siglas seguem recomendaygo
da IUCN, utilizando a grafia inglesa para facilitar a consult por
pesquisadores de diferentes nacionalidades.

Grupo Categorias de Ameaca
RE PE CR EN VU Total
Esponjas 1 2 3
Moluscos 6 11 17
Crusticeos 7 7
Insetos 7 11 18
Peixes 4 6 18 28
Anfibios 10 10
Repteis 5 12 17
Aves 2 8 31 42 45 128
Mamiferos 1 8 5 19 33
Total 2 9 43 72 135 261
RE Regionalmente extinto; PE Provavelmente extinto;
CR Criticamente em perigo; EN Em perigo; VU Vulnerivel.


Com relacao aos primatas, Alouatta caraya (Humboldt,
1812) e Alouatta guariba clamitans Cabrera, 1940 foram
classificadas como Vulnerivel (VU) e Cebus nigritus nigritus
(Goldfuss, 1809) como Dados Insuficientes (DD). A
destruigao e descaracterizagao dos habitats constituem as
principals fatores que representam ameaga as populagoes
de primatas no Estado. Como medidas para conservagao
desses tixons sao sugeridas a protegao e recuperacao do
habitat, juntamente com programs de educacao ambiental
e estudos de auto-ecologia.

Ana Alice B. de Marques, Universidade do Vale do
Rio dos Sinos, Avenida Unisinos, 950, 93022-000 Sao
Leopoldo, RS, Brasil, e-mail: , Carla
S. Fontana, Museu de Ciencias e Tecnologia, Pontificia
Universidade Cat61lica do Rio Grande do Sul, Avenida
Ipiranga, 6681, Caixa Postal 1429, 90619-900 Porto Alegre,
RS, Brasil, Eduardo V61lez, Museu de Ciencias Naturais,
Fundagao Zoobotanica do Rio Grande do Sul, Rua Dr.
Salvador Franga 1427, 90690-000 Porto Alegre, RS, Brasil,
Glayson A. Bencke, Laborat6rio de Ornitologia, Fundagao
Zoobotanica do Rio Grande do Sul, Rua Dr. Salvador
Franga 1427, 90690-000 Porto Alegre, RS, Brasil, Mauricio
Schneider, Departamento de Zoologia, Universidade
Federal do Rio Grande do Sul, Avenida Bento Gongalves
9500, Bloco IV, pr. 43435, 90540-000 Porto Alegre,
RS, Brasil, e Roberto E. dos Reis, Museu de Ciencias e
Tecnologia, Pontificia Universidade Cat61lica do Rio Grande
do Sul, Avenida Ipiranga, 6681, Caixa Postal 1429, 90619-
900 Porto Alegre, Rio Grande do Sul, Brasil.

Reference

Marques, A. A. B., Fontana, C. S., Vdlez, E., Bencke, G. A.,
Schneider, M. and Reis, R. E. 2002. Lista das Esp&cies da
Fauna Ameacadas de Extinao no Rio Grande do Sul. Decreto
n,41.672, de 11 de junho de 2002. FZB/MCT-PUCRS/
PANGEA, Porto Alegre. 52pp.


BRASIL TEM UM CENTRO DE PROTECDO DE
PRIMATAS

Para estudar e proteger as 95 especies de primatas presents
em territ6rio national, o Instituto Brasileiro do Meio
Ambiente e dos Recursos Naturais Renoviveis (IBAMA)
criou um centro especifico. 0 Centro de Protegao de
Primatas Brasileiros foi criado atraves da Portaria No. 148
de 18 de outubro de 2001. A sede operational do projeto
seri em Cabedelo, na regiao metropolitan de Joao Pessoa,
no estado da Parafba. 0 edificio e um antigo engenho de
agdcar, onde ji funcionam outras unidades de informadao
do Ibama e a semente do Centro de Protegao de Primatas
Brasileiros. 0 bi6logo do Ibama, Marcelo Marcelino,
responsivel pelo Centro, diz que os pesquisadores atuarao
em todo o territ6rio national, coletando dados sobre os
animals.

"Vamos estudar primatas de todos os tipos em todo o
Brasil. Queremos sair a campo para coletar informagoes
sobre definigoes geogrificas, doengas que afetam os animals
e ocorrencias em unidades de conservagao, alim de ordenar
os dados que ji existem e organizer o material para que
possa subsidiary polfticas de conservagao das especies", disse.
"Pretendemos ir ao campo ji no ano que vem e queremos
comegar a trabalhar logo na Amaz6nia, que e nosso alvo
preferential pela riqueza do espago geografico e pelo
ndmero de especies. Tambdm temos urgencia de pesquisar
o material que ji existe em unidades de conservagao, para
saber o que ji esti sendo protegido ou nao".

O Centro vai investigar ainda a ocorrencia de doengas entire
as populagoes de primatas. Para isso a equipe de Marcelino
deve recorrer aos arquivos pdblicos ji existentes. "Tem
muito material esquecido, que sequer foi publicado. N6s
vamos buscar tudo o que hi disponivel na literature e em
arquivos pdblicos para fazer uma ordena~io e gera~io de
informao6es", anunciou o pesquisador.

A equipe do projeto deve ser composta inicialmente por
cinco tecnicos, e o Centro vai concentrar toda a atuadao
do Ibama em relagao a primatas brasileiros: Conservagao,
pesquisa, com&rcio illegal, transport e a criacao e manejo
em cativeiro.

O Conselho Cientifico do Centro foi instituido atraves
da Portaria MMA/IBAMA No. 846, 19 de julho de
2002, para analisar as quest6es apresentadas pelo Centro
e oferecer subsidios tecnicos e cientificos para a tomada
de decis6es. E composto pelas seguintes especialistas:
Jdlio Cesar Bicca-Marques (Universidade Cat61lica do Rio
Grande do Sul), Adelmar F. Coimbra Filho (Academia
Brasileira de Ciencias), Stephen F. Ferrari (Universidade
Federal do Pari), Alfredo Langguth (Universidade Federal
da Parafba), Alcides Pissinatti (Centro de Primatologia do
Rio de Janeiro CPRJ), Anthony B. Rylands (Center for
Applied Biodiversity Science, Conservation International,
e a Universidade Federal de Minas Gerais) e Cliudio





Neotropical Primates 10(2), August 2002


Valladares-Padua (IPE Instituto de Pesquisas Ecol6gicos
e a Universidade de Brasilia).

Marcelo Marcelino, Chefe, Centro de Protegao de
Primatas Brasileiros, Instituto Brasileiro do Meio Ambiente
e dos Recursos Naturais Renovaveis (IBAMA), BR-230
Km 10, Mata da AMEM, 58.310-000 Cabedelo, Paraiba,
Brasil. E-mail: .



AREA DE PROTEcAO AMBIENTAL DA BACIA DO RIO
SAo JOAO/MIco-LEAO-DOURADO

No dia 26 de julho de 2002, foi criada a Area de Protedao
Ambiental (APA) da Bacia do Rio Sao Joao/Mico-Leao-
Dourado, corn 150,529 ha, espalhados por seis municipios
do estado do Rio de Janeiro. Engloba os principals
remanescentes de Mata Atlantica, onde ainda habitat o
mico-ledo-dourado, Leontopithecus rosalia, abrangendo
o entorno das Reservas Biol6gicas de Poco das Antas e
Uniao, ambas gerenciadas pelo Instituto Brasileiro do
Meio Ambiente e dos Recursos Naturais Renovaveis
(IBAMA). As duas reserves foram exclufdas da APA por
serem de categories mais restritivas, e as principals areas
de atuaqao da Associaqao Mico-Leao-Dourado (AMLD)
por meio do Programa de Conservacao da especie que
vem desenvolvendo desde 1983. Corn a APA, a regiao
passa a ter criterios para a ocupacao e o uso do solo que
profbem a degradacao ambiental e promovem a gestao
ambiental participativa e responsAvel por parte do poder
pdblico e da iniciativa privada. Corn isso, fica assegurada a
integridade da biodiversidade local e muitos outros services


prestados pela bacia do Rio Sao Joao, sendo a dnica fonte
hfdrica para abastecimento pdblico de toda a regiao dos
Lagos Fluminenses, desde Saquarema ate Rio das Ostras,
passando por Cabo Frio, Bdzios e Araruama.

Denise Margal Rambaldi, Diretora Executiva, Associacao
Mico-Leao-Dourado (AMLD), Rodovia BR 101 Km 214,
Caixa Postal 109.968, 28860-970 Casimiro de Abreu, Rio
de Janeiro, Brasil.



THE LION TAMARINS OF BRAZIL FUND

The Lion Tamarins of Brazil Fund receives donations from
zoos and captive breeding institutions which hold lion
tamarins to support conservation activities in the wild:
Brazilian field researchers, surveys, censuses, behavioral
and ecological studies, translocations and reintroductions,
public education, reforestation and land acquisition
(Neotropical Primates 2(suppl.), pp.4-7, 1993). As report-
ed by Jeremy J. C. Mallinson in Tamarin Tales (Volume 6,
2002), during the fiscal year (September 2000 August
2001) the fund received a record US$46,841, currently
being distributed directly to Brazilian conservation and re-
search programs on lion tamarins. Thirty-nine zoos, three
individuals, a regional zoo organization, and the Margot
Marsh Biodiversity Foundation were the contributors.
Since its creation in 1992, this fund has contributed enor-
mously to the success of the in situ conservation programs
for the four species, Leontopithecus rosalia, L. chrysomelas,
L. chrysopygus and L. caissara, in Brazil's Atlantic forest.





Neotropical Primates 10(2), August 2002


Jeremy Mallinson has retired as Co-Custodian of the
Fund. Bengt Holst, Vice-Director of the Copenhagen
Zoo and mastermind behind a highly successful cam-
paign (2001-2002) for the Atlantic forest and lion
tamarin conservation by the European Association
of Zoos and Aquaria (EAZA), has taken over the task
as the European counterpart of Devra Kleiman and
Jonathan Ballou who run the Fund on the other side of
the pond.

Contributions from North and South America should be
arranged with Jonathan D. Ballou, Department of Con-
servation Biology, National Zoological Park, Smithsonian
Institution, Washington, DC 20008, USA, Tel: +1 202 673
4828, Fax: +1 202 673 4686, e-mail: ,
and from Europe, Africa, Asia and Australasia with Bengt
Hoist, Copenhagen Zoo, Sdr. Fasanvej 79, DK-2000
Frederiksburg, Denmark, Tel: +45 72 200 220, Fax: +45
72 200 219, e-mail: .



8TH EUROPEAN STUDBOOK FOR SAGUINUS
IMPERATOR

With the help of Joana Lobo, and the Lisbon Zoo staff,
Orlando Silva and Patricia Vilarinho, Eric Bairrao Ruivo,
Animal Collections Coordinator of the Lisbon Zoo in Por-
tugal, has released the 8th European Studbook for the em-
peror tamarin, Saguinus imperator. The data is current to
31t December, 2001, and records 158 (84.66.8) tamarins,
all of the subspecies S. i. subgrisescens. No S. i. imperator
remain in Europe, only a small, non-breeding population
of eight hybrids (5.3.0). There were a total of 59 births
during 2001, although unfortunately their survival was
poor 33 died. There were 20 adult deaths and as such
the population remained quite stable, increasing by 7
from 2000. In his introduction, Eric Bairrao Ruivo credits
the high birth rate to the cooperative management of the
population. It is now spread through 54 institutions 41
with S. i. subgrisescens and five with non-breeding hybrids.
Besides European Zoos, the studbook also registers ani-
mals in Australia (currently, Perth- hybrids, 2.1.0; Sydney
- 2.0.0; and Melbourne- 2.0.0) and Singapore (0.1.0).
Aaalborg Zoo, Denmark, and Paris Zoo, France, received
the species for the first time in 2001. In 2002 a further
five institutions will enter the breeding programme. The
studbook provides a full historical listing of the species,
analyses of the demographics since 1964, age distribution,
inbreeding coefficients, and founder representation and
mean kinship. The recommendations for transfers and
breeding during 2002 are also detailed. The studbook in-
cludes an extensive report on a study of sexual differences
in behavioral patterns in captive S. i. subgrisescens carried
out by Ruivo himself.

Eric Bairrno Ruivo, EEP Co-ordinator for Emperor
Tamarin, Jardim Zool6gico de Lisboa, Estrada de Benfica
158, 1549-004 Lisboa, Portugal.


References

Ruivo, E. B. 2001. European Suul,,oo for the Emperor Tam-
arin Saguinus imperator ssp. 8th Edition, 2001. Lisbon
Zoological Garden, Lisboa, Portugal. 182pp. (Data cur-
rent through 31 December 2001)
Ruivo, E. B. 2001. Sexual differences of behavioral pat-
terns in emperor tamarins (Saguinus imperator subgrise-
scens): A case study for science communication in the
zoo community. In: European Srulb/ook for the Emperor
Tamarin Saguinus imperator ssp. 8th Edition, 2001, E. B.
Ruivo (compiler), pp.21-49. Lisbon Zoological Garden,
Lisboa, Portugal.



CONSERVATION INFORMATION SERVICE (CIS)

Primate-Science is pleased to announce the availability
of a new site devoted to primate conservation and the
management of primate habitats. The purpose of CIS
is to promote communication between primatologists
and funding sources. CIS will facilitate communication
between individuals seeking funding and organizations
or philanthropists that potentially could support their
work.

Individuals or groups whose projects relate to primate
conservation or the preservation of primate habitats
are invited to provide descriptions of their projects for
posting on the CIS site. These project descriptions will
be available to potential donors via the Internet. Because
it is important to understand primates in their ecological
setting, the emphasis of CIS is on maintaining primates
in their natural habitats, not on propagating endangered
species in captivity. To be listed, proposed projects must be
reviewed and endorsed by the Conservation Information
Service (CIS).

We hope that public access to a combination of
information and evaluation will encourage philanthropists
and foundations to increase their support at this critical
time when many primates face the threat of extinction.
We especially encourage postings from Non-Governmental
Organizations (NGO's), local organizations, and individuals
in developing countries where conservation efforts are
underway. We hope that facilitating direct contact between
donors and recipients will insure that funds are used most
efficiently.

Please visit the CIS web site where you will find instructions
for submitting proposals pin/cis/>. For questions or further information please
contact: Max Snodderly, CIS Steering Committee Chair,
Schepens Eye Research Institute and Harvard Medical
School, e-mail: . From: science@primate.wisc.edu>, 8 July, 2002.





Neotropical Primates 10(2), August 2002


RE-INTRODUCTION SPECIALIST GROUP

The Re-introduction Practitioners Directory, a registry
of reintroduction projects worldwide (both plants and
animals) was compiled and edited by Pritpal S. Soorae
and Philip J. Seddon in 1998 for the IUCN/SSC Re-
introduction Specialist Group (RSG) in collaboration
with the National Commission for Wildlife Conservation
and Development, Riyadh, Saudi Arabia (ISBN 9960
614 08 5, 97pp.). It is now on the IUCN website
and can be accessed from the Members
interest page:
or directly from: reintrddirectl998.pdf>. The project profiles for New
World primates include ( .-'-- ..r ..- Leontopithecus
rosalia, L. chrysopygus, Alouatta r,,//oI.r (Alouatta pigra) and
Brachyteles arachnoides.

Pritpal S. Soorae, Programme Officer, IUCN/SSC Re-
introduction Specialist Group (RSG), Environmental
Research & Wildlife Development Agency (ERWDA), P.O.
Box 45553, Abu Dhabi, United Arab Emirates (UAE), Fax:
(971) 2 681-7361, e-mail: .



THE LINCOLN PARK Zoo NEOTROPIC FUND

The Lincoln Park Zoo Neotropic and Africa/Asia Funds
support field research in conservation biology around the
world. The Neotropic fund focuses on projects undertaken
in Latin America and the Caribbean. Since 1986, the
fund has awarded over 146 grants in 19 countries. The
Africa/Asia fund, launched in 1997, focuses on projects
throughout Africa, Asia, and the Pacific. The funds
emphasize 1) the support of graduate students and other
young researchers, 2) direct impact on wildlife conservation
and/or conservation biology, 3) involvement by students
and/or local field assistants from Latin America, Africa,
or Asia at levels that engender appreciation for wildlife
conservation, and 4) links to either the Lincoln Park Zoo
animal collection or conservation activities of the zoo staff.
Each fund typically supports between five and ten projects
annually, including project renewals for a second year.
Most awards fall into the range of $3,000-$6,000. Initial
support is for up to 12 months from the date of award,
and the maximum duration of support is two years. The
current deadline for receipt of Neotropic and Africa/Asia
proposals is October 1st. For additional information
and application procedures go to conservation>, e-mail: , or
write to: Lincoln Park Zoo NF/AA Funds, Department
of Conservation and Science, Lincoln Park Zoo, 2001 N.
Clark St, Chicago, IL 60614.


2003 IPS MARTHA J. GALANTE AWARD CALL
FOR APPLICATIONS

SThe Martha J. Galante Award is
given annually by the International
Primatological Society (IPS) to professional
primatologists of primate habitat countries
to support conservation training. The IPS Conservation
Committee, chaired by Claudio Valladares-Padua, is
soliciting applications for the 2003 Award. The deadline
for applications is 1 May 2003. The award can be used for
conservation training: Transportation to the course or event
location, course or event fees, and expenses during the event
period. People interested in receiving this award should:

* Be officially enrolled in an academic institution or a
similar organization (either taking or giving courses or
doing research or conservation work),
* send information about the program of interest (courses,
congresses, symposia, field work, etc.),
* send a letter explaining his/her interests in participating in
the course or event (in English),
* send a CV in English,
* send a letter of acceptance for the respective course, and
* send two recommendation letters (including information
about referee).

The review of the application will be made by the
Conservation Committee, and the results will be
announced in August 2003. Grant proposals can be sent
by post or e-mail to: Dr. Claudio Valladares-Padua, IPS
Conservation Committee, IPL Instituto de Projetos e
Pesquisas Ecol6gicas, Caixa Postal 47, 12960-000 Nazard
Paulista, S~o Paulo, Brazil, E-mail: .







MARTIN TALES

Recently published was Number 6 (2002) of Tamarin
Tales, the newsletter of the International Committee
for the Conservation and Management of the Lion
Tamarins (ICCM). It is distributed to all institutions
which hold lion tamarins and participate in the captive
breeding programs. The first article, "Status of the Lion
Tamarins in the Wild", gives the current estimates for
the populations sizes of the four species: The golden lion
tamarin, Leontopithecus rosalia- 1000; the golden headed
lion tamarin, L. chrysomelas- 6,000-15,500; the black lion
tamarin, L. chrysopygus 1000; and the black-faced lion
tamarin, L. caissara- as few as 400. There follows a graphic
illustration of the loss of forest in Sao Paulo, once covering





Neotropical Primates 10(2), August 2002


about 80% of the state, and now reduced to about 3%!
The third article discusses the problems of the fragmented
habitats for the species and the establishment of the
metatopopulation management programs for the golden
and the black lion tamarins, including a breakdown of the
current population of the former: about 220 individuals
in the Pogo das Antas Biological reserve, 140 resulting
from animals translocated to the Uniao Biological Reserve,
about 400 in several separate populations resulting from
the reintroduction program, and about 250 animals
elsewhere, including the foothills of the Serra dos Orgaos.
Paula Proc6pio de Oliveira, who will defend her doctoral
thesis this year (Federal University of Minas Gerais, under
the supervision of Gustavo A. B. da Fonseca) reports on
her studies of golden lion tamarin ecology and the progress
of the translocated population in the Uniao Biological
Reserve. She also reports on future activities of the
program, which include the organization of a data base by
Vanessa Puerta Veruli, with all the information collected
during the translocation program since 1997 and, with
Fabiana Godoy and Leonardo Vieira of the Information
System Laboratory of the Associacao Mico-Leao-Dourado
(AMLD), the mapping of the trail system using satellite
images and GPS. Mariana Janzantti Lapenta will be
carrying out studies of seed dispersal in the reserve for a
doctoral thesis at the University of Sao Paulo.

Kristel De Vleeschouwer, postdoctoral researcher at
the Center for Research and Conservation of the Royal
Zoological Society of Antwerp, provides a short story of
her first impressions of the west side of the Una Biological
Reserve where she will be carrying out a field study
examining particularly the adaptations of golden-headed
lion tamarins to degraded and fragmented forests. Patricia
Matsuo also reports on the activities of the Conservation
Education Program for the golden lion tamarin, run by
the AMLD. They include teacher training for schools
in the vicinity of the reserves where lion tamarins have
been introduced, with numerous workshops being held
in the Adelmar F Coimbra-Filho Educational Center in
the Pogo das Antas Biological Reserve. Alex Howes of
the Friends of the National Zoo writes on some curious
aspects of tool use by free-ranging golden lion tamarins
in the National Zoo, Washington, DC, Zoo Atlanta
and Zoo Oregon, even though nothing of the sort has
been observed in the wild. Finally there are reports on
the retirement of the Jersey Zoo's long-time Director,
Jeremy J. C. Mallinson, effulgent warrior of lion tamarin
conservation, and the activities of the Lion Tamarins of
Brazil Fund (see page 101).

The newsletter is edited by Jonathan D. Ballou,
Department of Conservation Biology, National
Zoological Park, Smithsonian Institution, Washington,
DC 20008, USA, e-mail: . Please
send to him news and short articles concerning lion
tamarins and their conservation and biology in the wild for
the next issue.


SPECIAL ISSUE OF BEHAVIOR SOCIAL BONDS IN
PRIMATES

Volume 139, numbers 2-3 (February/March 2002) of
Behaviour- an International Journal of Behavioural Biology
(Brill, Leiden) is a special issue dedicated to the theme
"What are Friends For? The Adaptive Value of Social
Bonds in Primate Groups". The Guest Editor was Joan B.
Silk (University of California, Los Angeles), and the papers
result from a symposium held during the XVIIIth Congress
of the International Primatological Society, in Adelaide,
Australia, 7-12 January 2001. Contents. Introduction -
J. B. Silk; Ecological models of female social relationships
in primates: Similarities, disparities, and some directions for
future clarity L. A. Isbell & T. P. Young; How adaptive
or phylogenetically inert is primate social behaviour?
A test with two sympatric colobines A. H. Korstjens,
E. H. M. Sterck & R. Noe; An expanded test of the
ecological model of primate social evolution: Competitive
regimes and female bonding in three species of squirrel
monkeys (Saimiri oerstedii, S. boliviensis and S. sciureus) S.
Boinski, K. Sughrue, L. Selvaggi, R. Quatrone, M. Henry
& S. Cropp; Constraints on relationship formation among
female primates L. Barrett & S. P. Henzi; Friendship
among adult female blue monkeys (Cercopithecus mitis)
- M. Cords; Social dynamics of male muriquis (Brachyteles
arachnoides hypoxanthus) K. B. Strier, L. T. Dib & J.
E. C. Figueira; Reciprocity and interchange in the social
relationships of wild male chimpanzees D. P. Watts;
Affiliation and aggression among adult female rhesus
macaques: A genetic analysis of paternal cohorts A.
Widdig, P. Ntirnburg, M. Krawczak, W. J. Streich & E
Bercovitch; Relationship assessment through emotional
mediation F Aureli & C. M. Schaffner; Using the 'F'
word in primatology J. B. Silk.



AMAZONIANA A SPECIAL EDITION DEDICATED TO
HARALD SIOLI

Volume 16(3/4), December 2001, of the journal Ama-
zoniana, is dedicated to Prof. Dr. Harald Sioli, pioneer
researcher, limnologist and conservationist of the Amazon,
who enjoyed his 90th birthday in August 2000. His first
studies began in 1945, and from the 1960s he collaborated
closely with the National Institute for Amazon Research
(INPA), Manaus. In 1965, he co-founded Amazoniana
with Djalma Batista, then Director of INPA, and in 1966
he was appointed director of the Max-Planck Institute for
Limnology in Plan, Germany, and director of its newly
formed Department for Tropical Ecology. He retired in
1978. In a brief appraisal of his work, Wolfgang Junk (cur-
rent director of the Department of Tropical Ecology) wrote
the following: "The impact of Sioli's activities reaches far
beyond the impact of his scientific publications. His very
early warnings about the destruction of the Amazon rain
forest and his appeals to protect the area for the benefit of





Neotropical Primates 10(2), August 2002


nature, and the local populations including the Amerin-
dian tribes, were heavily criticized by some politicians and
development planners, but received endorsement from
Brazilian scientists and were enthusiastically accepted by
Brazilian students. This development fortified the ideas for
the need for environmental protection in Brazil. Some of
the students, influenced by his ideas during the sixties and
seventies, are today leading scientists and administrators
in Brazilian state and governmental organizations, and
introduce ecological aspects into politics, planning and
administration. Today Harald Sioli is Nestor of German
tropical ecology and one of the great tropical ecologists in
the world."(p.268).

Nothing on monkeys, but this edition of Amazoniana
(only Part 1 of the dedication to Prof. Sioli) is replete
with excellent review papers on Amazonian geology, geo-
morphology, paleohistory, geography, limnology, ecology
and biodiversity, and includes three important reviews of
the Amazonian refuge theory, and overviews of Amazon
deforestation and development. A sample of the contents:
Appraisal of the scientific work of Harald Sioli W.
J. Junk, pp.285-297; Birthday letter to Harald Sioli -
L. Schmidt, pp.299-301; The prehistoric human geogra-
phy of Brazil A. N. Ab'Saber, pp.303-311; Holy Ganga
and the mighty Amazon B. Gopal, pp.337-348; Amazo-
nia 2000: An evaluation of three decades of regional plan-
ning and development programs in the Brazilian Amazon
region G. Kohlepp, pp.363-395; Mangal communities
of the "Salgado Paraense": Ecological heterogeneity along
the Braganga peninsula assessed through soil and leaf
analysis E. Medina et al., pp.397-416; The mystery of
the Marajoara: An ecological solution B. J. Meggers,
pp.412-440; Comparative study of the litterfall in a pri-
mary and secondary terra firme forest in the vicinity of
Manaus, State of Amazonas, Brazil W. A. Rodrigues, K.
Furch & H. Klinge, pp.441-462; Amazonian deforesta-
tion: Regional and global issues E. Salati, C. A. Nobre
& A. A. dos Santos, pp.463-481; Land use dynamics in
the Amazon estuary and implications for natural resource
management N. J. H. Smith, pp. 517-537; Climatic
forcing of evolution in Amazonia during the Cenozoic:
On the refuge theory of biotic differentiation J. Haffer
& G. T. Prance, pp.579-607; A paradigm to be discarded:
Geological and paleoecological data falsify the Haffer and
Prance refuge hypothesis of Amazonian speciation P. A.
Colinvaux, G. Irion, M. E. Rdisanen, M. B. Bush & J. A.
S. Nunes de Mello, pp. 609-646; Ice age tropical South
America: What was it really like? T. van der Hammen,
pp.647-652.

Amazoniana is edited by Wolfgang Junk and Joachim
Adis (Max Planck Institute for Tropical Limnology,
Plan), Francisco de Assis Esteves (Federal University of
Rio de Janeiro) and Ulrich Saint-Paul (Bremen). ISSN
0065-6755. For subscriptions or single issues, contact. Kom-
missionsverlag Walter G. Miihlau, Holtenauer Str. 116,
D-24105 Kiel, Germany, Fax: +49 431 800 9050, e-mail:
.


GUIDELINES FOR NONHUMAN PRIMATE
RE-INTRODUCTIONS

The IUCN/SSC Re-introduction Specialist Group (RSG)
(Chair, Frederic J. Launay) was established in 1988 in
response to an increasing number of plant and animal
re-introductions worldwide. The IUCN Guidelines for Re-
introductions, published in 1998, covers key issues and is a
general policy document that applies to both animals and
plants. In June 2002, the RSG published a special issue of
Re-introduction NEWS dedicated to primates. In addition
to seven short case studies on primate re-introductions and
translocations, the special issue features a 29-page docu-
ment: Guidelines for Nonhuman Primate Re-introductions.

Edited by Lynne R. Baker, the guidelines are based on
current IUCN policy documents, a review of case histories,
and consultation across a broad range of disciplines.
Comments were solicited from a large group of experts
and interested parties, and a thorough review was carried
out by a Core Review Board with the following members:
Benjamin Beck (National Zoological Park, Smithsonian
Institution, Washington, DC), Thomas M. Butynski
(Africa Section, IUCN/SSC Primate Specialist Group
and Eastern Africa Biodiversity Hotspots, Conservation
International, Washington, DC), Ardith Eudey (Asian
Section, IUCN/SSC Primate Specialist Group), Elizabeth
L. Gadsby (Pandrillus, Drill Rehabilitation and Breeding
Center, Nigeria), Kenneth Glander (Duke University,
Durham, NC), William B. Karesh (Wildlife Conservation
Society, New York, NY), Devra G. Kleiman (National
Zoological Park, Smithsonian Institution, Washington,
DC), John Lewis (International Zoo Veterinary Group,
London), Russell A. Mittermeier (Chair, IUCN/SSC
Primate Specialist Group and Conservation International,
Washington, DC), John E Oates (Hunter College,
City University of New York), Anthony B. Rylands
(Neotropical Section, IUCN/SSC Primate Specialist
Group, and Center for Applied Biodiversity Science,
Conservation International, Washington, DC), Pritpal
S. Soorae (Senior Conservation Officer, IUCN/SSC Re-
introduction Specialist Group, Abu Dhabi, UAE), Shirley
S. Strum (University of California, San Diego), Caroline
Tutin (Centre International de Recherches Medicales de
Franceville, Gabon, and University of Stirling, Scotland),
Michael Woodford (Working Group on Wildlife Diseases,
World Organisation for Animal Health, Office International
des Epizooties, Paris).

The guidelines are divided into 11 sections: Executive
Summary; Context of Guidelines; Introduction; Definition
of Terms; Aims, Objectives, and Precautionary Principle;
Planning for Re-introduction; Disease Transmission
and Veterinary Requirements; Transport and Release
Implementation; Post-Release Monitoring; Considerations
for Translocation; and Acknowledgments. Four annexes
follow: Key References, Husbandry References, Key
Contacts, and Core Review Board. "Key Contacts"





Neotropical Primates 10(2), August 2002


includes a useful list of the Primate Taxon Advisory Groups
worldwide. The following is the Executive Summary:

"The IUCN/SSCRe-introduction Specialist Group: Guidelines
for Nonhuman Primate Re-introductions is intended as
a guide to re-introduction programs. The priority has
been to develop standards that are of direct, practical
assistance to those planning, approving, or implementing
re-introductions. The primary audience of these guidelines
is the re-introduction practitioner.

Because re-introduction projects are often restricted by
location, resources, and other limitations, this document
is a meant as a "best-practice" model, or an ideal code
of conduct. Re-introduction managers are strongly
encouraged to use this document as their principal guide to
primate re-introductions.

Each re-introduction project should develop written
guidelines that apply specifically to its taxon, region, legal
structure, etc. These customized documents should be up-
dated over time and eventually result in a re-introduction
manual for the taxon of interest. They should also directly
relate to this existing document, so that these guidelines
can be regularly updated with new information and
techniques.

Guidelines for Nonhuman Primate Re-introductions covers the
main steps of a re-introduction effort. The steps are listed
in a suggested order of execution, although some overlap
with one another. It is realized that many projects have been
operating for some time, so their managers should attempt
to integrate the guidelines as soon as possible into their
current operating procedures and protocol.

Before initiating any re-introduction project, managers
must clearly define why that project is needed and do a
rapid overall assessment to ensure that key requirements,
such as habitat suitability, are likely to be met. The main
goal of any re-introduction effort should be to re-establish
S- r populations of primates in the wild and
to maintain the viability of those populations. Although
exceptions to this, such as trial re-introductions of
common species and rescue/welfare releases, should also
adhere to these guidelines as much as possible, such
projects are not considered true re-introductions or
conservation approaches and are not specifically covered
in these guidelines.

Re-introduction practitioners are strongly encouraged to
contact the IUCN/SSC Re-introduction Specialist Group
(RSG) and present and discuss their re-introduction
proposals and results. As a result, a network of contacts can
be developed and information from various projects shared.

Details regarding the care of animals held in captivity prior
to release, such as enclosure enrichment, are not specifically
covered in these guidelines. However, where appropriate,
important points regarding these topics will be noted."


The special primate issue of Re-introduction NEWS includes
an introduction by the RSG Primate Section Chair, Devra G.
Kleiman, who discussed some divergence in the definitions
of the key terms used when compared to the 1998 IUCN
Guidelines for Re-introductions. She also reviewed the articles
in the special issue and emphasized that the RSG provides
guidance to those planning to re-introduce or translocate
animals or plants mainly for conservation purposes and that,
for a program to have significant conservation value, it must
result in a significant increase in the numbers and genetic
diversity of a threatened or endangered species in protected
habitat, preferably within the species' historic range.

The articles in the special issue are as follows: Re-
introduction and translocation as conservation tools
for golden lion tamarins in Brazil M. C. M. Kierulff,
B. B. Beck, D. G. Kleiman, & P. Proc6pio de Oliveira,
pp.7-10; Translocation of black howler monkeys in Belize
- R. H. Horwich, F Koontz, E. Saqui, L. Ostro, S. Silver,
& K. Glander, pp.10-12; Translocation of three wild
troops of baboons in Kenya S. C. Strum, pp.12-15;
Habitat ecologique et liberty des primates: A case study of
chimpanzee re-introduction in the Republic of Congo -
K. H. Farmer & A. Jamart, pp. 16-18; The release of captive-
bred black-and-white ruffed lemurs into the Betampona
Reserve, eastern Madagascar A. Britt, C. Welch, &
A. Katz, pp.18-20; Preparing for re-introduction: 10 years
of planning for drills in Nigeria E. L. Gadsby, pp.20-
23; Re-introduction of orang-utans in Indonesia K. S.
Warren & R. A. Swan, pp.24-26; Release of golden langurs
in Tripura, India- A. K. Gupta, pp.26-28.

The special primate issue was supported by the
American Society of Primatologists (ASP), International
Primatological Society (IPS), the Margot Marsh Biodiversity
Foundation, and the Primate Society of Japan. This issue of
Re-introduction NEWS was edited by Pritpal Soorae and
Lynne R. Baker.

To contact the RSG: Pritpal Soorae, Executive Officer,
IUCN\SSC Re-introduction Specialist Group Secretariat,
c/o Environmental Research & Wildlife Development
Agency (ERWDA), P.O. Box 45553, Abu Dhabi, United
Arab Emirates (UAE): Tel: (971) 2 693-4650, Fax: (971)
2 681-7361, E-mail: . The
guidelines are available on the RSG Website at http://iucn.
org/themes/ssc/programs/rsg.htm.

Lynne R. Baker, Conservation Biology Department,
University of Minnesota, 180 McNeal Hall, 1985
Buford Avenue, St. Paul, MN 55108, USA, e-mail:
, Devra G. Kleiman, RSG
Primate Section Chair, Research Associate, National
Zoological Park, Washington, DC 20008, USA, e-mail:
, and Pritpal Soorae, IUCN/
SSC Re-introduction Specialist Group Secretariat, c/o
Environmental Research & Wildlife Development Agency
(ERWDA), P.O. Box 45553, Abu Dhabi, United Arab
Emirates (UAE), e-mail: .





Neotropical Primates 10(2), August 2002


References

Baker, L. R. (ed.). 2002. IUCN/SSC Re-introduction
Specialist Group: Guidelines for Nonhuman Primate Re-
introductions. Re-introduction News (21): 29-57. Website:
.
IUCN. 1998. IUCN Guidelines for Re-introductions. Pre-
pared by the IUCN/SSC Re-introduction Specialist
Group. The World Conservation Union (IUCN), Gland,
Switzerland. Website: policy/reinte.htm>.
Soorae, P. S. and Baker, L. R. (eds.). 2002. Re-introduction
NEWS: Special Primate Issue, Newsletter of the IUCN/SSC
Re-introduction Specialist Group, Abu Dhabi, UAE. (21):
60pp. (ISSN 1560-3709).



NEW PUBLICATIONS REGARDING REINTRODUCTION
AND THE PLACEMENT OF CONFISCATED ANIMALS

The IUCN Guidelines for the Placement of Confiscated
Animals (2002) were developed after an extensive review
process, and became official IUCN Policy at the 51"t
Meeting of the IUCN Council in February, 2000. The
guidelines are designed to provide options for the place-
ment of animals confiscated from illegal or irregular trade.
The Environmental Research and Wildlife Development
Agency (EWRDA) supported the publication. Copies
(limited) are available from: IUCN/SSC Re-introduction
Specialist Group Secretariat, c/o Environmental Research
& Wildlife Development Agency (ERWDA), P.O. Box
45553, Abu Dhabi, United Arab Emirates (UAE): Tel:
(971) 2 693-4650, Fax: (971) 2 681-7361, e-mail:
. Website: intranet/DocLib/Docs/IUCN735.pdf>.

The booklet Quarantine and Health Screening Protocols
for '1 j.-- Prior to Translocation and Release into the Wild
(2002) was edited by Michael H. Woodford. It describes
many of the disease risks attending wildlife translocation
projects. Suggestions are made for the development of sys-
tematic procedures for the reduction of these risks at both
the source of the founder animals and at the release site, and
covers the following taxa: Artiodactyla, Perissodactyla, Pri-
mates, Carnivora, Marine mammals, Rodentia, Lagomor-
pha, marsupials (New and Old World), Monotremata, Chi-
roptera, Aves, Reptilia and Amphibia, and fishes. Available
from: Care for the Wild International, Ashfolds, Rusper,
West Sussex RH12 4QX, UK, Tel: +44 1293 871596, Fax;
+44 1293 8715022, e-mail: .



BOOKS

Diversidade Bioldgica e Cultural da Amazonia, edited by
Ima Cdlia Guimaraes Vieira, Josd Maria Cardoso da Silva,
David Conway Oren and Maria Angela D'Incao. 2001.


Museu Paraense Emilio Goeldi, Belkm, Para, Brazil. 421pp.
ISBN 85 7098 067 1. Price $25.00 (+ US$5.00 p&p
outside of Brazil). In English and Portuguese. The results
of a symposium celebrating the 130' anniversary of the
Museu Paraense Emilio Goeldi, Belkm, 23-27 October
1996 "The Biological and Cultural Diversity of Amazonia
in a World of Transformation". The book covers three basic
questions: What is the origin of Amazonian biodiversity?;
What is the origin of the region's cultural diversity?; and
How to promote the sustainable use of biodiversity in
the Amazon? Contents. Part I. Origin of Biodiversity in
Amazonia. The Amazonian rainforest only some 6-5 million
years old N.-A. Mbrner, D. Rosetti & P. M. de Toledo,
pp.3-18; Paleoecology of Amazonia T. Van der Hammen,
pp.19-44; Hypotheses to explain the origin of species in
Amazonia J. Haffer, pp.45-118; Avian diversification
in Amazonia: evidence for historical complexity and
a vicariance model for a basic diversification pattern
- J. Bates, pp.119-137; Molecular phylogenetics and the
diversification of Amazonian mammals J. Patton & M.
N. F da Silva, pp.139-164. Part II. Human and Cultural
Diversity. Diversidade genetica de populao6es humans na
Amazonia. D. de F Lobato da Silva, A. K. C. Ribeiro dos
Santos & S. E. Batista dos Santos, pp.167-193; Amazonia
socioambiental sustentabilidade ecol6gica e diversidade
social D. Lima & J. Pozzobon, pp.195-251; Um aspect
da diversidade cultural do caboclo R. H. Mauns, pp.253-
272; Science and the representation of nature in Amazonia:
from La Condamine through Da Cunha to Anna Roosevelt
- D. Cleary, pp.273-296. Part III. Sustainable Use of
Biodiversity in Amazonia. As ciencias, o uso de recursos
naturais na Amazonia e a nocao de desenvolvimento
sustentavel: por uma interdisciplinaridade ampla E de
Assis Costa, pp.299-318; Natural vs. social science concepts
in applied research on Amazonia: a critical assessment M.
Nitsch, pp.319-346; Domestication of Amazonian fruit
crops past, present, future C. R. Clement, pp.347-
367; Dinamica evolutiva em royas de caboclos amaz6nicos
- P. S. Martins, pp.369-384; Influence of habitat on the
sustainability of mammal harvests in the Peruvian Amazon
- R. Bodmer, P. Puertas, R. Aquino & C. Reyes, pp.385-
402; Biodiversity: today's and tomorrow's importance W
Kerr, pp.403-409. Part IV. Tributes. La Penha: gerador e
gerenciador de ciencia L. M. E Bassalo, pp.413-416;
Paulo Sodero: mestre por excelencia I. C. G. Vieira,
p.417; Jorge Pozzobon, agora no cdu com diamantes, M.
Meira, pp.419-421. Available from: Biblioteca, Museu
Paraense Emilio Goeldi, Caixa Postal 399, 66040-170
Belkm, Para, Brazil. Website: .

A Guide to Careers in PhysicalA. .-' ..... .. edited by Alan
S. Ryan. 2002. Bergin and Garvey. Price: $69.95. ISBN 0-
89789-693-9. Physical anthropology focuses on biological
evolution of humans and their ancestors, the relationship
of humans to other organisms, and patterns of biological
variation within and among human populations. Physical
anthropology is sometimes referred to by another name
- biological anthropology. There are several specialties of
physical anthropology including primate studies, paleo-





Neotropical Primates 10(2), August 2002


anthropology, and human variation. Because of its broad
scope, physical anthropology has borrowed principles
from evolutionary biology, comparative anatomy, genetics,
medicine, paleontology, zoology, geology, and demography.
The subject of this book is careers in physical anthropol-
ogy. Most physical anthropology graduate students have
traditionally aspired to a career as a college or university
faculty member in an anthropology department. Because
physical anthropology has a strong biocultural emphasis
and its subject matter is enormously diversified, today's
students of physical anthropology have a wealth of potential
nontraditional career opportunities. Contents: Introduction
- A. S. Ryan; 1. The meaning of physical anthropology -
A. S. Ryan; 2. Teaching physical anthropology in a univer-
sity: The traditional career C. W Wienker; 3. Teaching
physical anthropology in the community college P. L.
Stein; 4. The practice of physical anthropology in a museum
environment D. H. Ubelaker; 5. Paleoanthropology at
home and in the field A. Kramer; 6. Primatology as a
career K. D. Hunt; 7. The post-doc experience: Is there a
light at the end of the tunnel? A. C. Stone; 8. Krogman, his
cleft palate collection, and me: or, what can an auxologist do
today? E, J. Bowers-Bienkowski; 9. Teaching anatomy at a
university M. F Teaford; 10. Research faculty in medical,
nursing, and public health schools; S. T. McGarvey & G.
D. James; 11. Physical anthropology, medical genetics, and
research B. B. Little; 12. Opportunities in public health
and international nutrition R. Martorell; 13. Having fun
- A jock in two worlds: Kinesiology and human biology R.
M. Malina; 14. Government research: links to biomedicine
and public health R. M. Garruto; 15. Private industry: Re-
search for profit A. S. Ryan; 16. Independent consulting:
Making your own rules M. R. London; 17. Journalism:
Bringing science to the public K. Wong; 18. Forensic sci-
ence as a new arena for a human biologist M. S. Schanfield.
Where to order Greenwood Publishing Group, Inc., 88 Post
Road West, Westport, CT 06881, USA, Tel: 800-225-5800,
Fax: 203-750-9790. Website: .



ARTICLES

Bales, K., French, J. A. and Dietz, J. M. 2002. Explaining
variation in maternal care in a cooperatively breeding
mammal. Anim. Behav. 63(3): 419-535.
Bardi, M. and Petto, A. J. 2002. Parental failure in captive
common marmosets ((C .- jacchus): A comparison
with tamarins. Folia Primatol. 73(1): 46-48.
Bicca-Marques, J. C., Garber, P A. and Azevedo Lopes, M.
A. 0. 2002. Evidence of three resident adult male group
members in a species of monogamous primate, the red titi
monkey (( .... cupreus). Mammalia 66(1): 138-142.
Custance, D. M., Whiten, A. and Fredman, T. 2002. Social
learning and primate reintroduction. Int. J. Primatol.
23(3): 479-499.
DeJoseph, M., Taylor R. S. L., Baker, M. and Aregullin, M.
2002. Fur-rubbing behavior of capuchin monkeys. J. Am.
Acad. Dermatol. 46(6): 924-925.


Dixson, A, and Amderson, M. 2002. Sexual selection and
the comparative anatomy of reproduction in monkeys,
apes and human beings. Ann. Rev. Sex Res. 12: 121-144.
Giudice, A. M. 2002. Analisis del comportamiento de
Cebus ,//da en jardins zool6gicos. Boletin de laAsociacidn
Primatoldgica Espanola 9 (2): 20-21.
Goodman, S. and Check, E. 2002. The great primate
debate. Nature, Lond. 417: 684-687.
Harcourt, A. H., Coppeto, S. A. and Parks, S. A. 2002.
Rarity, specialization and extinction in primates.
J. Biogeog. 29(4): 445-456.
Hernandez-L6pez, L., Parra, G. C., Cerda-Molina, A. L.,
Prez-Bolafios, S. C., Sanchez, V. D. and Mondragon-
Ceballos, R. 2002. Sperm quality differences between
the rainy and dry seasons in captive black-handed spider
monkeys (Ateles, ..rr .., 1 Am. J. Primatol. 57(1): 35-41.
Hernandez-Lopez, L., Parra, G. C., Cerda-Molina, A. L.,
Prez-Bolafios, S. C. and Mondrag6n-Ceballos, R. 2002.
Digestion by trypsin enhances assessment of sperm pa-
rameters in the black-handed spider monkey (Ateles geof
froyi). Lab. Prim. Newsl. 41(3): 4-6.
Hladik, C. M., Pasquet, P. and Simmen, B. 2002.
New perspectives on taste and primate evolution:
The dichotomy in gustatory coding for perception of
beneficent versus noxious substances as supported by
correlations among human thresholds. Am. J. Phys.
Anthropol. 117(4): 342-348.
Jones, C. B. 2002. How important are urinary signals in
Alouatta? Lab. Prim. Newsl. 41(3): 15-17.
Kostan, K. M. and Snowdon, C. T. 2002. Attachment
and social preferences in cooperatively-reared cotton-top
tamarins. Am. J. Primatol. 57(3): 131-139.
Kralik, J. D. and Hauser, M. D. 2002. A nonhuman
primate's perception of object relations: experiments
on cottontop tamarins, Saguinus oedipus. Anim. Behav.
63(3): 419-535.
Lawler, R. R. and Stamps, C. 2002. The relationship
between tail use and positional behavior in Alouatta
palliata. Primates 43(2): 147-152.
Lynch, J. W., Ziegler, T. E. and Strier, K. B. 2002. Individual
and seasonal variation in fecal testosterone and cortisol
levels of wild male tufted capuchin monkeys, Cebus j,e//.
nigritus. Horm. Behav. 41(3): 275-287.
Montoya, G. E., Vernot, J.-P. and Patarroyo, M. E. 2002.
Partial characterization of the CD45 phosphatase cDNA
in the owl monkey (Aotus vociferans). Am. J. Primatol.
57(1): 1-11.
Moorman, E. A., Mendoza, S. P., Shideler, S. E. and Lasley,
B. L. Excretion and measurement of estradiol and pro-
gesterone metabolites in the feces and urine of female
squirrel monkeys (Saimiri sciureus). Am. J. Primatol.
57(2): 79-90.
Price, E. C., Piedade, H. M. and Wormell, D. 2002. Popu-
lation densities of primates in a Brazilian Atlantic forest.
Folia Primatol. 73(1): 54-56.
Prieto, 0. H., Santa Cruz, A. M., Scheibler, N., Borda, J.
T. and G6mez, L. G. 2002. Incidence and external mor-
phology of the nematode Trypanoxyuris (Hapaloxyuris)
callithricis, isolated from black-and-gold howler monkeys





Neotropical Primates 10(2), August 2002


(Alouatta caraya) in Corrientes, Argentina. Lab. Prim.
Newsl. 41(3): 12-14.
Rasmussen, D. R., Chapin, B. L. and Chambers, C. M.
2002. Mantled howlers cause a decrease in distance
between members of a group of rufous-naped tamarins:
A field experiment. Lab. Prim. Newsl. 41(2): 10-14.
Sanchez, S., Pelaez, F and Gil-Btirmann, C. 2002. Why do
cotton-top tamarin female helpers carry infants? A pre-
liminary study. Am. J. Primatol. 57(1): 43-49.
Sechrest, W., Brooks, T. M., Fonseca, G. A. B. da, Konstant,
W. R., Mittermeier, R. A., Purvis, A., Rylands, A. B. and
Gittleman, J. L. 2002. Hotspots and the conservation
of evolutionary history. Proc. Natl. Acad. Sci. 99(4):
2067-2071.
Serio-Silva, J. C. and Rico-Gray, V. 2002. Influence of mi-
croclimate at different canopy heights on the germination
of Ficus (urostigma) seeds dispersed by Mexican howler
monkeys (Alouatta palliata mexicana). Interciencia 27(4):
186-190.
Silk, J. B. 2002. Females, food, family and friendship. Evol.
Anthropol. 11(3): 85-87.
Soltis, J., Bernhards, D., Donkin, H. and Newman, J.
D. 2002. Squirrel monkey chuck call: Vocal response
to playback chucks based on acoustic structure and
affiliative relationship with the caller. Am. J. Primatol.
57(3): 119-130.
Spinozzi, G. and Truppa, V. 2002. Problem-solving and
hand preferences for a multicomponent task by tufted
capuchins (Cebus q",,/'/). Int. J. Primatol. 23(3): 621-
638.
Tsujimoto, S. and Sawaguchi, T. 2002. Working memory
of action: A comparative study of ability to selecting
response based on previous action in New World
monkeys (Saimiri sciureus and C .- jacchus). Behav.
Processes 58(3): 149-155.
Urioste, J. A. de and Bethencourt, M. J. 2002. Fundaci6n
Neotr6pico. Boletin de la Asociacidn Primatoldgica
Espanola 9(2): 3-5.



ABSTRACTS

Dew, J. L. 2002. Synecology and seed dispersal in woolly
monkeys (Lagothrix lagothricha poeppigii) and spider
monkeys (Ateles belzebuth belzebuth) in Parque Nacional
Yasuni, Ecuador. Diss. Abst. Int. B62(7): 3041.
Di Bitteti, M. S. 2002. Food associated calls in the tufted
capuchin monkey (Cebus ,//cl.). Diss. Abst. Int. B62(9):
3883.
Reeder, D. M. 2002. The biology of parenting in the
monogamous titi monkey (C .... moloch). Diss. Abst.
Int. B62(9): 4263.
Rodriguez, S. M. 2002. Analisis demografico, conductual y
gen&tico del mono aullador (Alouattapalliata mexicana) y
mono arafia (Ateles ...rr .., vellerosus) en un paisaje frag-
mentado del sur de Veracruz, Mexico. Lab. Prim. Newsl.
41(2): 3.


Selected abstracts from Anthropological Science 110(1),
2002.

Izawa, K. 2002. The mobbing call and long call of wild
spider monkeys. II, p.89.
Kobayashi, S., Natori, M., Pessoa, L., Oliveira, J., Langguth,
A. and Setoguchi, T. Taxonomic status of tufted capuchin
monkey (Cebus ,I,/li) in adjacent area of the Atlantic
rain forest. III. Analyses of nonmetric characters on the
skull, p.96.
Kondo, S., Natori, M. and Hanamura, H. Odontometrical
study of the deciduous and permanent molars in the
squirrel monkey (Saimiri), p.69.
Kuwahata, H., Kuroshima, H., Anderson, J. R. and Fujita,
K. Do monkeys prefer regular visual patterns? p. 113.
Nakano, Y., Hirasaki, E., Oka, K., Hirokawa, Y. and
Kumakura, H. The change of the primate locomotion on
the inclined substrata, p.75.
Nishimura, A. Home range of woolly monkeys (Lagothrix
lagotricha) at La Macarena, Colombia: Seasonal change
and a comparison with sympatric spider monkeys (Ateles
belzebuth), p.126.
Saito, A., Ueno, Y., Kawamura, S. and Hasegawa, T. Food
search behavior of trichromatic and dichromatic capuchin
monkeys (Cebus ,,//l"'), p.134.
Shimooka, Y. Intraday grouping pattern of wild spider
monkeys, p.89.
Takenaka, N., Hirai, M. and Kawamura, S. Y-chromosomal
LWS/MWS visual pigment genes of owl monkey, p.81.
Tsujimoto, S. and Sawaguchi, T. 2002. A comparative
study of 'working memory of action' in New World mon-
keys, p.83.

Selected abstracts of the Spring Meeting 2002 of the
Primate Society of Great Britain, Oxford Brookes
University, Oxford, 15-16 April, 2002. In Primate Eye
(77), June 2002.

Bearder, S. Tackling primate conservation from the sharp
end and from the blunt end, p.3.
Brend, S. Primate sanctuaries: Animal welfare or
conservation? pp.3-4.
Brend, S. Primate Society of Great Britain Captive Care
Working Party, pp.13-14.
Day, R. Neophilia, innovation and social learning:
A study of intergeneric differences in callitrichid monkeys,
pp.4-5.
Feistner, A. Primate Society of Great Britain Conservation
Working Party, p.14.
Lee, P. Ethics, attitudes and futures for primates: Beyond
bushmeat, pp.5-6.
Semple, S., Cowlishaw, G. and Bennett, P. Immune system
evolution among anthropoid primates: Pathogens,
injuries and predators, p.10.
Smith, A., Buchanan-Smith, H., Surridge, A., Mundy, N.
and Osorio, D. The effect of colour vision phenotype on
the detection and selection of fruits by tamarins (Saguinus
spp.), p.11.





Neotropical Primates 10(2), August 2002


Stamer, C. and Bearder, S. An introduction to the MSc
in primate conservation at Oxford Brookes University,
p.17.

Selected abstracts from the 16th Annual Meeting of the
Society for Conservation Biology, 14-19 July 2002,
co-hosted by the Durrell Institute of Conservation and
Ecology (DICE), the University of Canterbury, Kent,
and the British Ecological Society. In the Programme
and Abstracts.

Alvarez-Romero, J. G. and Medellfn, R. A. Introduced
mammals of Mexico: Diversity, distribution, potential
impact and prioritisation for their control, p.A3.
Anderson, A. B. and Bickford, S. M. Engaging stakeholders
to implement biological corridors: International case
studies, p.A5.
Ayres, J. M. A new model for conserving biodiversity in
Central Amazonia, p.A5.
Bager, A. and Amaral, F. P. Analysis of a fauna protection
system implanted in federal protected areas in southern
Brazil, p.A6.
Barlow, J. and Peres, C. Forests, flames and feathers:
Effects of El Nifio mediated forest fires on Amazonian
understorey bird assemblages, p.A9.
Beck, B. B., Martins, A. E, Ballou, J. D. and Mickelberg,
J. Demographic evaluation of the golden lion tamarin
reintroduction program, p.A1 1.
Becker, C. D. Evolving value for biodiversity preservation
in rural Ecuador, pp.A11-12.
Bennett, E. L., Robinson, J. G. and Eves, H. The scale of
hunting and wild meat trade in tropical forests today,
p.A12.
Bodmer, R. and Puertas, P. Density dependent responses to
hunting in Amazonian mammals, pp.A15-16.
Bowen-Jones, E., Mew, J. and Valencia, L. Community
conservation through land purchase lessons from the
Awacachi Corridor Project, Ecuador, p.A17.
Brandon, K. Parks as islands: Ecological and social
implications of edges, p.A17.
Ceballos, G. Global patterns of mammal species diversity,
endemism and endangerment: Implications for
conservation, p.A23.
Christen, C. At home in the field: Development of
Smithsonian Tropical Field Stations in Panama, p.A25.
Dietz, J., Ballou, J. D. and Baker, A. J. Effects of intense
predation on the demography and genetic effective size of
isolated populations, p.A36.
Dietz, L. A., Aquino A. L., Di Bitetti, M. D., Placci, G.,
and Maltez, H. M. From vision to reality, implementing a
tri-national Atlantic forest biodiversity corridor, p.A36.
Dorsey, M. Tracking political economies and ecologies in
the upper Amazon basin, p.A37.
Ferraz, G., Russell, G. J., Stouffer, P. C., Bierregaard, R. 0.,
Pimm, S. L. and Lovejoy, T. E. Rate of species loss from
Amazonian forest fragments, p.A45.
Hanazaki, N. and Begossi, A. Terrestrial fauna used by
caizaras from the Atlantic forest coast, Brazil, p.A58.


Harcourt, A. H., Parks, S. A. and Coppeto, S. A. Human
influences on extinction: A global analysis, pA59.
Howell, C. A. Applications of coarse-filter theory to
Ecuador, p.A64.
Jenkins, C. and Anderson, A. B. Using conservation
priorities to design a biological corridor in the Atlantic
forest of Brazil, p.A66.
Kleiman, D. G. and Moosbruker, J. Structure and function
of FWS-AZA partnerships in Endangered Species
Recovery Programmes, p.A73.
Laurance, W F. Rapid erosion of plant biodiversity in
Amazonian forest fragments, p.A78.
Leme, A., Begossi, A. and Tamashiro, J. Plant and animal
use in medicine by caboclos from the middle Negro river,
Amazonia, Brazil, p.A79.
Mace, G. M. Making choices in species conservation
assessing threats and priorities, p.A85.
Mcaliley, L. R., Haynie, M. L., Brant, J. G., Phillips, C.
J., Jones, C. and Baker, R. J. Examination of a proposed
corridor in Ecuador: Does the Rio Pastaza restrict gene
flow?: A pilot study, p.A90.
Melnick, D. Species-based approaches to broader scale
conservation of genetic diversity, pp.A92-93.
Millington, A., Tanner, T., Bradley, A., Lazcano, J.-M.,
Espinoza, D., Leo, M., Casaretto, C., Vehkamaki, S. and
Beckmann, S. Compromised conservation? Evaluating
the success of montane forest national parks in Argentina,
Bolivia and Peru, p.A94.
Morsello, C. Indigenous peoples and market integration:
Paradise, hell or purgatory in Brazil? A97.
Nelson, K. C. Local response to global environmental
initiatives: Carbon mitigation projects in Chiapas,
Mexico, p.A102.
Otterstorm, S. M. and Schwartz, M. Is fire a necessary
disturbance in the tropical dry forests of Mesoamerica?
pp.A108-109.
Pearl, M., Stanley-Price, M. and Williams, E. Future
directions for species-based conservation in human-
modified landscapes, p.A1 12.
Peres, C. Spatial heterogeneity in game vertebrate biomass
and bushmeat hunting patterns in Amazonian forests,
p.A1 13.
Perez-Sweeney, B., Valladares-Padua, C. and Melnick, D.
J. Using genetics for black lion tamarin (Leontopithecus
chrysopygus) metapopulation management, p.A1 13.
Pressey, B., Watts, M. and Barrett, T. Priority conservation
areas: Testing alternative approaches with simulations of
future land uses, p.A1 16.
Pyhala, A. Participation, institutions and protected area
management in Peruvian Amazonia, p.A1 17.
Redford, K. H. and Sanderson, S. E. Contested relationships
between biodiversity conservation and poverty alleviation,
pp.A119-120.
Regan, H. M., Davis, F. W. and Andelman, S. J. The use
of decision-making tools in systematic conservation
planning, p.A120
Regan, T. J., Burgman, M. A., McCarthy, M. A. and
Andelman S. J. Threatened species classifications and





Neotropical Primates 10(2), August 2002


population viability analysis: Shall ever the twain meet?
p.A120.
Rice, R. Conservation concessions: A new tool for
biodiversity conservation in the tropics, p.A121.
Rodrigues, A. S. L. and Gaston, K. J. How large do reserve
networks need to be? p.A123.
Roeder, A. D., Poinar, H. N. and Morin P. A. Collection
and storage methods for faecal samples prior to DNA
extraction, p.A124.
Rosendo, S. Institutional synergies in extractive reserves in
Amazonia, p.A125.
Schwind, K. Focus on farmers: Addressing the political and
economic causes of deforestation in Chiapas, Mexico,
p.A131.
Seymour, R., Bruford, M., Macleod, N. and Leader-
Williams, N. Subspecies and ESUS: The value of a holistic
approach to determining conservation units, p.A132.
Silva, J. M. C., Pinto, L. P., Cavalcanti, R. B. and Kierulff,
M. C. M. Establishing biodiversity corridors in the
Brazilian Atlantic forest, p.A134.
Valladares-Padua, C. and Martins, C. S. Metapopulation
management of black lion tamarins (Leontopithecus
chrysopygus) in remnants of Atlantic forest of the interior,
pp.A147-148.
Vizquez, M. A., Almeida, D., Nogales, E, Santander, T.,
Bonoccorso, E., Freile, J. F, Boada, C., Roman, H.,
Chiriboga, C., Andrade, K. and Aguirre, Z. The last
dry forest of a megadiversity country: Biological and
socioeconomic approach, pp.A149-150.
Velhzquez Rocha, I. A Nicaraguan dry forest reserve:
Balancing international private interests with local
community needs, p.A150.







The American Zoo and Aquarium Association (AZA)
Annual Conference, 10-14 September 2002, Fort Worth
Zoological Park, Fort Worth, Texas. The conference
program is geared toward the many disciplines in the
zoological profession directors, animal curators, keepers,
society members, scientists, gift shop merchandisers, and
practitioners in public relations, development, education,
and government affairs will all find something of interest.
Most of the AZA committees and special interests groups
meet in conjunction with the Annual Conference. For more
information: .

19th Annual Conference of the European Association
of Zoos and Aquaria (EAZA), 17-22 September 2002.
Hosted by Barcelona Zoo, Spain. The main theme of
the Conference will be Central and South America, with
emphasis on their current fund-raising and awareness
campaign the Atlantic forest of Brazil, Argentina and
Paraguay. The meeting will be held in the Pompeu Fabra
University, next to Barcelona Zoo. Website: www.eaza.net/index.html>.


III Congresso Brasileiro de Unidades de Conservacao,
22-26 de setembro de 2002, Centro de Convenq6es Edson
Queiroz, Fortaleza, Ceara. Realizaqao; Rede Nacional
Pr6-Unidades de Conservacao, Fundacao 0 Boticario de
Protegao a Natureza e Associaqao Caatinga. Patrocinio: The
Nature Conservancy. 0 event esta organizado de maneira
a permitir a apresentacao e discussao de grandes temas do
manejo de unidades de conservacao atraves de conferencias,
palestras e das sess6es paralelas: seminarios e apresentadao
de trabalhos t&cnicos-cientfficos. Informao6es sobre
Inscrio6es: Rowam Eventos, Telefax: 0** (41) 342-9078,
e-mail: <3cbuc@brturbo.com>.

VIII Congreso Latinoamericano y II Congreso
Colombiano de Botinica, 13-18 de octubre de 2002,
Cartagena de Indias, Colombia. "Nuestros conocimientos
al servicio de la sociedad". Informes: Enrique Forero, e-
mail: , o cias.unal.edu.co>. Website: eventos/congrbot/>.

Colloque 2002 Soci&t6 Francophone de Primatologie,
23-25 October, 2002. Dou&-la-Fontaine. This 14d annual
meeting of the Francophone Primate Society has the theme
of "Reproduction of Primates", but also regular sessions
on paleontology, anthropology, conservation, medical
research, ethology, and ecology, as well as a round table
on animal ethics. For more information visit the web
site: fontaine.htm>. For further information on the society visit:
.

X Congresso Brasileiro de Primatologia, 10-15
November 2002, Universidade Federal do Para, Belem.
Hosted by the Sociedade Brasileira de Primatologia (SBPr).
For more information: Stephen Ferrari, Departamento
de Psicologia, Universidade Federal do Para, Campus do
Guami, Caixa Postal 8607, 66075-150 Belem, Pard, Brazil,
e-mail: . Note: On 8th July 2002, the
organizing committee informed that the Congress had
been moved forward to November from the previously
announced dates of 25-30 August, 2002.

Foro de Primatologia 2002 Estaci6n de Biologia "Los
Tuxtlas", 21-22 de noviembre, 2002, Instituto de Biologfa
'Los Tuxtlas", Universidad Nacional Aut6noma de Mexico.
El objeto de esta reuni6n es actualizar e intercambiar
informaci6n acerca de investigaciones en curso con primates
nativos (Alouatta palliata, A. pigra y Ateles ...rr . I en el
sureste de Mexico y revisar los problems de conservaci6n
de las poblaciones. Esto permitirA determinar cual es el
estado de conocimiento acerca de la distribuci6n actual
de las poblaciones y su estado de conservaci6n, asf como
conocer los tipos de investigaciones bAsicas y aplicadas que
se llevan a cabo actualmente con primates silvestres en el sur
de Mexico. Tres dreas son de interns especifico: Poblacion y
ecologia -reconocimientos demogrificos, relaciones primate-
planta, recursos alimenticios, dispersi6n de semillas, y otros;
Conducta- ecologfa del comportamiento, conduct social; y





Neotropical Primates 10(2), August 2002


Conservacidn distribuci6n actual de las species, estado de
conservaci6n de las poblaciones, impact demografico de la
fragmentaci6n del habitat, destrucci6n y fragmentaci6n del
habitat, cacerfay trafico, proyectos de conservaci6n. Se desea
participar, comunicarse al correo
6 al fax + (294) 942-4668. Indicar si participaci6n es
como asistente o como presentaci6n de trabajo. Si es lo
segundo, enviar resdmen (max 250 palabras) antes del 5
de Noviembre, indicando si se trata de presentaci6n oral o
tipo cartel. Ndmero de asistentes al foro sera limitado, por
lo que se sugiere comunicar su participaci6n con suficiente
anticipaci6n. Los participants seran hospedados en las
instalaciones de la Estaci6n de Biologfa Los Tuxtlas. A los
asistentes cuyos trabajos sean aprobados para presentaci6n
se les cubriran gastos de estancia y alimentaci6n en la
Estaci6n Los Tuxtlas del IB-UNAM.

Primate Society of Great Britain (PSGB) Winter Meeting
2002, 29 November, 2002, Zoological Society of London,
Regent's Park, London, UK. The theme is "Primate Evolu-
tion and Adaptation". For information: Dr Sarah Elton,
Department of Anthropology, University of Kent at Can-
terbury, Canterbury CT2 7NS, Kent, UK, Tel: +44 (0)1227
823232, Fax: +44 (0)1227 827289, e-mail: @ukc.ac.uk>.

Dynamics and Conservation of Genetic Diversity in
Forest Ecosystems, 2-5 December, 2002. Strasbourg,
France. The conference will be divided into two main
parts: Part A, processes and mechanisms promoting genetic
diversity in forest ecosystems and Part B, implementations
in conservation strategies. Speakers will be presenting
information on forest trees and other short generation
species. A webpage for the conference is available at: //www.pierroton.inra.fr/genetics/Dygen/>. For further
information contact: DYGEN conference secretariat,
Dr. Marie-Pierre Reviron, INRA BP 45, 33610 Cestas,
France, Tel: +33 5 57 12 28 32, Fax: +33 5 57 12 28 81,
e-mail: .

XXIth Annual Conference of the Australasian Primate
Society, 6-8 December, 2002, Melbourne Zoo, Mel-
bourne, Australia. Organizers are Amanda Embury (Royal
Melbourne Zoological Gardens) and Debbie Williams
(CSL). For more details and to download a registration
form, please visit , or contact:
Amanda Embury, APS Conference Organizer, c/o Mel-
bourne Zoo. Australia, e-mail: .

2003

VI Congress Internacional en Gesti6n de Recursos
Naturales, 20 el 24 de enero de 2003, Hotel Villa del
Rio, Valdivia, Chile. Este event esta siendo organizado
por el Centro de Estudios Agrarios & Ambientales (CEA)
y cuenta con el auspfcio de importantes organizaciones
nacionales e internacionales. Este VI Congreso esta
estructurado en simposios: VIII Simposio de Manejo de
Vida Silvestre y Conservaci6n de la Biodiversidad, VI


Simposio Iberoamericano de Educaci6n y Comunicaci6n
Ambiental y VI Simposio de Desarrollo Sustentable,
I Simposio de Humedales y Recursos Hidricos y I
Simposio de Sistemas de Informaci6n Geograficos en
la Gesti6n de Recursos Naturales. Toda la informaci6n
relacionada con objetivos, program, estadia, inscripciones,
auspicios etc., esta en Internet en la direcci6n: //www.ceachile.cl/congresoVI.html>. Claudia Gil Cordero,
Comite Organizador VI CIGRN, Casilla 164, Valdivia.
Chile, Tel: 56-63-215846, Fax: 56-63-299065, e-mail:
o . Visite nuestra
pagina institutional en .

Student Conference on Conservation Science, 26-28
March, 2003, Conservation Biology Group, Department
of Zoology, University of Cambridge. "Building links
among young conservation scientists and practitioners".
Plenary lectures: Elizabeth Bennett (Wildlife Conserva-
tion Society), Andrew P. Dobson (Princeton Univer-
sity), Bob Pressey (New South Wales National Parks and
Wildlife Service) and Achim Steiner (Director-General,
IUCN World Conservation Union). Web site: www.zoo.cam.ac.uk/sccs/index.html>.

Primate Society of Great Britain (PSGB) Spring Meeting
2003, 10-11 April, 2003, School of Psychology, University
of St. Andrews, Fife, Scotland. Abstracts for oral presen-
tations, deadline: 10 January, 2003. Plenary talks will be
on Primate Cognition. Invited speakers include: Andrew
Whiten, Hannah Buchanan-Smith, Kevin Laland and
Debbie Custance. For more information: Dr. Klaus Zu-
berbuhler, e-mail: , or Gillian Brown,
e-mail: .

4th European Congress of Mammalogy, 27 July
- 1 August, 2003, Brno, Czech Republic. Hosted by the
Institute of Vertebrate Biology, Academy of Sciences of
the Czech Republic. Information and the pre-registration
form are available on the website .
Any questions about organization should be directed to
Jan Zima, Organising Committee, e-mail: .
The first information and the pre-registration form are now
available on the website: < http://www.ivb.cz>.

28th International Ethological Conference, 20-27 August
2003, Costao do Santinho Resort, Florianopolis, Brazil.
On behalf of the International Council of Ethologists
and hosted by the Brazilian Society of Ethology. Deadline
for submission of symposia: 31 January 2003. Deadline
for submission of abstracts, financial aid applications,
and standard reduced registration rate: 20 February
2003. For more information on the conference contact:
Professor Kleber del Claro, e-mail: ,
or on the scientific program, contact Professor Regina
Macedo, e-mail: . Web site: //www.iec2003.org/home.htm>.




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