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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
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Publication Date: June 1996
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Full Text
ISSN 1413-4703

A Newsletter of the Neotropical Section of the IUCN/SSC Primate Specialist Group
Editors: Anthony B. Rylands and Emesto Rodriguez Luna
PSG Chairman: Russell A. Mittermeier
PSG Deputy Chairman: William R. Konstant


Jr \


, r. .-

Neotropical Primates 4(2), June 1996



In their review on the current status of New World
monkey classification at the species and subspecies level,
Mittermeier and Coimbra-Filho (1981) discussed the
taxonomic problems of each genus and pointed out
weaknesses and the need for further research. They
indicated that Ateles geoffroyi is one of the species in
need of a major taxonomic revision, which in our opinion
is becoming an important problem in terms of its
conservation status. In this note, we comment this and
other authors' (e.g., Konstant et al., 1985; Mittermeier
et al., 1988) notes on the taxonomic status of Ateles
geoffroyi vellerosus, A. g. yucatanensis and A. g. pan,
and compare them to the observations we have made
and recorded in study sites from three countries; Mexico,
Belize, and Guatemala.

Mittermeier and Coimbra-Filho (1988) referred to Ateles
as a "variable genus" and suggested that "...some
rearrangement and perhaps reduction of the species and
subspecies recognized by Kellogg and Goldman is
probably necessary." Roosmalen and Klein (1988),
referring to Hernindez-Camacho and Cooper (1976),
noted that "Differences between species and subspecies
are based almost entirely upon the pelage characteristics.
These are, to some extent, variable within populations
and may intergrade between populations over large parts
of their range...". However, considering the subspecies
vellerosus, yucatanensis, and pan, to what extent are
the pelage characteristics "variable" within populations,
and how much may they "intergrade" between
populations over large parts of the species' range? In
our experience, the species' pelage characteristics are
widely variable and therefore not sufficiently reliable to
be considered as one of the major traits supporting
subspecies' distinctions in the cases of vellerosus,
yucatanensis, and pan. In other words, we believe the
pelage characteristics of Ateles geoffroyi should not be
regarded as the measure of distinctive taxonomic traits
among the three subspecies. The following observations
are summarized to support our statement.

A. geoffroyi vellerosus

In their assessment of Ateles geoffroyi's current
taxonomic situation, Konstant et al. (1985) described
A. g. vellerosus as a subspecies in which "...dorsal
surfaces...range from black to dark brown, except for a
light band across the lumbar region, and contrast strongly
with its lighter abdomen and inner limbs. Exposed flesh-

colored skin is often present about the eyes." This
description is compatible with the field observations of
Gilberto Silva-L6pez, Joaquin Jim6nez-Huerta, Maria
Rebeca Toledo-Cardenas, and Jorge Benitez-Rodriguez
on A. g. vellerosus at Sierra de Santa Marta, Veracruz,
Mexico, except for the fact that these researchers also
found several adult individuals in which: (a) the dorsal
surfaces were not as dark, (b) the allegedly lighter band
across the lumbar region was not very marked, and (c)
the contrast between the color and tones of the dorsal
surfaces and the inner limbs was not at all clear. Konstant
et al. (1985.) also pointed out that the "...subspecies can
apparently be distinguished from A. b. belzebuth and
the darker variety of A. belzebuth hybridus by the
absence or marked reduction of a white triangular
forehead patch and sideburns", bit observations at Santa
Marta suggest that the white trianguilairfbrehead patch
may be very common in A. g. vellerosus. Biologists J.
Jim6nez-Huerta and J. Benitez-Rodriguez (pers. comm.),
for example, observed several individuals with this
characteristic, and even came to the point of
distinguishing one of the female dominated subgroups
by the presence of"Blanca", an adult female with a large
white triangular forehead patch, a trait which was shared
with two of the infant females that were forming part of
Blanca's subgroup.

These observations suggest that the vellerosus subspecies
has a wide variety of pelage colors and tones which,
according to campesinos of the Sierra, may vary between
"muy negro a muy blanco" (very black to very white).
Mexican researcher Alvar Gonzalez Christen (pers.
comm.) had the chance to observe the "dirty white"
coloration of an adult spider monkey in the crater of the
Santa Marta volcano, and Hall and Dalquest (1963,
p.262) noted that "One man reported an albino spider
monkey in the hills west of Jimbal (Veracruz)." Our team
also observed a whitish vellerosus spider monkey while
at the National Park of Tikal, Guatemala, with a darker
distal third of the tail. Both A. Gonzalez-Christen in
Veracruz, and our team in Guatemala, found no evidence
to conclude these monkeys were albinos, but merely
individuals with a markedly whitish coloration.

A. geoffroyi yucatanensis

Konstant et al. (1985) described A. g. yucatanensis as
"... brownish-black on its head, neck and shoulders,
becoming lighter brown on the lower back and hips and
contrasting with its silvery white underside, inner limbs
and sideburns..." They also noted that the subspecies
"...may be confused with lighter individuals of A.
geoffroyi vellerosus." None of the papers by Elizabeth
Watts and Victor Rico-Gray (e.g., Watts et al., 1986;
Watts and Rico-Gray, 1987), who studied spider
monkeys in the Yucatan Peninsula, provide evidence

Cover photograph by Russell A. Mittermeier: Alouatta pigra from the Bermuda Landing, Belize.

Page 41

Neotropical Primates 4(2), June 1996

on the subject and only note that "The taxonomic status
of (Ateles geoffroyi yucatanensis) is controversial and
little is known of (its) natural history and ecology". The
general description of the subspecies by both Dr. DamiAn
Rumiz and Gilberto Silva-L6pez at the Rio Bravo
Conservation and Management Area of Belize (Fragoso
et al., 1990; Silva-L6pez and Rumiz, 1995) is quite
similar to that of William Konstant, Russell Mittermeier
and Stephen Nash, but D. Rumiz and G. Silva-L6pez
also noted that "The pattern of color observed in the
spider monkeys more closely resembled that of the
Mexican subspecies (A. g. vellerosus) than the Yucatan
subspecies (A. g. yucatanensis), contrary to what was
previously reported for Belize (Kellogg and Goldman,
1944)", and continue "We might say there was certain
inter-individual variation in the color, which made it
difficult to assign individuals to a particular subspecies.
In fact, some individuals showed patterns of color that
are intermediate between the subspecies' descriptions
and color representations made by Konstant et al.

A. geoffroyi pan

The Guatemalan spider monkey, Ateles geoffroyi pan,
was reported by Konstant etal. (1985) as "...very similar

Figure 1. Presumed geographical range of Ateles geoffroyipan (see
Konstant et al., 1985). The figure has been modified for this report
(see text). Legend: BA: Barillas; AL: Aldea Juil; FC: FincaChelemhi;
BI: Biotopo Mario Dary Rivera; CH: Chilasc6. The thick line delimits
the geographical range proposed for A. g. vellerosus; diagonal lines
indicate tropical forest, the dotted line is the Panamerican Highway,
the dashed lines show the Sierras of ChamA (A), Chuactis (B), and
Cuchumatanes (C). The numbers indicate the Departments within
the range: 1. Huehuetenango, 2. Quich&. 3. Alta Verpaz, 4. Baja
Verapaz, 5. El Progreso, 6. Guatemala, 7. Sacatepdquez, 8.
Chimaltenango, 9. Escuintla, 10. Suchitep6quez, 11. Solola, 12.

Page 42

to the darker colored individuals of A. geoffroyi
vellerosus", and described it as "...having a thick black
coat, and is said to occur at high altitudes. It differs from
A. g. vellerosus in that its dorsal coloration does not
contrast as markedly with that of its ventral surface, and
it does not possess a lighter-colored saddle on its lumbar
region." However, the description of the subspecies,
which is "...supposed to occur in the central mountains
of Guatemala..." was based "...on only 3 animals of
unknown geographic origin (Kellogg and Goldman,
1944)...", which encouraged Konstant and coworkers
to conclude pan "...may not be a valid taxon."

Our study of the subspecies' geographical range, as
proposed by Kellogg and Goldman (1944, in Konstant
et al., 1985), seems to support the remark by Konstant
et al. regarding the validity of pan as a distinct
subspecies. The area of reference appears in more detail
in Figure 1. As can be seen, the area includes important
portions of several Guatemalan departments. The main
vegetation type in the area is coniferous forest, with
Pinus, Quercus, and Liquidambar among the dominant
genera. Some remnants of tropical forest can be found
in the lowlands of Alta Verapaz and Quich6 (including
the locality of Barillas), to the north; near Chilasc6 and
in the Biotopo Mario Dary Rivera, in the east; and in
Escuintla and Retalhuleu, in the south. The latter,
however, is a very disturbed area surrounding a segment
of the Panamerican Highway, with extensive cultivation
of maize, rice, banana, and beans.

Portions of three important sierras form part of the area,
including: the Sierra de Chami (in Alta Verapaz and
Quich6), with elevations ranging from 300 to 1,500 m;
the Sierra de Chuacils (in Baja Verapaz and Quiche),
with elevations ranging from 600 to 2100 m; and the
Sierra de los Cuchumatanes (in Huehuetenango and
Quiche), with elevations between 1,500 and 2,700 m.
Barillas is located on the northern slope of the
Cuchumatanes, but Alouattapalliata is the only monkey
species reported for the area. The same species was
reported by Villar (1994) in the Biotopo Mario Dary
Rivera, which is located in the highlands of the Sierra
de Chuactis (a zone also known as the highlands of the
interior). In Chilasc6, another locality included in our
records, the only species reported is Alouatta pigra.
Chilasc6 forms part of the Sierra de las Minas. The
literature and field reports we have indicate spider
monkeys have not been recorded in this area.

Captive monkeys have also been studied. In 1990-1991,
Johanna Motta carried out a detailed survey of captive
spider monkeys in four Guatemalan zoos (three
individuals from La Aurora Zoo, 11 individuals from
The Jungle Zoo [IRTRA], nine individuals from the
Minerva Zoo, and three individuals from the Petencito

Page 43 Neotropical Primates 4(2), June 1996

Zoo), one safari park (Autosafari Chapin, 19
individuals), and two private collections (one individual
from Finca Nacional Santo Tomis, and four individuals
from the Finca Enrique Ponce), taking footprints,
pictures, measuring and weighing every single
individual, as well as obtaining blood samples of the
monkeys and inquiring about their history. Her purpose
was to conduct a chromosome study of the monkeys,
supported by accurate records from the sampled
individuals. Dr. Anne Baker, then of the Brookfield Zoo,
Chicago, helped in the analyses of the blood samples in
1991, but unfortunately the results were not conclusive.
However, the interviews, observations, and analyses of
photographs of the 50 individuals led her to conclude
that: (a) no single color pattern was dominant among
the individuals studied, (b) the individual range of
patterns of color and tone varied remarkably, from very
dark-colored to very whitish-colored animals, and (c)
the study provided insufficient evidence to conclude that
pan and yucatanensis, if considered valid taxa, are
among Guatemala's captive spider monkeys.

The spider monkeys at La Aurora Zoo, Guatemala City,
provide an example of the wide spectrum of colors and
tones in Ateles geoffroyi. We had the chance to observe
La Aurora's spiders and are in agreement with the
researcher Lorena Calvo (see Konstant et al., 1985) that
the subspecies kept at the zoo is A. g. vellerosus.

All these observations and records have led us to
conclude that:
1. The coloration pattern of A. g. vellerosus includes
a broader spectrum of color and tones than the
previously considered;
2. A. g. yucatanensis may be considered a valid taxon,
but only after more evidence (photographs,
morphological studies, caryological studies, and field
observations) can be obtained from several localities
in the known geographical range of the subspecies;
3. based on the available evidence (maps, vegetation
types, and existing records) and supporting the
observations of Konstant et al. (1985) on the subject,
we conclude that A. g. pan should not be considered a
valid taxon.


Along the years, support from The Charles A. and Anne
Morrow Lindbergh Foundation, the WWF-U.S. Primate
Program, the International Primatological Society (IPS),
the Chicago Zoological Society, the Program for Belize,
the Manommet Bird Observatory, the Program for
Studies in Tropical Conservation of the Department of
Wildlife Ecology and Conservation, University of
Florida, Gainesville, NYZS The Wildlife Conservation
Society, CONACYT Mexico (# 54800), and the Instituto

de Investigaciones Biol6gicas of the Universidad
Veracruzana, have made possible the field observations
included in this report. GSL also thanks A. B. Rylands
for his encouragement. This a contribution from the
Program Bioconservaci6n of the Instituto de
Investigaciones Biol6gicas, Universidad Veracruzana.

Gilberto Silva-L6pez, Programa Bioconservaci6n,
Institute de Investigaciones Biol6gicas, Universidad
Veracruzana (IIB-UV), Apartado Postal 294, Xalapa,
Veracruz, 91000 Mexico, Johanna Motta-Gill,
Researcher, Guatemala, and Alonso I. SAnchez
HernAndez, Programa Bioconservaci6n, Instituto de
Investigaciones Biol6gicas, Universidad Veracruzana
(present address: Jardin BotAnico de Tlaxcala).


Fragoso, J., Rumiz, D., Hunter, T., Silva-L6pez, G. and
Smith, R. 1990. Wildlife inventory of the Rio Bravo
Conservation and Management Area, Belize: A report
to the Manommet Bird Observatory and the
Programme for Belize. Program for Studies in Tropi-
cal Conservation, University of Florida, Gainesville.
Hall, E. R. and Dalquest, W. W. 1963. The Mammals of
Veracruz. University of Kansas Publ. Mus. Nat. Hist.
HernAndez-Camacho, J. and Cooper, R. W. 1976. The
non-human primates of Colombia. In: Neotropical
Primates: Field Studies and Conservation, R. W.
Thorington, Jr. and P. G. Heltne (eds.), pp.35-69. Na-
tional Academy of Sciences, Washington, D. C.
Kellogg, R. and Goldman, E. A. 1944. Review of the
spider monkeys. Proc. U. S. Nat. Mus. 96:1-45.
Konstant, W., Mittermeier, R. A. and Nash, S. D. 1985.
Spider monkeys in captivity and in the wild. Primate
Conservation (5):82-109.
Mittermeier, R. A. and Coimbra-Filho, A. F. 1981. Sys-
tematics: species and subspecies. In: Ecology and Be-
havior ofNeotropical Primates, Vol. 1, A. F. Coimbra-
Filho and R. A. Mittermeier (eds.), pp.29-110.
Academia Brasileira de Ci8ncias, Rio de Janeiro.
Mittermeier, R. A., Rylands, A. B. and Coimbra-Filho,
A. F. 1988. Systematics: species and subspecies an
update. In: Ecology and Behavior ofNeotropical Pri-
mates, Vol. 2, R. A. Mittermeier, A. B. Rylands, A. F.
Coimbra-Filho and G. A. B. da Fonseca (eds.), pp. 13-
75. World Wildlife Fund, Washington, D. C.
Roosmalen, M. G. M. van and Klein, L. L. 1988. The
spider monkeys, genus Ateles., In: Ecology and Be-
havior of Neotropical Primates, Vol. 2, R. A.
Mittermeier, A. B. Rylands, A. F. Coimbra-Filho, and
G. A. B. da Fonseca (eds.), pp.455-537. World Wild-
life Fund, Washington, D. C.
Silva-L6pez, G. and Rumiz, D. 1995. Los primates de
la reserve Rio Bravo, Belice. La Cienciay el Hombre

Neotropical Primates 4(2), June 1996

Page 43

Neotropica/ Primates 4(2), June 1996 Page 44

Villar, L. 1994. Untitled short note in the mural news-
paper of the Centro de Estudios Conservacionistas of
the Universidad de San Carlos de Guatemala.
Watts, E. S., Rico-Gray, V. and Chan, C. 1986. Mon-
keys of the Yucatan Peninsula, Mexico: Preliminary
survey of their distribution and status. Primate Con-
servation (7): 17-22.
Watts, E.S. and Rico-Gray, V. 1987. Los Primates de la
Peninsula de Yucatan, Mexico: studio preliminary
sobre su situaci6n actual y estado de conservaci6n.
Bi6tica 12(1):57-66.


La clasificaci6n taxon6mica de los monos del Nuevo
Mundo ha sufrido constantes reordenamientos desde el
pasado siglo. Sin embargo, se arrib6 a un consenso mAs
o menos generalizado al separarlos en 2 families:
Callitrichidae (Callithrix, Cebuella, Leontopithecus,
Saguinus), y Cebidae, agrupando a todos los g6neros
restantes (Simpson, 1945; Cabrera, 1958; Simons, 1972);
esto se transform en la clasificaci6n traditional que en
muchos casos continia vigente, con el objetivo principal
de distinguir a los "Callitrichidae", aquellos platirrinos
de pequefio tamafio corporal, poseedores de garras en
lugar de ufias plans, que han perdido el tercer molar y
que dan a luz dos crias. No obstante, todos estos
caracteres fueron observados como adquisiciones
derivadas (Ford, 1980) en contrast con la hip6tesis de
que son caracteres primitivos retenidos desde los
platirrinos ancestrales (Hershkovitz, 1977). Pero es obvio
que la gran diversidad del Infraorden Platyrrhini va mis
alli de esta dicotomia familiar.

Rosenberger (1981) consider que los Cebidae reonen
s6lo a Cebus y Saimiri (Cebinae) con Callithrix,
Cebuella, Leontopithecus, Saguinus y Callimico
(Callitrichinae), mientras que los restantes se agrupan
en la Familia Atelidae, subdividida en Atelinae (Ateles,
Lagothrix, Brachyteles y Alouatta) y Pitheciinae
(Pithecia, Chiropotes, Cacajao, y Callicebus y Aotus
como taxones hermanos mAs distantes). El esquema de
Ford (1986) difiere en la exclusion de Callicebus y Aotus
fuera de los Pitheciinae, agrupandolos con Cebus y
Saimiri en la Familia Cebidae, aunque s6lo a los efectos
de preservar el amplio uso de "Cebidae" y la
contraposici6n hist6rica con Callitrichidae (Callithrix,
Cebuella, Leontopithecus, Saguinus y Callimico, sensu
Ford). No obstante, Ford aclara la escasa sustentaci6n
de su Familia "Cebidae", en especial la posici6n de Cebus
en relaci6n a los restantes platirrinos. Kay (1990)
establece que Aotus represent el tax6n hermano de los
Atelinae (Ateles, Brachyteles, Lagothrix y Alouatta), en

tanto Saimiri se relacionaria con los Callitrichinae; Cebus
se consider aqui una forma mas primitive que divergi6
antiguamente de los restantes grupos.

De estos esquemas, podemos ver que existe consenso
en algunos clados, pero la problematica se centra en los
gCneros Aotus, Callicebus, Cebus y Saimiri. Thorington
y Anderson (1984), en respuesta a las diferentes
hip6tesis, reunieron a todos los platirrinos en la 6nica
Familia Cebidae, subdivididaen subfamilias; aquiAotus,
Callicebus, Cebus y Saimiri son separados en las
subfamilias monotipicas Aotinae, Callicebinae, Cebinae
y Saimiriinae, respectivamente. Los restantes clados se
conservan como fue sefialado mis arriba, a excepci6n
de Alouatta, tambi6n separado en Alouattinae; pese a
que cominmente se lo agrupa con los Atelinae, aun no
es clara la posici6n de Alouatta, puesto que conserve
ciertos caracteres de la dentici6n cuya polaridad es

Al parecer no existen sinapomorfias dentarias exclusivas
de todos los platirrinos (Szalay y Delson, 1979; Kay,
1980; Rosenberger, 1981); por el contrario, hallamos la
que posiblemente sea la tinica sinapomorfia craneal,
consistent en el contact entire parietal y yugal, evitando
la conexi6n entire frontal y aliesfenoides, en la region
pt6rica (Ashley-Montague, 1933; Le Gros Clark, 1959;
Rosenberger, 1977; Delson y Rosenberger, 1980; Ford,
1986). Tambi6n existirian al menos tres caracteres
postcraneales finicos para todos los platirrinos, a
excepci6n de su possible presencia en ciertos especimenes
del Oligoceno de Egipto (Ford, 1986). Es decir que en
principio no poseemos suficiente informaci6n confiable
para reunir a todos los platirrinos en un clado

Tabla 1. Clasificacion de los Platyrrhini.
Familia Atelidae
Subfamilia Atelinae
Subfamilia Alouattinae
Subfamilia Pitheciinae
Subfamilia Callitrichinae
Subfamilia Cebinae
Subfamilia Saimiriinae
Subfamilia Aotinae
Subfamilia Callicebinae

Neolropical Primates 4(2), June 1996

Page 44

Page 45 Neotropical Primates 4(2), June 1996

monofil6tico (la carencia de sinapomorfias puede
trasladarse a todo el Orden Primates, dado que la finica
que poseen estos es la bula formada por una extension
del petroso, en el complejo temporal), y luego se
presentan a6n mis inconvenientes al buscar caracteres
derivados exclusivos de todos los "Cebidae" de la
clasificaci6n traditional.

Es sabido que los f6siles deben contribuir, y de hecho
son la parte fundamental, en la compleja construcci6n
taxon6mica de las formas vivientes; entonces hallamos
que los f6siles mAs abundantes y mejor preservados
consistent en dientes aislados o in situ en especimenes
fragmentarios. Tambidn asumimos que estos dientes nos
proven de una important informaci6n sistemAtica, por
lo cual, pese a deducir que no existirian sinapomorfias
dentarias que sean exclusivas de los Platyrrhini, podemos
aproximarnos a una historic evolutiva lo suficientemente
fundamentada dentro del infraorden, mas ain teniendo
en cuenta el gran incremento de hallazgos f6siles en las
Oltimas dos d6cadas. Los analisis de la dentici6n, entire
otros, nos muestran entonces que no existe una "Familia
Cebidae" que agrupe a los no-Callitrichinae, y
consecuentemente el registro f6sil corrobora esta

Como resultado de un analisis cladistico de 67 caracteres
dentarios (Tejedor, ms.) se evidenci6 una estrecha
relaci6n entire los Callitrichinae (representados en este
studio por Callithrix, Leontopithecus y Callimico) y
Saimiri y Cebus, este ultimo como el tax6n hermano
mAs distant. Rosenberger (1981) obtuvo id6nticos
resultados basado en caracteres craneodentarios;
asimismo, recientes studios moleculares demuestran
similar agrupamiento (Schneider et al., 1995, incluyendo
ademis Aotus y Callicebus) o excluyen s6lo a Cebus
(Meldrum et al., 1993). Por otra parte, en el mencionado
analisis cladistico, Callicebus y Aotus aparecen como
taxones hermanos de los restantes platirrinos, y mAs
distant atn se halla Homunculus, un f6sil del Mioceno
temprano de Patagonia, de cercanas vinculaciones con
Callicebus y, en menor grado, con Aotus. Los otros
g6neros utilizados en este analisis fueron Ateles (como
representante de los Atelinae, excluyendo Alouatta) y
Pithecia y Cacajao (representando a los Pitheciinae),
separados ambos grupos en clados bien distintivos, cuya
monofilia es indudable. En estas instancias, y recordando
el esquema de Thorington y Anderson (1984), es
pertinente sugerir el uso de subfamilias para separar los
distintos clados monofil6ticos de los Platyrrhini (Tabla
1), donde filogen6ticamente puede corroborarse, luego
de extensos debates, la estrecha relaci6n entire los
Callitrichinae y Cebus (Cebinae) y Saimiri (Saimiriinae).
Provisionalmente, Alouatta es considerado como inico
integrante de la Subfamilia Alouattinae, aunque restan
studios mas amplios para comprender su exacta

posici6n filogenetica. No fue incluido Homunculus
dentro de esta clasificaci6n porque s6lo se utilize en el
analisis cladistico para concluir si existia una cercana
relaci6n con Callicebus y Aotus, ya que previas
observaciones indicaban una gran afinidad morfol6gica,
y mas precisamente Callicebus muestra importantes
caracteres primitivos compartidos con Homunculus
(Tejedor, 1995a, 1995b). La prolongada evoluci6n
independiente de Callicebus y Aotus conduce a
separarlos en las subfaiT.lias monotipicas Callicebinae
y Aotinae, respectivamente; ambos presentan caracteres
diferenciales en los incisivos, siendo en Aotus mas
espatulados y de implantaci6n mas vertical, en tanto que
en Callicebus son mas estrechos y procumbentes. Los
caninos son reducidos y escasamente proyectados sobre
el plano oclusal, mientras que los molares tienen c6spides
bien desarrolladas, mas prominentes en Callicebus, quien
posee ademas un gran hipocono y marcado cingulo
lingual en los molares superiores. Algunas de las
tendencies derivadas en otros platirrinos actuales
conducen a extremar la procumbencia de los incisivos
(Pitheciinae, Callithrix, Cebuella), incrementar el
tamafio de los caninos, con su mAxima expresi6n en los
Pitheciinae, reducir el tercer molar (Cebus, Samiri, e
incluso se pierde en los Callitrichinae, except
Callimico), acentuar el desarrollo de crestas en los
molares (Brachyteles, Alouatta), reducir el relieve
oclusal de premolares y molares (Pitheciinae), o
aumentar notablemente el espesor del esmalte dentario
(Cebus). Parte de l a polaridad tentative de estos
caracteres se deduce de la informaci6n que provee
Homunculus, como antiguo representante del infraorden
(Tejedor, ms.).

Es oportuno insistir en la utilizaci6n del nombre familiar
Atelidae (Tabla 1), que agrupa segin este studio a todos
los platirrinos, en reemplazo de "Cebidae". Delson
(comunicaci6n personal; ver tambidn Rosenberger et al.,
1990) advierte que el t6rmino supragen6rico Atelina
Gray 1825, basado en el g6nero Ateles, antecede a Cebina
Bonaparte 1831, basado en Cebus, y dado que- a
instancias del C6digo Internacional de Nomenclatura
Zool6gica todos los nombres del "grupo familiar"
(Superfamilia, Familia, Subfamilia, Tribu) deben estar
basados en el mismo g6nero con la misma prioridad en
todas las jerarquias dentro de ese grupo familiar, no
existen razones para continuar utilizando "Cebidae".

Reiterando que estos studios estan basados en el analisis
de la dentici6n exclusivamente, es imprescindible contar
con mayores contribuciones a partir de otras alternatives
de trabajo, para elucidar la compleja radiaci6n
filogen6tica de los platirrinos.

Marcelo F. Tejedor, CAtedra de Anatomia Comparada,
Facultad de Ciencias Naturales y Museo, Paseo del

Page 45

Neotropical Primates 4(2), June 1996

Neotropical Primates 4(2), June 1996 Page 46

Bosque, 1900, La Plata, Argentina.


Ashley-Montague, M. F. 1933. The anthropological sig-
nificance of the pterion in primates. Am. J. Phys.
Anthropol. 18:159-336.
Cabrera, A. 1958. CatAlogo de los mamiferos de Amdrica
del Sur, I. Rev. Mus. Arg. Cienc. Nat. "Bernardino
Rivadavia" 4(1): 1-307.
Delson, E. y Rosenberger, A. L. 1980. Phyletic Perspec-
tives on Platyrrhine Origins and Anthropoid Relation-
ships. En: Evolutionary Biology of the New World
Monkeys and Continental Drift, R. L. Ciochon y A. B.
Chiarelli (eds.), pp. 445-458. Plenum Press, New York.
Ford, S. M. 1980. Callitrichids as phyletic dwarfs, and
the place of the Callitrichidae in Platyrrhini. Primates
Ford, S.M. 1986. Systematics of the New World mon-
keys. En: Comparative Primate Biology, Vol. I: Sys-
tematics, Evolution and Anatomy, D. Swindler y J.
Erwin (eds.) pp.71-135. Alan R. Liss, New York.
Hershkovitz, P. 1977. Living New World Monkeys
(Platyrrhini), With an Introduction to Primates, Vol.
I. Chicago University Press, Chicago.
Kay, R. F. 1980. Platyrrhine origins: a reappraisal of
the dental evidence. En: Evolutionary Biology of the
New World Monkeys and Continental Drift, R. L.
Ciochon y A. B. Chiarelli (eds.), pp. 159-188. Plenum
Press, New York.
Kay, R.F. 1990. The phyletic relationships of extant and
fossil Pitheciinae (Platyrrhini, Anthropidea). J. Hum.
Evol. 19: 175-208.
Le Gros Clark, W. E. 1959. The Antecedents of Man.
University of Edinburgh Press, Edinburgh.
Meldrum D. J., Kay, R. F. y Chiu, C.-H. 1993. Phyloge-
netic relationships of Cebus and Saimiri inferred from
mitochondrial DNA sequences and dental anatomy.
Am. J. Phys. Anthropol. 16:144-145.
Rosenberger, A.. L. 1977. Xenothrix and ceboid phy-
logeny. J. Hum. Evol. 6:461-481.
Rosenberger, A. L. 1981. Systematics: the higher taxa.
En: Ecology and Behavior of Neotropical Primates,
Vol. 1, A. F. Coimbra-Filho y R. A. Mittermeier (eds.),
pp.9-27.: Academia Brasileira de Cidncias, Rio de
Rosenberger, A. L., Setoguchi, T. y Shigehara, N. 1990.
The fossil record of callitrichine primates. J. Hum.
Evol. 19:209-236.
Schneider, H., Schneider, M. P. C., Sampaio, I., Harada,
M. L., Barroso, C. M. L., Czelusniak, J. y Goodman,
M. 1995. DNA evidence on platyrrhine phylogeny
from two unlinked nuclear genes. Am. J. Phys.
Anthropol. suppl. 20:191.
Simons, E. L. 1972. Primate Evolution. Macmillan, New

Simpson, G. G. 1945. The principles of classification
and a classification of mammals. Bull. Am. Mus. Nat.
Hist. 85:1-350.
Szalay, F. S. y Delson, E. 1979. Evolutionary History of
the Primates. Academic Press, New York.
Tejedor, M. F. 1995a. Descripci6n de nuevos restos
dentarios asignados a Homunculus patagonicus (Pri-
mates, Platyrrhini), procedentes de la localidad de
Monte Observaci6n (Santacrucense), Provincia de
Santa Cruz. Restimenes XI Jornadas Argentinas de
Paleontologia de Vertebrados, Tucumdn, Mayo de
Tejedor, M. F. 1995b. La posici6n de Aotus y Callicebus
en la filogenia de los primates Platyrrhini. Resutmenes
X Jornadas Argentinas de Mastozoologia, La Plata,
Noviembre de 1995.
Thorington, R. W., Jr. y Anderson, S. (1984). Primates.
En: Orders and Families of Recent Mammals of the
World, S. Anderson, S. & J. Knox Jones, Jr. (eds.),
pp.187-217. Wiley and Sons, New York.


The discovery of two nearly complete skeletons of large
Pleistocene primates from Bahia, Brazil, was announced
in these pages three years ago (Cartelle, 1993).
Preliminary analysis of these fossils is now complete,
and they are identified as two different genera of very
large atelines (Fig. 1). One skeleton is an adult individual
referred to Protopithecus brasiliensis Lund, 1838
(Hartwig and Cartelle, 1996), and the other is a nearly

Figure 1. Frontal and lateral views of the skull of Protopithecus
(left) and Caipora (right). Illustration by Humberto do Espirito Santo.

Neotropical Primates 4(2), June 1996

Page 46

Table 1. Cranial measurements for Protopithecus, Caipora, and the four genera of living ateline New World monkeys. All measurements in
Protopithecus Caipora Ateles Brachyteles Lagothrix Alouatta
n=1 n=I n=92 n=ll n=73 n=25
NCL 110.1 94.1 77.9 86.6 73.7 61.2
68.5-84.4 79.8-91.7 67.0-81.2 54.9-68.8
NCB 72.8 75.4 60.8 61.9 58.6 51.0
54.9-65.7 57.7-65.1 53.8-63.1 47.3-56.2
TSL 150.5 131.5 114.1 114.8 104.9 107.6
104.0-122.0 100.0-122.0 97.2-114.6 96.3-121.4
BAS-NAS 83.4 77.2 63.3 68.2 63.1 64.9
55.4-71.9 58.2-74.4 57.9-70.0 57.6-78.2
PL 43.8 40.6 34.4 38.7 31.6 39.9
29.9-40.7 34.2-44.1 26.2-37.4 34.7-59.9
BOB 70.8 63.3 55.6 57.3 54.0 52.3
49.9-64.2 52.0-61.2 47.9-59.0 47.2-60.7
NCL = Neurocranial length, NCB = Neurocranial breadth, TSL = Total skull length, BAS-NAS = Basion-nasion, PL = Palate length, BOB =
Biorbital breadth.

Table 2. Postcranial measurements for Protopithecus, Caipora, and the four genera of living ateline New World monkeys. All measurements
in millimeters, except where indicated. Asterisk denotes incomplete growth of the fossil.
Protopithecus Caipora Ateles Brachyteles Lagothrix Alouatta
n=1 n= I n=31 n=3 n= 17 n=25
FHD 25.0 22.9 17.9 18.2 15.0 13.4
15.8-20.2 16.9-19.8 14.0-15.7 11.8-15.9
FND 16.2 15.2 9.9 10.3 8.3 7.9
8.0-11.8 9.5-11.2 7.4-9.6 6.4-10.8
FL 237 216* 205.6 202.0 166.4 154.2
190.5-226.0 186.5-212.0 157.5-176.5 139.0-171.0
BCB 45.4 38.5 31.8 29.0 27.1 23.9
29.1-34.9 28.0-30.9 24.2-29.3 21.4-26.7
HHD 28.4 25.1 20.5 19.8 20.1 19.8
17.8-24.1 18.2-21.6 18.6-22.2 16.9-23.1
HMST 18.5 18.5 11.0 10.4 10.2 9.5
10.0-12.4 9.4-11.6 9.2-11.4 7.3-12.1
BIEPI 48.0 37.7 30.9 30.0 28.0 26.6
28.5-33.2 26.7-33.0 25.6-30.0 22.5-30.8
1-1 1.04 1.06* 1.05 1.07 0.98 0.95
1.01-1.07 1.05-1.08 0.96-1.0 0.92-0.98
FV 90 ml 70 ml 40 ml (n=l) 20 ml (n=1) 20 ml (nl)
FHD = Femoral head diameter, FND = Femoral neck diameter, FL = femoral length, BCB = femoral bicondylar breadth, HHD = humeral head
diameter, HMST = humeral midshaft thickness, BIEPI = humeral biepicondylar breadth, I-I = Intermembral index (forelimb length/hindlimb
length), FV = femoral volume (measured by water displacement in a graduated cylinder).

mature subadult recently described as a new genus and
species, Caipora bambuiorum Cartelle and Hartwig,
1996 (Fig. 1).

The complete skeleton of Protopithecus is remarkable
for its large size, calculated to be approximately 25 kg
based on extrapolations from postcranial measurements
(Tables 1 and 2). It is further remarkable because the
cranium distinctly resembles Alouatta, the howler
monkey, while the postcranium bears the same
adaptations to suspension and brachiation as Ateles and
Brachyteles. The type specimen discovered in Minas
Gerais by Lund in 1836 was known only from a proximal
femur and a distal humerus, and from these it was
assumed that Protopithecus was a large Pleistocene
muriqui (Hartwig, 1995). The presence of a flat, posterior
nuchal plane, and an extended basicranium in the skull

of the new skeleton was entirely unexpected.
Protopithecus presents as unique mixture of size and
morphology, and shows that atelines were more diverse
in the recent past.

The Caipora skeleton resembles living spider monkeys
in both cranial and postcranial details. The skull is
sufficiently different from Protopithecus in the shape
of the neurocranium and basicranium to warrant its own
genus. It differs from Ateles in having a much wider,
and thus larger braincase. The limb bones are not fully
grown, but suggest that this Caipora individual weighed
approximately 20 kg.

Together, Protopithecus and Caipora represent the
largest South American primates, the most complete
fossil platyrrhines, and the first substantial record of

Neotropical Primates 4(2), June 1996

Page 47

Neotropical Primates 4(2), June 1996 Page 48

Pleistocene primate evolution on the continent.

Castor Cartelle, Instituto de Geociencias, Universidade
Federal de Minas Gerais, 31270-901 Belo Horizonte,
Minas Gerais, Brazil, and Walter Carl Hartwig,
Department of Anthropology, University of California,
Berkeley, California 94720, USA.


Cartelle, C. 1993. Achado de Brachyteles do Pleistoceno
Final. Neotropical Primates 1(1): 8.
Cartelle, C. and Hartwig, W. C. 1996. A new extinct
primate among the Pleistocene megafauna of Bahia,
Brazil. Proc. Nat. Acad. Sci. 93:6405-6409.
Hartwig, W. C. 1995. A giant New World monkey from
the Pleistocene of Brazil. J. Hum. Evol. 28: 189-195.
Hartwig, W. C. and Cartelle, C. 1996. A complete skel-
eton of the giant South American primate
Protopithecus. Nature, Lond. 381: 307-311.


In Argentina the black howler monkey, Alouatta caraya,
inhabits upland semideciduous forests, and flooded
forests on the islands of the large rivers, Parand and
Paraguay (Cabrera, 1939; Brown and Zunino, 1994).
Field studies done on this species in these habitats
revealed differences in density, social organization, diet,
and behavior, related to the floristic structure (Rumiz,
1987, 1990; Rumiz et al., 1986; Zunino, 1986, 1989;
Bicca-Marques, 1994; Brown and Zunino, 1994).

Currently, A. caraya is not considered as threatened
(Emmons, 1990; Rylands et al., 1995). However, in
Argentina the continuous extensive deforestation for
timber, land use, and dam construction, suggests a
progressive degradation and reduction of the habitat
available for this species. The aims ofthe project reported
here consisted in the definition of three types of habitat
occupied by A. caraya, evaluating as such the relation
between habitat and population characteristics and the
effects of alteration.

Study Site and Methods

The study was carried out in the northwest of the
Corrientes Province, Argentina (270 30'S, 58' 50'W),
comprising riparian forest patches along the Riachuelo
river, and flooded forest in the Parana river.

In the Riachuelo river area, the forest is a mosaic of tall

and low patches of about 10 ha separated by grasslands.
The forest patches show different degrees of alteration,
some suffering a selective logging while in others most
of the trees have been eliminated.

On the islands, the terrain is inundated almost yearly
defining as such the floristic structure which is
characterized by the presence of fast-growing tree
species (Rumiz et al., 1986). Occasionally, floods persist
over long periods resulting in the loss of trees, and
reducing dramatically the presence of howler monkeys.
Due to the low quality of the timber, and the fact that
the soils are not suitable for agriculture, logging is not a
serious problem.

To compare the habitats, inventories of trees were carried
out in three sites inhabited by the howler monkeys. On
the Riachuelo river, a tall forest patch was selected which
had suffered little alteration (BPA), along with a second
which had been heavily exploited (BMA). The criteria
used to select these sites were based on qualitative
evaluations of the tallness of the canopy, the presence
of species of economic importance, abundance of thin-
trunked trees, and evidence of exploitation. The study
of the flooded forest patch (SI) was carried out on the
island of Brasilera, near the confluence of the Parana
and Paraguay rivers.

The species, height, and diameter at breast height (DBH)
were recorded for all trees with a DBH greater than 10
cm. They were plotted in quadrats of 10 x 10 m. For
comparisons between habitats we employed the
following variables: NiH>10 m = Number of trees
belonging to each species with a height greater than 10m;
DBHm = Mean DBH; Nsp = Number of species; Ni/
Nsp = Number of individuals of each species with respect
to the total number of species; and NiDBH>20 cm =
Number of individuals with a DBH greater than 20 cm.
We also calculated the density, and used Shannon's index
as a measure of diversity. Differences between habitats
were analyzed by applying a discriminant analysis. To
evaluate the effect of logging, we compared our results
with previous inventories of 174 ha at BPA and BMA
in 1987.

The density of howler monkeys was estimated in BPA,
BMA, and SI. Daily censuses by transect were carried

Table 1. Habitat characteristics. Mean values and SD of tree density
expressed as individuals per ha (Di); Total number of species (Nsp);
Shannon's Index (H'); Diameter at breast height (DBH); Mean height
(Hm); BPA: Unexploited forest; BMA: Disturbed forest; SI: Flooded
Site Di Nsp H' DBH Hm
BPA 500 (26.8) 20 4.61 16.68 (7.5) 6.58 (1.6)
BMA 745 (75.8) 14 1.92 19.95 (2.9) 5.99 (1.5)
SI 300(76.5) 7 1.15 32.47 (11.4) 15.15(3.6)

Neotropical Primates 4(2), June 1996

Page 48

Neotropical Primates 4(2), June 1996

Table 2. Results of classification among the three habitats.
BMA 20 70.0 25.0 5.0
BPA 19 36.8 63.2 0.0
SI 29 0.0 6.2 93.1

out during at least three weeks at each site. The area
surveyed was estimated by multiplying the distance
traveled by 40 m. The estimated density values were
obtained applying the Non-Linear Density Plot Method
(Struhsaker, 1981).

Results and Discussion

The floristic analysis (Table 1) showed that the BPA
and BMA had more slender trees than the SI. BMA
showed the greatest abundance of trees with a DBH
between 10 and 30 cm, with a few individuals in the
largest categories. At BPA we also observed a
predominance of slender trees (DBH=10-30 cm), but
some larger trees were also present. SI showed the
highest frequency of large trees, with a peak of DBH
between 20 and 50 cm.

Small trees were most abundant at BMA; almost 50%
of those recorded were between 4-6 m. BPA also showed
a high frequency of low trees, with more than 40%
grouped between 6-8 m. SI, on the other hand, showed
the greatest spread, with individuals ranging in height
from eight to 24 m.

The most diverse of the three plots was BPA due to the
high number of tree species. The diversity values in
BMA were approximately half those of BPA, suggesting
the possibility that a wider range of food sources was to
be found in the less altered environments. At SI,
however, diversity was lower than in the other two
habitats, due to fewer species. Finally, the highest density
values were obtained at BMA, were found to be
intermediate in BPA, and lowest in SI.

The vegetation quadrats were grouped in 3 OTUS, each
corresponding to the previously defined habitats. The
first discriminant function explained 95% of the
variance, and suggested that the first two functions could
explain the relationships between the ecological factors.
The variables with greatest discriminant value were: Hm,
Ni, and Ni/Nsp on the first function, and Hm on the
second. The first discriminant function separated most
of the SI quadrats with respect to BPA and BMA (Table
2). However, the analysis did not discriminate clearly
between BPA and BMA. The partial overlap suggested
a degree of similarity between them. This was expected
due the common origin of the two areas, differentiated
only by the degree of exploitation.

Regarding the howler census, we found that the mean

Table 3. Population characteristics of black howler monkeys. Mean
values (SD), and comparisons by ANOVA among habitats.
Variable BMA BPA SI P
Density (inds./ha) 0.50 (0.17) 0.88 (0.18) 2.37 (0.74) <0.01
Group size 4.60 (1.42) 7.80 (1.64) 13.00 (7.10) <0.01
Sex ratio 1.60 (0.62) 2.00 (0.70) 1.90 (0.60) >0.05

group size was significantly lower at BMA, and only
slightly smaller at BPA than at SI (Table 3). However,
larger groups were more frequent at SI than at BPA.

The density values showed significant differences
between habitats, the lowest was at BMA, and the highest
at SI. In spite of the differences in density and group
size, sex ratio was not significantly different between
habitats. The sex ratio appeared as a constant,
independent of the environmental differences, but with
many multimale groups in SI and a predominance of
unimale groups in the other habitats.

A continuous reduction in quality and size of the useful
habitat for howlers was observed for the Riachuelo area
between 1987 and 1994. Currently 21.14% of the 123
ha of BPA studied in 1987 is being exploited, which is
consequently becoming more similar to BMA, and
considering the 174 ha of habitat comprising BPA and
BMA, 16.1% has been eliminated altogether.
Considering the islands occupied by howlers, the
recently inaugurated dam of Yaciret6 has eliminated
habitats containing between 4000 and 14000 howlers
due to inundation (Neris et al., 1994; Zunino and Ruiz,

The density of howler monkeys in the flooded forest is
higher than has been recorded for other populations of
Alouatta living in tropical environments (Crockett and
Eisenberg, 1987). The flooded forest offers a less diverse
diet (Brown and Zunino, 1994), but seasonality is less
marked than in the terra firm forest patches at the same
latitude (Rumiz et al., 1986). The availability of potential
food resources in the flooded forest appears to be more
uniform in space and time.

In the last decades forest exploitation in the study site
has resulted in a loss of habitat for the black howler
monkeys, reducing the populations and increasing the
isolation of the groups. If this trend continues in the
future, the mainland population will be seriously
threatened. The status of A. caraya in flooded forest,
however, remains uncertain. Other projects threaten this
environment, such as the Corpus and ParanA Medio dams
and the waterway project that intends to permit
navigation as far as Bolivia. Protected areas where these
howlers occur are limited to the National Parks of Chaco
and Pilcomayo, with an estimated population of about
10,000 howlers, and none of them includes flooded
forests, the most important habitat for this species

Page 49

Neotropical Primates 4(2), June 1996

(Brown and Zunino, 1994).

Gabriel E. Zunino, Susana Bravo, Museo Argentino
de Ciencias Naturales "Bernardino Rivadavia", Avenida
Angel Gallardo 470, 1405 Buenos Aires, Argentina,
FlAvia Murad Ferreira, Aracruz Cellulose, Vit6ria,
Brazil, and Carolina Reisenman, Facultad de Ciencias
Exactas y Naturales, Universidad de Buenos Aires,


Bicca-Marques, J. C. 1994. Padrao de utilizago de uma
ilha de mata por Alouatta caraya (Primates: Cebidae).
Rev. Brasil. Biol. 54(1):161-171.
Brown, A.D. and Zunino, G. E. 1994. Habitat, densidad
y problems de conservaci6n de los primates de la Ar-
gentina. Vida Silvestre Neotropical 3(1):30-40.
Cabrera, A. 1939. Los monos de la Argentina. Physis
Crockett, C. M. and Eisenberg, J. F. 1987. Howler varia-
tions in group size and demography. In: Primate Soci-
eties, B. B. Smuts, D. L. Cheney, R. M. Seyfarth, R.
W. Wrangham and T. T. Struhsaker (eds.). pp.54-67.
The University of Chicago Press, Chicago.
Emmons, L. H. 1990. Neotropical Rainforest Mammals.
A Field Guide. The University of Chicago Press, Chi-
Neris de Colman, N. N., Placci, G., Ruiz, J. C. and
Zunino, G. E. 1994. Panel de experts en monos
caraya. Informe Final. Entidad Binacioal Yaciret6,
Ayolas, Paraguay. 16pp.
Rumiz, D. I. 1987. Ecologia poblacional de Alouatta
caraya en el norte de la Argentina. Tesis de Doctorado.
Universidad Nacional de La Plata, Buenos Aires,
Rumiz, D. I 1990. Alouatta caraya: Population den-
sity and demography in northern Argentina. Am. J.
Primatol. 21(4):279-294.
Rumiz, D. I., Zunino, G. E., Obregozo, M. L. and Ruiz,
J. C. 1986. Alouatta caraya: Habitat and resource uti-
lization in northern Argentina. In: Current Perspec-
tives in Primate Social Dynamics, D. M. Taub and F.
A. King (eds.), pp: 175-193. Van Nostrand Reinhold,
New York.
Rylands, A. B., Mittermeier, R. A. and Rodriguez Luna,
E. 1995. A species list for the New World primates
(Platyrrhini): Distribution by country, endemism, and
conservation status according to the Mace-Lande sys-
tem. Neotropical Primates 3(suppl.): 113-164.
Zunino, G. E. 1986. Algunos aspects de la ecologia y
etologia del mono aullador negro (Alouatta caraya)
en habitats fragmentados. Tesis de Doctorado,
Universidad de Buenos Aires, Buenos Aires, 152 pp.
Zunino, G. E. 1989. Habitat, dietay actividad del mono
aullador negro (Alouatta caraya) en el noreste de la

Argentina. Bol. Primatol. Latinoamericano 1(1):74-
Zunino, G. E. and Ruiz, J. C. 1995. Reintroducci6n y
translocaci6n de primates en la Argentina:
Antecedentes, riesgos y beneficios. In: Simposio:
Primatologia en Argentina, Reszimenes. X Jornadas
Argentinas de Mastozoologia, pp.77-78. Sociedad
Argentina para el Estudio de Los Mamiferos
(SAREM), 14-17 November, 1995, La Plata, Argen-



Division of labor based on age or size may reflect the
reproductive condition of individuals in social groups.
In 1967, West proposed the general hypothesis that
hierarchical relations may be advantageous to both
dominants and subordinates and that individuals of low
rank may be inferior reproductive who benefit
genetically from associations with and contributions to
reproductively superior individuals. Since increasing age
or size eventually entails decreasing reproductive value
(Vx), several authors have noted that the display of social
behavior, such as foraging behavior that benefits all
members of a group, should increase with age as the
benefits from individual (selfish) reproduction decline
(e.g., West-Eberhard, 1975; Hrdy and Hrdy, 1976). As
individual reproductive value decreases, benefits
(genetic or other) from assisting the reproduction of
conspecifics (social behavior) may increase because
costs (genetic or other) of social behavior decrease with
decreased benefits from individual reproduction. In order
to test this hypothesis, I studied the relationship between
adult female age, dominance rank, reproductive value,
and social foraging behavior (food search and pursuit)
for adult female mantled howler monkeys (Alouatta
palliata Gray).

Subjects and Methods

During an extended period of study at Hacienda la.
Pacifica, Cafias, Guanacaste, Costa Rica, I studied two
marked, aged groups of mantled howler monkeys in two
tropical dry forest habitats (see Jones, 1980; Table 1).
For this species, age and dominance rank are negatively
correlated in both sexes (Jones, 1978, 1980).

Foraging was operationally defined as the behavioral
series: feed-rest-move (at least 100m) feed, by a unit
of more than three adults. These criteria were adopted
in order to standardize measurement and to eliminate
periods of food search within unusually large patches

Page 50

Neotropical Primates 4(2), June 1996

and by consort pairs. I identified which females in the
primary study groups initiated foraging sequences and
analysed these observations by age.

My null hypothesis held that the frequency of foraging
by females of any age class would be proportional to
the total number of females who foraged in an age class.
Two of the 15 females in one group (both young adults)
were never observed to direct foraging sequences and
are excluded from analysis. Three females were aged
on the basis of physical and behavioral traits other than
tooth wear, and assignment to age classes for these
females was made independent of the present analysis.
Two of these females were observed from sub-adult
through adult growth and classified as young adults; a
third female, classified as middle-aged, was the mother
of a sub-adult and a juvenile offspring, a highly unlikely
combination for any other age class (see Glander, 1980).
In my analysis of the second group (eight adult females),
two young adult immigrant females were never observed
to forage socially and were excluded from analysis. The
pattern of results reported here would remain unaffected
by alternative treatments of the raw data.

A monthly foraging rate for each forager was computed
by dividing the frequency of foraging by the female's
number of months resident in a group, a period of time
varying. from 10-14 months since some females
emigrated during the study. These rates were compared
with a female's age class, on the one hand, and
dominance rank, on the other, to assess the relationship

2 5:


y'= 0.62x 0.67


** *

0 A MA MO 0
Age Class
Figure 1. Social foraging rate as a function of age for young adult
(YA), middle-aged (MA), middle-aged to old (MO) and old (0)
female howlers. Each point represents one adult female expect where
asterisk indicates two.
asterisk indicates two.

Table 1. Age class, estimated age in years, number of females in
each age class (N), observed (0) and expected (E) frequencies of
social foraging, and cumulative chi square (X2) values for a test of
the null hypothesis.
Age class N 0 E (O-E)'/E
Young adult (5-7) 5 15 42.4 17.71
Middle-aged (7-10) 5 35 42.4 1.29
Middle-aged to Old (10-15) 1 18 8.1 12.11
Old (15+) 1 33 8.1 76.54
Total 12 101 101.0 107.65

between the display of social foraging behavior and rank,
and reproductive value (Vx, population data in
Malmgren, 1979, Table 23; equation after Wilson and
Bossert, 1971) where the relative contribution to future
generations of an individual of a given age is quantified.

Results and Discussion

Table 1 presents the results of my analysis for the first
group of foraging frequency as a function of female age,
including expected frequencies. Computing "goodness
of fit" led to an unequivocal rejection of the null
hypothesis (P<0.001, x2 = 107.64, df= 3). Thus, old age
and foraging frequency are significantly related. Young
adult females initiate foraging significantly less than
expected on the basis of their numbers (P50.001),
suggesting that such individuals are relatively "selfish"
or are conserving time and energy, possibly for
reproduction or competition. Table I also shows that
the middle-aged to old female foraged more than
expected by chance (P50.01), and this female succeeded
the oldest and lowest ranking female as the most frequent
forager when the old female emigrated in 1977 (personal

Additional observations support the reliability of the
above patterns. The oldest female in the second group
foraged more frequently than any other (P< 0.001, X2 =
17.29, df = 2). Similarly, the relationship between
foraging rate and age class (Fig. 1) yields a significant
positive correlation (r = +0.629, p<0.05). Related to this,
the correlation between foraging rate and dominance
rank (Fig. 2) is significant but negative (i.e., the higher
the foraging rate, the lower the dominance rank, r =
-0.63, p50.05). Thus, the initiation of foraging is
significantly associated with female age and dominance

It was hypothesized above that the expression of social
behavior should increase with increasing age since
reproductive value (Fig. 3) decreases with age and with
it the benefits from selfish reproduction. Fig. 3 shows
the reproductive value curve for the population of howler
monkeys at Hacienda da Pacifica. Comparing Fig. 3 with
Figs. I and 2, consistent with expectation, a strong
negative association appears to exist between
reproductive value and rate of foraging. Reproductive

Page 51

Neotropical Primates 4(2), June 1996

y'= 0.199x + 0.153

2 4 6 8
Dominance Rank

10 12

Figure 2. Social foraging rate as a function of individual dominance
rank; each point represents one adult female. Note that low numbers
mean high rank.

value in the four adult age classes is negatively and
significantly correlated with social foraging rate/month
(r. = -0.95, p<0.02). These results support the view that
increasing age or size eventually entails decreasing
reproductive value and that the display of social behavior
should increase with age as the benefits from individual
(selfish) reproduction decline.

What features of the howlers' environment might favor
temporal division of labor? On 52 occasions, I was able
to record the specific resource upon which foraging
sequences terminated. Forty-four (85%) of these
sequences terminated on ephemeral food, while eight
(15%) sequences terminated with feeding on mature
leaves (P_0.001, X2 = 49, df= 1). Thus, the initiation of
foraging sequences appears to be associated with food,
the local distribution of which is temporally uncertain;
new leaves, flowers, and fruit. The old female initiated
21 of the 52 (40%) bouts, 20 of these for ephemeral

An old female's presumed experience with the mosaic
of her home range might enhance her efficiency as a
forager so that her foraging activity may yield an
energetic and nutritional gain to other group members.
Temporal uncertainty of preferred food resources may
favor individuals that are the beneficiaries of the foraging
activity of others when reproductive value is low.
Division of labor through differential social roles may
be a function of relative reproductive value, and
behavioral roles may be understood within the context
of life history patterns.

2 3 6 7 s 11 -12 13 14 15 M 1G 1 1 19 2 1 22 22 24 s
Age in Years
Figure 3. The reproductive value (V,) curve for the howler population
at Hacienda I a Pacifica, Cafias, Guanacaste, Costa Rica. Reproductive
value was computed for the mid-point of each age class since discrete
ages could not be determined year-by-year.


I appreciate the comments of R. C. Lewontin, E. 0.
Wilson, M.-J. West-Eberhard, I. S. Bernstein, W. C.
Dilger, and K.E. Weber on an earlier draft of this note. I
thank the W. Hagnauer family for permission to work
on their property, Hacienda La. Pacifica, and for logistic
assistance. The work was supported by grants from the
National Fellowships Fund and the National Research

Clara B. Jones, Institute of Animal Behavior, Rutgers
University Newark, 101 Warren Street, Newark, New
Jersey 07102, U.S.A.


Glander, K.E. 1980. Reproduction and population
growth in free-ranging mantled howling monkeys. Am.
J. Phys. Anthropol. 53:25-36.
Hrdy, S. B. and Hrdy, D. B. 1976. Hierarchical rela-
tions among female hanuman langurs (Primates:
Colobinae, Presbytis entellus). Science 197:913-915.
Jones, C. B. 1978. Aspects of reproductive behavior in
the mantled howler monkey, Alouatta palliata Gray.
Dissertation, Cornell University, Ithaca, New York.
Jones, C. B. 1980. The functions of status in the mantled
howler monkey, Alouattapalliata Gray: Intraspecific
competition for group membership in a folivorous
neotropical primate. Primates 21:389-405
Malmgren, L. A. 1979. Empirical population genetics
. of golden mantled howling monkeys (Alouatta
palliata) in relation to population structure, social dy-
namics, and evolution. Ph.D. Dissertation, University
of Connecticut, Storrs.
West, M. J. 1967. Foundress associations in polistine
wasps: Dominance hierarchies and the evolution of


Page 52

Neotropical Primates 4(2), June 1996

social behavior. Science 157:1584-1585.
West-Eberhard, M. J. 1975. The evolution of social be-
havior by kin selection. Quart. Rev. Biol. 50:1-33.
Wilson, E. 0. and W. H. Bossert. 1971. A Primer of
Population Biology. Sinauer Associates, Stamford,


Common marmosets (Callithrix jacchus) are endemic
to northeastern Brazil and live in a variety of habitat
types (Rylands et al., 1993). Previous research on the
behavior and ecology of this species has been restricted
to the semi-deciduous coastal forests of Permambuco,
Paraiba, and Rio Grande do Norte (e.g., Alonso and
Langguth, 1989; Digby and Barreto, 1993; Hubrecht,
1984; Scanlon et al., 1989). However, the majority of
the common marmoset geographic range encompasses
the very different vegetation found in the interior of
Brazil. Here we report on preliminary surveys of a
marmoset population in the northeastern Brazilian state '
of Piaui.

Vegetation of the Brazilian Northeast

The northeastern interior is dominated mainly by two
types of vegetation: the cerrado (savanna forest) and
the caatinga (dry thorn scrub). The cerrado covers over
2.01 million km2 within Brazil and is second only to the
Amazon forest in the area it covers (Rizzini et al., 1988).
The term cerrado (sensu lato) encompasses a wide range
of subtypes of xeromorphic vegetation from the campo
limpo (open grasslands) to the cerraddo (dense savanna
forests). Trees are semi-deciduous with broad and rigid
leaves and thick bark that allows them to survive frequent
savanna fires (Eiten, 1972). The caatinga covers an
addition 0.91-1 million km2 and is characterized by a
semi-arid climate. Herbs and grasses grow in the
caatinga only during the rainy season, and vegetation is
xerophytic or deciduous (Rizzini, 1977; Rizzini et al.,
1988). The flora of the Brazilian cerrado is estimated to
include 7,000 species compared to the 60,000 species
of the Amazon flora and 2,000 species of the flora found
in the northeastern caatinga (Castro, 1994).

The Study Site

Surveys were conducted at the Sete Cidades National
Park in the municipalities of Piripiri and Piracuruca, Piaui
(04 05-09'S, 41 30-45'W; alt. 100-300 m). The park
encompasses 6,221 ha and includes a small hostel,

restaurant, and administrative offices. The primary
tourist attractions in the park are a series of dramatic
rock formations and rock paintings (Brazil, IBDF/FBCN,

The park exists in a cerrado-caatinga transition zone
resulting in a mosaic of habitats. In relatively level areas
with good drainage, plant species characteristic of the
cerrado predominate [e.g., "lixeira" (Curatella
americana), "barbatimao" (Stryphnodendron
coriaceum), "cascudo" (Terminallafagifolia), "faveira-
de-bolota" (Parkiaplatycephala), and"piqui" (Caryocar
coriaceum)]. In areas with poor drainage (and subject
to flooding) open grassland is found, and along stream
beds, riparian forests of the cerraddo. Mixed into many
of these habitats are species characteristic of the riparian
forests [e.g., "jatobi-de-mata" (Hymenaea courbaril var.
stilbocarpa), "pau-marfim?' (Agonanadra brasiliensis),
and "pau-pombo" (Sclerolobium paniculatum or
Tapirira guianensis)] and species characteristic of
caatinga [e.g., "sabid" (Mimosa caesalpiniifolia), "pau-
d'Arco-de-sete-folhas" (Tabebuia aurea), "aroeira"
(Miracrodruon urundeuva), "macambira"(Bromelia
laciniosa), and "xique-xique-' Pilosocereus gounellei)
(Barroso and GuimbirAes, 1980).

Marmoset Surveys

Informal surveys were carried out during two periods:
July 1994 (three days; eight surveys of 2-5 hours
duration) and July 1995 (17 days, total of 66 hours of
surveys). Surveys involved one to four observers in five
different areas within the park. Particular attention was
paid to locating and identifying gum-producing plants
bearing characteristic marmoset gouge-holes.

Direct sightings or indirect evidence of common
marmosets were found in three areas. In Area 1, a patch
of cerraddo with no standing water, a group of at least
three individuals was sighted during the 1994 survey.
Vocalizations were heard in this same general area in
1995. Though the area contained trees known to be gum
sources for this species [e.g., "cajui" or cashew
(Anacardium occidentale var. microcarpum)], no trees
with gouge-holes were found. In Area 2, pristine riparian
forest in a section of the park closed to tourists, three
gum trees were found with gouge marks typical of those
created by the marmosets (see below). Area 3 consisted
of a semi-disturbed gallery forest adjacent to the park
office and hostel. Here, a marmoset group containing at
least seven individuals was followed for 9.5 hours over
10 days. The group consisted of three adults (at least
one male and one female), two juveniles (estimated at
6-7 months of age based on size and pelage), and two
infants (estimated at about one month of age). During
the brief observation period animals used approximately

Page 53

Neotropical Primates 4(2), June 1996 Page 54

one to 1.5 ha of an estimated 15 hectare patch of forest.

A total of eight individual trees of four species were
found with gouge holes typical of those produced by
marmosets. Three cashew trees (A nacardium occidental
var. microcarpum) were found marked. One (in Area 3)
was seen being used by the marmosets and was heavily
marked with fresh gouges. The other two trees contained
only older, dry gouges. Three "pau terra" trees (Qualea
grandiflora and Q. parviflora) were identified. Again,
marmosets in Area 3 were observed feeding on one of
the three trees. This is the first record of this genus being
used by common marmosets, though both species have
been reported as a food source for the black-tufted
marmoset (Callithrix penicillata) (Fonseca and Lacher,
1984). One "jatoba-de-chapada" (Hymenaea stigono-
carpa) was found with only three gouges, and it is
unlikely that this tree was used as a food source. The
fourth species was unidentified, but was heavily marked
with old, dry gouges. Overall, while several gum trees
were found, their apparent densities were much lower
than those found in the Atlantic coastal forests (e.g.,
Alonso and Langguth, 1989; Scanlon et al., 1989).


Marmosets have become increasingly well known for
their unusual social organization and variable
reproductive strategies. While we have been able to
broaden our understanding and documentation of their
behavior, it is currently unclear what ecological factors
may be responsible for this flexibility. Seasonality in
food availability is likely to affect marmoset population
densities and home range sizes which, in turn, are likely
to have profound effects on the social organization and
reproductive strategies of this species. As the seasonal
caatinga and cerrado habitats make up the majority of
the common marmoset geographic range, a long-term
study of the groups at the Parque Nacional de Sete
Cidades should add considerably to our understanding
of this species.

Preliminary results from Sete Cidades already indicate
some key differences between this marmoset population
and those found in the coastal forests. Of particular note
is the apparent low density of trees being used as gum
sources by the marmosets (less than two gouged trees/
hectare in Area 3). In comparison, Scanlon et al. (1989)
found a minimum of 54 gum-producing trees per hectare
(most with gouge marks), and Alonso and Langguth
(1989) identified 25 gum-producing trees in their group's
home range (5 trees/hectare; all with gouged holes). The
relatively low density of gouged trees at Sete Cidades
suggests that the marmoset themselves are at low
densities, or that they rely on other food sources. The

low frequency of visual or auditory contacts with animals
during surveys supports the former, that groups are
indeed at low densities compared to coastal populations
(e.g., Digby and Barreto, 1993; Scanlon et al., 1989).
The density of gouged trees and marmoset groups at
Sete Cidades are also low in comparison to those for the
black-tufted-ear marmoset (C. penicillata) (Fonseca and
Lacher, 1984; Faria, 1984). Additional behavioral
observations will be necessary before it can be
determined whichalternative food sources area available
to the Sete Cidades marmosets.

These results, while preliminary, already suggest
important differences in the ecology of common
marmosets living in the cerrado-caatinga transition zone
compared to groups living in the coastal forests. The
confirmation of the presence of common marmosets at
the Sete Cidades National Park is a further important
step in the understanding of this species' ecology, in
particular because of the lack of protected areas in the
semi-arid caatinga and cerrado habitats within its
geographical range. Of the seventeen conservation units
(excluding two that contain "possibly introduced and
mixed populations") cited for the species by Rylands et
al. (1993), only three protect semi-arid habitats. Despite
its ecological flexibility, natural populations of common
marmosets are increasingly vulnerable to habitat
destruction. The protection of a relatively large area of
native habitat such as that at Sete Cidades will be
important for the conservation of common marmosets
(and that of other cerrado/caatinga fauna) over the long
term, as will the collection of more detailed data on the
behavior and ecology of this species.


Fieldwork in the Parque Nacional de Sete Cidades was
authorized by IBAMA Piaui. We are especially grateful
to our field assistants Elena Davis, William Brandao
Pimentel, and Jose Orlando Soares Oliveira. We also
thank Eug6nia Vit6ria e Silva de Medeiros, Francisco
Celso de Medeiros, Sandra Dantas, and Jos6 Arribamar
de Carvalho for their assistance. This project was funded
by Duke University Research Council and the Duke-
UNC Program in Latin American Studies.

Leslie Digby, Biological Anthropology and Anatomy,
Duke University, Box 90383, Durham, North Carolina
27708, USA, Stephen F. Ferrari, Departamento de
Psicologia, Universidade Federal do Para, CP 8607,
66.075-150 Beldm, ParA, Brazil, and A. Alberto Jorge
F. Castro, N6cleo de Referencia em Ciencias
Ambientais do Tr6pico Ecotonal do Nordeste
(TROPEN), Programa de Fitodiversidade e
Fitossociologia dos Cerrados Marginais do Nordeste

Neotropical Primates 4(2), June 1996

Page 54

Page SS Neotropical Primates 4(2), June 1996

(FITCEM), Universidade Federal do Piaui, Av.
Petr6nio Portela 1310, Campus de Ininga, 64.003-600,
Teresina, Piaui, Brazil.


Alonso, C. and Langguth, A. 1989., Ecologia e
comportamento de Callithrixjacchus (Callitrichidae,
Primates) numa ilha de floresta Atantica, Rev.
Nordestina Biol. 6:105-137.
Barroso, G. M. and Guimaraes, E. F. 1980. Excursao
botanica ao Parque Nacional de Sete Cidades, Piaui.
Rodrigugsia 32(53):241-267.
Brazil, IBDF/FBCN. 1979. Plano de Manejo Parque
Nacional de Sete Cidades. Institute Brasileiro de
Desenvolvimento Florestal (IBDF) and FundaqAo
Brasileira para a Conservacao da Natureza. FBCN,
Castro, A. A. J. F. 1994. Comparaqao floristica de
esp6cie do cerrado. Silvicultura 15(58):16-18.
Digby, L. L. and Barreto, C. E. 1993. Social Organiza-
tion in a wild population of common marmosets
(Callithrixjacchus). Part I: group composition and
dynamics. Folia Primatol. 61:123-134.
Eiten, G. 1972. The cerrado vegetation of Brazil. Bo-
tanical Rev. 38(2):201-341.
Faria, D.S. de. 1984. Uso de arvores gomiferas do
cerrado por Callithrix jacchus penicillata. In: A
Primatologia no Brasil, M. T. de Mello (ed.), pp. 83-
96. Sociedade Brasileira de Primatologia, Brasilia.
Fonseca, G. A. B. da and Lacher, T. E., Jr. 1984. Exu-
date feeding by Callithrix jacchus penicillata in
semidecidous woodland (cerraddo) in central Bra-
zil. Primates 25:441-50.
Hubrecht, R. C. 1985. Home-range size and use and
territorial behavior in the common marmoset,
Callithrixjacchusjacchus, at the Tapacurd field sta-
tion, Recife, Brazil. Int. J. Primatol. 6:533-550.
Rizzini, C. T. 1979. Tratado de Fitogeografia do Bra-
zil: Aspectos Sociol6gicos e Floristicos. HUCITEC/
EDUSP, Sao Paulo.
Rizzini, C. T., Coimbra Filho, A. F. and Houaiss, A.
1988. Ecossistemas Brasileiros/Brazilian Ecosystems.
Editora Index, Rio de Janeiro.
Rylands, A. B., Coimbra-Filho, A. F. and Mittermeier,
R. A. 1993. Systematics, geographic distribution, and
some notes on the conservation status of the
Callitrichidae. In: Marmosets and Tamarins: System-
atics, Behaviour, and Ecology, A. B. Rylands (ed.),
pp.11-77. Oxford University Press, Oxford.
Scanlon, C. E., Chalmers, N. R. and Monteiro da Cruz,
M. A. 0. 1989. Home range use and the exploitation
of gum in the marmoset Callithrixjacchusjacchus.
Int. J. Primatol. 10:123-136.


Located in northwestern Rond6nia (Figure 1), the 99,813
ha Estagao Ecol6gica Serra dos Tres Irmaos was decreed
in 1990 as part of statewide network of conservation
units. Tres Irmaos is the only component of this network
located on the left or west bank ofthe Rio Madeira, which
plays an important role in the zoogeography of the
region's primates (Rylands and Bernardes, 1989; Ferrari
and Lopes, 1992), in addition to a number of other
mammals (Emmons, 1990). Most of these taxa are not
found elsewhere in Rond6nia, which emphasizes the
importance of this Ecological Station's role in the
conservation of the biodiversity of this state, one of the
most intensely-colonized areas of Brazilian Amazonia.

Two different areas of the Station were surveyed in
October and December 1995 in order to identify its
diurnal mammal species and evaluate their population
densities. Nine primate species were observed during
these surveys (Table 1). In addition to nocturnal
sightings, a group of four owl monkeys was seen in
activity on one occasion at mid-morning. A tenth species
not observed during surveys, Alouatta seniculus, was
encountered on the left bank of the Madeira, 5 Km from
the southern limit of the Ecological Station.

Local residents interviewed all reported that howlers are
found only in areas close to the Rio Madeira. This,
together with the lack of any indirect evidence
(vocalizations or feces) of the occurrence of Alouatta
within the Ecological Station, which at its closest point
is 3 km from the Madeira, indicates that the distribution
of A. seniculus in this area may be restricted to a
relatively narrow corridor, perhaps less than a kilometer
in width, on the left bank of this river. Howlers are
nevertheless more widespread further downstream
(Ferrari and Lopes, 1992).

A similar distribution was indicated by local residents
for two other species not observed during the present
study Ateles belzebuth and Cebuella pygmaea. The
presence of Ateles would be expected from its known

Table 1. Primates observed in the Tres Irmaos
Ecological Station, Rond6nia.
Aolus nigriceps
Callicebus caligalus
Cebus albifrons
Cebiis apella
Lagolhrix lagotricha cana
Pithecia irrorata
Saguinus fiuscicollis weddelli
Saguinus labiatus labiatus
Saimiri sciureuss) boliviensis

Page 55

Neotropical Primates 4(2), June 1996

Neotropical Primates 4(2), June 1996

distribution (Kellogg and Goldman, 1944), but that of
Cebuella, while not improbable (Rylands et al., 1993),
would constitute an important extension of its
geographical range. It is hoped further fieldwork, planned
for 1996, will not only confirm the occurrence of these
two species in the area, but will also provide insights into
the factors determining their local distribution, and that
of others such as Alouatta seniculus.

Two-hundred kilometers of line transect censusing were
carried out during the present study, during which all but
two species A. nigriceps and C. albifrons were
recorded. A third species, S. boliviensis, was sighted on
only one occasion. The most abundant species were L.
lagotricha, P. irrorata, S. fuscicollis and S. labiatus,
which together contributed 86.4% of sightings. Lagothrix
appeared to be particularly abundant at the site, a good
indication of a lack of hunting within the reserve.

The relative scarcity of the Cebus species, normally
among the most abundant primates in western Amazonian
communities, whether hunted or not (Peres, 1990), raises
some interesting questions, especially in the light of the
local distribution of Alouatta, for example. Pithecia, on
the other hand, was recorded twice as frequently as Cebus
at Tres Irmaos, the opposite of the situation recorded at
most other western Amazonian sites.

Fortunately, the Serra dos Tres Irmaos Ecological Station
is relatively isolated from Rond6nia's principal areas of
human colonization, which lie to the east/south of the
Rio Madeira. The Station is accessible only by boat, and
appears to suffer little encroachment, except by local
fisherman. The results of the present study nevertheless
indicate the need for the extension of the Station's limits
to the left bank of the Madeira in order to protect fully
the area's mammalian communities. This has been
recommended to the state environment secretariat, and is
currently under study.

The fieldwork reported here was supported by the United
Nations Development Program (PNUD) and SEDAM -
Rond6nia. We would also like to thank Jorge Lourengo,
Renato Cintra, Hdlio Pensador, and local residents within
the study area.

Stephen F. Ferrari, Ernesto H. Cruz Neto, Simone
Iwanaga, H. Kitia M. Correa, Universidade Federal
do Para, Caixa Postal 8607, 66075-150 Bel6m, Pard, and
Paulo C. S. Ramos, PNUD/SEDAM, Rond6nia, Brazil.


Emmons, L. H. 1990. Neotropical Rainforest Mammals
A Field Guide. Chicago University Press, Chicago.
Ferrari, S. F., and Lopes, M. A. 1992. New data on the

distribution of primates in the region of the confluence
of the Jiparand and Madeira rivers in Amazonas and
Rond6nia, Brazil. Goeldiana Zool. 11:1-12.
Kellogg, R., and Goldman, E. A. 1944. Review of the
spider monkeys. Proc. U. S. Nat. Mus. 96:1-45.
Peres, C. A. 1990. Effects of hunting on western Ama-
zonian primate communities. Biol. Conserv. 54:47-59.
Rylands, A. B., and Bernardes, A. T. 1989. Two prior-
ity regions for primate conservation in the Brazilian
Amazon. Primate Conservation (10):56-62.
Rylands, A. B., Coimbra-Filho, A. F. and Mittermeier,
R. A. 1993. Systematics, geographic distribution, and
some notes on the conservation status of the
Callitrichidae. In: Marmosets and Tamarins: System-
atics, Behaviour, and Ecology, A. B. Rylands (ed.),
pp. 11-77. Oxford University Press, Oxford.


In 1985, a family of common marmosets was moved
from a laboratory setting to a "wild" environment
(Chamove and Rohrhuber, 1989). The group was
composed of a pair of 2-year-old laboratory-born
common marmosets (Callithrixjacchus) and their first
set of laboratory-born twin sons (9 months old). The
four lived together in a wire-mesh cage 3 x 2.1 x 1.4 m
prior to release and were fed on a normal laboratory
diet. Soon after moving to the garden, twins were born
and were 1.2 months old at the time of this observation.
The outside area included what was once a walled garden
- long neglected containing trees, shrubs, and vines.
There was continuous woodland for several kilometers
and the animals could move throughout a wide area
without needing to go to the ground. Ivy covered most
of the wall and extended out from it over 1.4 m in a
tangle of old and new stems. Toads (Bufo vulgaris) and
semi-wild domestic cats were also seen, but were never
observed being approached by the monkeys. This is in
contrast to Kleiman et al. (1986) who reported that lion
tamarins showed great interest in toads. (Presumably
other indigenous Scottish wildlife were present although
not seen). Upon release, the marmoset family appeared
to adapt quickly (Wendt, 1962). The most striking
change in the behavior of the animals was the branch
type they chose to use. When in the cages they spent
most of the time on flat mesh surfaces and horizontal
branches with infrequent, brief (0.8/min) visits to the
floor. Unrestricted outside, they spent most of the time
(89%) in the dense network of thin flexible ivy vines,
where they could not be seen at a distance. They rarely
visited more open shrubs (10%) or trees (1%). The
monkeys were never observed on the ground. Although
having no prior experience with gums, the monkeys were
regularly observed feeding from gouges they had made

Page 56

Neotropical Primates 4(2), June 1996

in elm trees (Elnus) preferring those of smaller girth.

In the laboratory, the monkeys reacted to soaring birds
as well as to aeroplanes, giving alarm calls and then
approaching the skylights to search for the bird once it
had disappeared. On the outside the monkeys responded
to soaring birds of all kinds, gulls were common, by
leaping into dense bushes and remaining still. The
marmosets appeared to ignore a large rubber snake
located on the ground or in the branches, even when the
head was made to move [see Heymann, 1987]. I found
a dead weasel (Mustela nivalis) that had been flattened
laterally by a car, and wedged its dry form into some
branches in a life-like position. Three of the four
marmosets mobbed the animal, giving alarm calls and
directing threats at the predator. The father, carrying the
1-month-old babies, approached most hesitantly,
remaining about 1.5 m away; the adult female
approached closest, to within 10 cm, and appeared to be
the most active in the mobbing. Surprisingly, the juvenile
males were not the most vigorous mobbers (Millar,
Evans, and Chamove, 1988). After about five minutes,
and when the weasel did not move off, the marmosets'
interest decreased. They moved away, still giving
sporadic alarm calls, and looking back at the immobile

When the study was published, there were few reports
of responses to predators of South American primates,
and we reported the response of the marmosets as being
presumably abnormal, maladaptive, and unlike what
would be expected from wild animals. The report by
Philips (1995) seems to suggest that this might not be
the case, with monkeys approaching predators more
closely than humans would judge as safe. Philips' white-
faced capuchins were mobbing a tayra, approaching to
within 2 m. Only one animal approached, but it was one
of the two adult males. The monkey (female) carrying
an infant did not approach closely. The remaining group
members were intermediate in distance. Just like the
tayra, our weasel made no aggressive response towards
the monkeys in response to their mobbing. Could it be
that there is a single animal that is the prime defender in
a group; that is has the "role" of defender (Chamove,

Arnold S. Chamove, Psychology Department, Massey
University, Palmerston North, New Zealand.


Chamove, A. S. 1983. Role or dominance in macaque
response to novel objects. Motivation and Emotion 7:
Chamove, A. S. and Rohrhuber, B. 1989. Moving
callitrichid monkeys from cages to outside areas. Zoo

Biol. 8: 151-163.
Heymann, E. W. 1987. A field observation of predation
on a moustached tamarin (Saguinus mystax) by an
anaconda. Int. J. Primatol. 8:193-194.
Kleiman, D. G., Beck, B. B., Dietz, J. M., Dietz, L.,
Ballou, J. D. and Coimbra-Filho, A. F. 1986. Conser-
vation program for the golden lion tamarin: Captive
research and management, ecological studies, educa-
tional strategies, and reintroduction. In: Primates: The
Road to Self-sustaining Populations, K. Benirschke
(ed.), pp.413-444. Springer Verlag, New York.
Millar, S. K., Evans, S and Chamove, A. S. 1988. Old-
est offspring contact novel objects soonest in
callitrichid families. Biology of Behaviour 13:82-96.
Phillips, K. 1995. Differing responses to a predator (Eira
barbara) by Alouatta and Cebus. Neotropical Primates
Wendt, H. 1962. Erfolgreiche Zucht des
Baumwollkoepfchens oder Pincheaeffchens,
Leontocebus (Oedipomidas) oedipus, in
Gefangenschaft. Saugetierkiindliche Mitteilungen,


Thermotolerant campylobacters have been shown to be
one of the most important etiological agents of acute
enteritis in humans, but in other mammalian species the
bacteria are present in an apparently healthy carrier-state
in the majority of cases (Rosef et al., 1983). In order to
determine the importance of wild primates as reservoirs
of these zoonotic microorganisms, rectal swabs were
obtained from a total of 43 individuals representing nine
species (Table 1) from different areas in the vicinity of
the town of Iquitos.

All samples were immediately placed into the transport
and enrichment medium (Fernandez, 1992) and
cultivated within eight hours on modified Skirrow's
medium (Fernandez, 1983), at 420C for 48h, in
microaerophillic conditions. Suspected colonies were
identified (Luechtefeld et al., 1981b) using catalase and
oxidase tests (both positive) and the morphological
features observed in Gram-stain (curved S-shaped rods).
Later, the thermotolerant Campylobacter species were
identified using the criteria proposed by Lior (1984) and
Goossens and Butzler (1992).

Campylobacters were isolated fron 9 (20.9%) of the
animals studied (Table 1). However, none of the animals
showed signs of enteritis or other illness. This isolation
rate was higher than that reported by Luechtefeld et al.

Page 57

Neotropical Primates 4(2), June 1996

Table 1. Prevalence of fecal cultures positive for thermotolerant campylobacters in wild primates.
Species No. of individuals Culture positive Culture negative Total no. positive
C jejuni C coli C lari
Saguinus labiatus 12 1 2 0. 9 3 (25,0)
Saguinus mystax 7 0 2 0 5 2 (28.6)
Pithecia monachus 6 0 2 0 4 2 (33.3)
Saimiri sciureus 4 0 0 0 4 0
Lagothrix lagotricha 4 0 0 0 4 0
Cebus apella 4 1 1 0 2 2(50.0)
Cebus albifrons 2 0 0 0 2 0
Ateles paniscus 2 0 0 0 2 0
Aotus sp. 2 0 0 0 2 0
Total 43 2 (22.2)* 7 (77.8)* 0 34 9 (20.9)
Numbers in parentheses = %. Numbers in parentheses with an asterisk = % of isolates.

(1981a) (9.3%) and similar to the results obtained by
FernAndez et al. (1987) in Brazil (19.0%). As reported
previously (Russell et al., 1988; Gozalo et al., 1991),
and supported by the present study, C. coli was the
predominant bacteria isolated from the primates.

It is highly likely that mammalian species differ in their
susceptiblility to intestinal colonization by C. jejuni and
C. coli, regardless of the degree of exposure to these
bacteria. This circumstance may help to explain the
different carriage rates detected in this and similar studies
(Rosefet al., 1983). Our data provide evidence that wild
primates from Iquitos appear to be important reservoirs
and infection sources of these bacteria for man. Further
studies are required to clarify and understand the
epidemiology of campylobacteriosis that is evidently a
complex phenomenon in developing countries.

Alvaro Tresierra-Ayala, Freddy Espinoza, Maria E.
Bendayan, Alfonso Bernuy and R6ger Ruiz Romero,
Departamento Acaddmico de Microbiologia.
Universidad Nacional de la Amazonia Peruana, Apartado
751, Iquitos, Peru.

Escamilla, J. 1991. A survey for Campylobacter in feral
and captive tamarins. Elsev. Sc. Publ., B.V. 675-676.
Lior, H. 1984. New, extended biotyping scheme for
Campylobacter jejuni, Campulobacter coli and
"Campylobacter laridis". J. Clin. Microbiol. 20:636-
Luechtefeld, N.W., Cambre, R.C. and Wang, W.L.L.
1981. Isolation of Campylobacter fetus subsp. jejuni
from zoo animals. J Am. Med. Ass. 199:1119-1122.
Luechtefeld, N.W., Wang, W.L.L., Blaser, H.J. and
Reller, L.D. 1981. Campulobacter fetus subsp. jejuni:
background and laboratory diagnosis. Lab. Med.
Rosef, 0., Gondrosen, B., Kapperud, G. and Underbal,
B. 1983. Isolation and characterization of
Campylobacter jejuni and Campylobacter soli from
domestic and wild mammals in Norway. Appl.
Environ. Microbiol. 46:855-859.
Rossell, R.G., Krugner, L., Tasai, C. and Ekstrom, R.
1988. Prevalence of Campylobacter in infant, juve-
nile and adult laboratory primates. Lab. Anim. Sci.

References NEWS

Fernindez, H. 1983. Thermophilic species of
Campylobacter: bacteriological, epidemiological and
pathogenical aspects. Unpubl. Doctoral thesis, Escola
Paulista de Medicina, Sao Paulo.
FernAndez, H. 1992. Increase of Campylobacter isola-
tion rates using an enrichment medium. Rev.
Microbiol. (S. Paulo). 23:5-7.
FernAndez, H., Rocha, M.B. and Trabulsi, L.R. 1987.
Ocorrencia de Campylobacter jejuni em animals do
zool6gico Rev. Fac. Med. Vet. Zootec. Univ. S. Paulo,
Goossens, H. and Butzler, J.P. 1992. Isolation and iden-
tification of Campylobacter spp. In: Campylobacter
jejuni: Current Status and Future Trends, 1.
Nachamkin, J. M. Blaser and L. S. Topkins (eds.).
American Society for Microbiology, Washington, D.C.
Gozalo, A., Block, K., Montaya, E., Moro, J. and


The existence of a new and undescribed marmoset near
the Rio Madeira was first brought to our attention in
June 1994 by Dr. Jose MArcio Ayres of the Wildlife
Conservation Society, New York. Information available
at the time indicated that it occurred in the region of the
Rio Abacaxis in the east of the state of Amazonas. As a
result, we organized an expedition to the area, made
possible through collaboration with Drs. Horacio
Schneider and J6lio Pieczarka of the Department of
Genetics of the Federal University of Para, and Dr. Jose
Augusto Muniz of the National Primate Center, National
Health Foundation, Beldm. The expedition was financed
by the John D. and Catherine T. MacArthur Foundation

Page 58

Page 59 Neotropical Primates 4(2), June 1996

and a Manaus-based tourist company, Amazon Ecopark
Hotdis e Turismo Ltda.

Further information was first obtained in the town of
Olinda do Norte. Describing what we knew of the
species, a local inhabitant indicated the Rio Canuma, as
the region where it occurred. Arriving at the mouth of
Rio Canuma, we obtained immediate confirmation of
the existence of the monkey, with a local trader, Sr. Pedro
Coelho, having two pet marmosets in his backyard, one
being Callithrix chrysoleuca from the west bank of the
Rio Canumd and the other the new species, which he
reported occurred on the east bank, where he lived. We
stayed in the area for four days, time enough to collect
two specimens of the new species. We also observed C.
chrysoleuca at Santa Barbara on the left bank of the

From there, we traveled to the Parand Uraria in the
direction of the Rios Abacaxis and Marimari, where we
were able to observe and collect specimens of the new
species as well as Callithrix mauesi, first described by
Mittermeier, Schwarz and Ayres in 1992, at the localities
of Santa Maria, Abacaxis and Sdo Jodo.

The description of this new species will be published in
the near future in the periodical Goeldiana Zoologia, a
publication of the Museu Paraense Emilio Goeldi,
Bel6m, with financial support from, Washington, D. C.
Conservation International. Stephen D. Nash of the State
University of New York kindly provided the illustrative

The new taxon has been named Callithrix saterei Silva
Jr. and Noronha 1996 after the Satere-Mauds Indians of
the region. The type series holotypee and six paratypes)
has been placed in the mammal collection of the Museu
Paraense Emilio Goeldi (MPEG), represented by the
series MPEG23955-23961. The type locality is "the
mouth of the Rio Canuma, right bank ofthe Rio CanumA,
municipality of Borba, Amazonas, Brazil (030 59'S, 59
05'W)", and the type series comes from three localities.

Callithrix saterei was assigned to the bare-eared
"argentata" marmoset group of Hershkovitz (1977), and
is evidently allopatric with regard to its nearest relations,
C. argentata, C. leucippe, C. melanura, C. emiliae and
C. nigriceps. It is, however, larger (average weight 430
g) and presents some autapomorphic traits, especially
with regard to an unusual morphology of the genitalia
of both sexes, and a combination of pigmented black
ears but loss of pigmentation on the face. It also has a
number of synapomorphies with the species C. nigriceps,
C. emiliae and C. intermedia to the south.

The conservation status of C. saterei remains unknown.

However, a large part of its geographic range is covered
by the Coati-Laranjal Indigenous Area of the National
Indian Foundation (FUNAI), shared by the Sater8-Mauds
and Mundurucus Indian tribes. This implies indirect
protection for the region's fauna and flora. In the area
we surveyed, outside the Indigenous area, it appears to
be a common species in secondary terra-firme forest in
varying stages of succession, as well as seasonally
inundated black-water forest (igap6), and even near
slash-and-burn cultivation.

Jose de Sousa e Silva Jr., Departamento de Zoologia,
Museu Paraense Emilio Goeldi, Caixa Postal 399,
66040-170 Bel6m, Pard, Brazil, and (current address)
Laborat6rio de Vertebrados, Departamentos de Ecologia
e de Gen6tica, CCS, Universidade Federal do Rio de
Janeiro, Caixa Postal 68020, 21941-970 Rio de Janeiro,
Rio de Janeiro, and Mauricio de Almeida Noronha,
Fundag o Floresta Amaz6nica, Rua dos Jatobis 142,
Coroado III, 69085-380 Manaus, Amazonas, Brazil.


Hershkovitz, P. 1977. Living New World Monkeys
(Platyrrhini), With an Introduction to Primates, Vol.
1. Chicago University Press, Chicago.
Mittermeier, R. A., Schwarz, M. and Ayres, J. M. 1992.
A new species of marmoset, genus Callithrix Erxleben,
1777 (Callitrichidae, Primates) from the Rio Mauds
region, state of Amazonas, Central Amazonia, Brazil.
Goeldiana Zoologia (14): 1-17.
Silva JOnior, J. S. and Noronha, M. de A. 1996. On a
new species of bare-eared marmoset, genus Callithrix
Erxleben 1977, from Central Amazonia, Brazil.
Goeldiana Zoologia (18). In press.


El present studio es parte del monitoreo de un grupo
de monos aulladores que fue liberado em la isla
Agaltepec, Veracruz, Mexico, dentro de un program
de translocaci6n con fines conservacionistas (Rodriguez-
Luna et al., 1993). La liberaci6n del grupo progenitor
termin6 em 1989 y la tropa qued6 consituida por 9
animals adults (8 hembras y 1 macho) y un infante; a
partir de entonces, se ha presentado un crecimiento
considerable a lo largo de estos aflos (Cortds-Ortiz et
al., 1994).

Un aspect important a estudar en un grupo de monos
recidn formado y en crecimiento, es la forma en que se

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Neotropical Primates 4(2), June 1996

Neotropical Primates 4(2), June 1996 Page 60

organizan socialmente para reproducirse. Para la especie
se han reportado diferentes tamafios de grupo, variando
desde 8 hasta 20 individuos (Milton, 1982; Estrada,
1982; Crockett y Eisenberg, 1986). La organizaci6n
social ha sido descrita como grupos o subunidades
poliginicos que pueden ser unimacho, multimachos y
de edad graduada (Jones, 1985; Crockett y Eisenberg,
1986) y algunos investigadores han reportado la forma
en que hembras y machos se asocian para copular
(Carpenter, 1965; Glander, 1980; Jones, 1985; Clarke,
1983). Sin embargo, mientras algunos afirman que el
macho dominant en turno tiene acceso exclusive a las
hembras adults (Clarke, 1984) y que aun cuando haya
mas de un macho adulto en las tropas, el sistema de
organizaci6n social es muy similar al sistema de harem
unimacho (Crockett y Eisenberg, 1986); outros aseguran
que las hembras en estro pueden establecer relaciones
de consorte y copular con various machos durante este
period (Carpenter, 1965), y han observado que los
machos dominantes copulan en la parte media del estro,
que ha sido relacionado con el "pico" de la fertilidad, y
los de menor rango durante otros moments antes o
despu6s de este pico (Glander, 1980; Jones, 1985).

Debido a estas diferencias, se planted la necesidad de
estudiar el tipo de organizaci6n social que opera en el
grupo de monos aulladores de Agaltepec. Las
observaciones pra este studio se realizaron durante 3
periods (marzo-abril 1994, noviembre 1994, mayo-julio
1995), cumpli6ndose con un total de 1225 horas de
observaciones y pudi6ndose observer 25 estros de 12
hembras diferentes. Al moment de iniciar las
observaciones la tropa estaba constituida por 30
individuos: 6 machos adults, 12 hembras adults (3 de
las cuales todavia no habian tenido crias, ni se habian
observado en interacciones secuales), 9 juveniles y 2
infants (3 infants mAs nacieron durante este periodo;
conforme transcurrieron los periodos de observaciones,
various machos y hembras entraron en la madurez sexual
e intervinieron en algunos episodios reproductivos de
la tropa; asimismo, una hembra adulta muri6. Al finalizar
las observaciones el grupo estaba constituido por 36
individuos: 9 machos adults, 14 hembras adults (9 que
habian tenido por lo menos una cria, 2 gestantes y tres
nuliparas), 11 juveniles y 2 infants.

Con base en la informaci6n obtenida a partir de la
revision bibliogrAfica, se formularon 4 hip6tesis
alternatives sobre el tipo de asociaci6n que se Ilevaria a
cabo entire machos y hembras en Agaltepec. Estas
hip6tesis son las siguientes:

1. Las hembras de la especie Alouatta palliata copulan
con various machos cualesquiera que estos sean, durante
sus periodos de estro (Carpenter, 1965).
2. Las hembras de la especie Alouatta palliata copulan

con various machos durante su period de estro; sin
embargo, .durante un period restringido, dondo se
supone hay mayor fertilidade, copulan preferen-
temente con un solo macho (Jones, 1985; Glander,
3. Las hembras de la especie Alouattapalliata copulan
con un solo macho durante sus periods de estro, pero
siempre con el mismo (Clarke, 1983).
4. Las hembras de la especie Alouatta palliata copulan
con un solo macho durante un period de estro, pero
pueden cambiar de pareja durante los estros siguientes.

Estas hip6tesis fueron enviadas, en noviembre de 1994,
a cuatro de los investigadores que han estudiado
poblaciones de monos aulladores en libertad y que han
hecho algunos comentarios sobre el sistema de
apareamiento en sus articulos (Dr. Margaret Clarke, Dr.
Clara Jones, Dr. Kenneth Glander y Dr. John Eisenberg),
para que las analizaran y asignaran, segin su experiencia,
la probalidad de cada una de las hip6tesis en tropas
silvestre de Alouatta palliata.

Los investigadores consultados difirieron en su
apreciaci6n sobre la probable ocurrencia de las hip6tesis
planteadas; lo cual sugiere que esta organizaci6n no es
estdtica, sino que varia de acuerdo a condiciones
particulares. Todos los investigadores coincidieron en
que esta organizaci6n depend de la estructura del grupo
y de su estabilidad social.

Los datos obtenidos durante el trabajo de campo
realizado para este studio, actualmente estAn siendo
analizados para identificar cuil o cuiles de estas hip6tesis
se cumplen para la tropa de Agaltepec, y si existen
preferencias individuals sobre alguno de los patrons
de apareamiento referidos o sobre la pareja. Sin embargo,
los datos para este grupo multimacho, sugieren que la
mayoria de las hembras copulan con mas de un macho
durante un period de estro.

En Mexico, la mayoria de las selvas donde se pueden
encontrar poblaciones de monos aulladores, se
encuentran severamente fragmentadas, y los grupos
quedan aislados en estos remanentes de vegetaci6n,
provocando la imposibilidad de emigraci6n de machos
y hembras hacia otros grupos o la inmigraci6n de nuevos
adults a las tropas, como sucede de manera natural en
habitat extenso. Esta situaci6n conlleva a la formaci6n
de un grupo cerrado, como lo es el grupo de Agaltepec,
con presencia de un mayor nOmero de machos de los
que regularmente se presentan en condiciones naturales
para la especie; por lo cual es muy probable que un tipo
de organizaci6n social muy.similar al de Agaltepec estd
operando en muchas tropas de monos aulladores en
habitat fragmentado.

Neotropical Primates'4(2), June 1996

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Page 61 Neotropical Primates 4(2), June 1996


Agradecemos a los doctors Margaret Clarke, Clara
Jones, Kenneth Glander y John Eisenberg, su disposici6n
para apoyar este trabajo y sus valiosos comentarios y
sugerencias sobre el mismo.

Liliana Cortes-Ortiz, Instituto de Neuroetologia,
Universidad Veracruzana, Apartado Postal 566, C.P.
91000, Xalapa, Veracruz, M6xicoy Manuel Martinez-
Morales, Maestria en Inteligencia Artificial, Universidad


Carpenter, C. T. 1965. The howlers of Barro Colorado
Island. En: Primate behavior: Field Studies of Mon-
keys and Apes, I. DeVore, I. (ed.), pp. 250-291. Holt,
Reinhart and Winston, Inc, New York.
Clarke, M. R. 1983. Infant killing and infant disappear-
ance following male takeovers in a group of free-rang-
ing howling monkeys (Alouattapalliata) in Costa Rica.
Am. J. Primatol. 5:241-247.
Clarke, M. R. y Glander, K. E. 1984. Female reproduc-
tive success in a group of free-ranging howling mon-
keys (Alouatta palliata) in Costa Rica. En: Female
Primates: Studies by Women Primatologists, M. F.
Small (ed.), pp. 111-126. Alan R. Liss, New York.
Cort6s-Ortiz, L., Rodriguez-Luna, E., Martinez-Morales,
M. y Carrera-Sanchez, E. 1994. Parametros
demograficos y reproductivos de un grupo de monos
aultadores (Alouatta palliata) en semilibertad. La
Ciencia y el Hombre (18): 141-166.
Crockett, C. M. y Eisenberg, J. F. 1986. Howlers: Varia-
tion in group size and demography. En: Primate Soci-
eties, B. B. Smuts, D. L. Cheney, R. M. Seyfarth, R.
W. Wrangham y T. T. Struhsaker, (eds), pp. 54-68.
University of Chicago Press. Chicago.
Estrada, A. 1982. Survey and census of howler mon-
keys (Alouatta palliata) in the rain forest of "Los
Tuxtlas", Veracruz, Mexico. Am. J. Primatol. 2:363-
Glander, K. E. 1980. Reproduction and population
growth in free-ranging mantled howling monkeys. Am.
J. Phys. Anthropol. 53:25-36.
Jones, C. B. 1985. Reproductive patterns in mantled
howler monkeys: estrus, mate choice and copulation.
Primates 26(2):130-142.
Milton, K. 1982. Dietary quality and population regula-
tion in a howler monkey population. En: The Ecology
of a Tropical Forest: Seasonal Rhythms and Long-
term Changes, E. G. Leigh Jr., A. S. Rand y D. M.
Windsor (eds.), pp. 273-289. Smithsonian Institution
Press, Washington, D. C.
Rodriguez-Luna, E., Garcia-Ordufia, F. y Canales-
Espinosa, D. 1993. Translocaci6n del mono aullador

Alouattapalliata: una alternative conservacionista. En:
Estudios Primatol6gicos en Mixico, Vol. 1, A. Estrada,
E. Rodriguez-Luna, R. L6pez-Wilchis y R. Coates-
Estrada (eds.), pp. 129-177. Biblioteca Universidad
Veracruzana. Xalapa, Veracruz.


The occurrence and distribution of the brown howler
monkey (Alouattafusca clamitans) is being studied in
the municipal district of Porto Alegre (300S 51W), the
capital of Rio Grande do Sul, the southernmost state of
Brazil. The presence of the brown howler monkey was
mentioned in the "List of the Vertebrates of the Great
Porto Alegre" (Fundaqgo Zoobotinica do Estado do Rio
Grande do Sul, 1976). However, very little is known
about where in Porto Alegre it occurs and how well (or
poorly) preserved its populations are. Only by gathering
more information on the status of the species will it be
possible to define strategies for its conservation in the

Porto Alegre is near the southern boundary of the
distribution of Alouatta fusca clamitans (Prates et al.,
1990). The municipality covers 47,152,28 ha and has a
population of about one million 200 thousand people,
mostly concentrated in the northern part. As the city
develops, and formerly forested areas are occupied, the
distribution of A. fusca is becoming more and more
restricted. The study aims to provide data on the status
of the populations of the brown howler monkey in the
face of the current urban growth and to establish a basis,
through the analysis of habitat quality, for the
management and preservation of the remaining

The area was divided into a 25 ha grid and the squares
with remaining forests are being visited and thoroughly
searched for the presence of the howlers. Occurrences
are being registered on the basis of actual sightings and
also by the presence of feces. Vocalizations and reports
by local inhabitants are considered as additional
information. Subsequently, vegetation analyses will be
carried out in some of the 25 ha squares visited to assess
any relationship between habitat structure and the
presence or otherwise of the brown howler monkey. The
study is divided into three phases, each covering a third
of the area of Porto Alegre, and will last twelve months.
The work is being developed by a group of researchers
from the Department of Zoology of the Federal
University of Rio Grande do Sul (Luis Fernando
Guimaraes Brutto, Sidnei Dornelles, Rodrigo Cambard
Printes, Gerson Buss and MArcia M. de Assis Jardim,
under the coordination of Helena Piccoli Romanowski

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Neotropical Primates 4(2), June 1996

Neotropical Primates 4(2), June 1996 Page 62

and Luis Flamarion B. de Oliveira; and is receiving
support from the Fundaqo de Amparo A Pesquisa do
Estado do Rio Grande do Sul (FAPERGS), the Pr6-
Reitoria de Pesquisa e P6s-Graduaq9o da Universidade
Federal do Rio Grande do Sul" (PROPESP-UFRGS) and
the Secretaria Municipal do Meio Ambiente. Porto
Alegre (SMAM-PMPA).

Gerson Buss, Departamento de Zoologia, Universidade
Federal do Rio Grande do Sul, Avenida Paulo Gama s/
n., Prddio 12125, Sala 326, 90049-060 Porto Alegre,
Rio Grande do Sul, Brazil.


Fundaqao Zoobotanica do Estado do Rio Grande do Sul.
1976. Preceituagdo Ecol6gicapara a Preservaago das
Areas Naturais da Grande Porto Alegre. Editora
Sulina, Porto Alegre. 151 pp.
Prates, J. C., Kunz Jr., L. F. and Buss, G. 1990.
Comportamento postural e locomotor de Alouatta
fusca clamitans (Cabrera, 1940) em floresta subtropi-
cal (Primates, Cebidae). Acta Biologica Leopoldensia
12(1): 189-200.


The Carlos Botelho State Park of 37,432 ha (240 44'-
240 15'S, 470 46'480 10'W) in the south of the state of
Sao Paulo takes in part of the municipalities of Sao
Miguel Arcanjo, Capdo Bonito and Sete Barras. It
protects an important part of the Atlantic forest of the
Serra do Mar, more precisely the Serra de Paranapiacaba.
The primates occurring in the park include Cebus apella
nigritus, Alouatta fusca clamitans, and the muriqui,
Brachyteles arachnoides. The first study of the muriqui
population there was carried out in 1985-1986 by
Paccagnella (1991; see also Mittermeier et al., 1987),
who estimated a population of 500-800 animals,
concentrated in the forest above 600 m altitude. The
importance of this site for muriquis resulted in the
establishment of a research program there in August
1988 (Strier, 1992), initially involving the location and
habituation of study groups, a difficult task in the steep,
mountainous terrain, but resulting eventually in
successful studies of the diet and distribution of food
resources, ranging behavior, and activity patterns
(Moraes, 1992a, 1992b, 1994; Carvalho Jr., 1994). A
more recent study was begun in January 1994, and since
then 26 months of data have been obtained on the diet
and feeding behavior, ranging and daily activity patterns
of one of the groups. Samples of food sources were
collected during this period in order to carry out
nutritional analyses, comparing especially seasonal

differences (Gomes, 1994). Data was also collected on
their feeding postures and hand preferences when
manipulating the food. The study is jointly supervised
by Dr. Karen B. Strier, Department of Anthropology,
University of Wisconsin Madison, and Dr. C6sar Ades,
Institute of Psychology, University of Sao Paulo, Sio
Paulo. Financial support has been kindly provided by
the U. S. National Science Foundation (Grant
BNS9582998), the Liz Clayborne and Art Ortenberg
Foundation, and the Chicago Zoological Society (all to
Dr. Karen Strier) and the Brazilian Science Council
(CNPq) and FINEP, Rio de Janeiro (to Dr. C6sar Ades
and M. Talebi Gomes). The study forms parts of the
requirements for a Master's thesis for the Institute of
Psychology, University of Sao Paulo.

Mauricio Talebi Gomes, Departamento de Psicologia
Experimental, Universidade de Sao Paulo, Avenida
Professor Mello Moraes 1721, Caixa Postal 66261,
05508-900 Sao Paulo, Sao Paulo, Brazil.


Carvalho Jr., 0. 1994. Ecologia e comportamento do
mono-carvoeiro, Brachyteles arachnoides, no Parque
Estadual Carlos Botelho (PECB) SP. In: Resumos.
VI Congress Brasileiro de Primatologia, p. 66.
Universidade Federal do Rio de Janeiro, Rio de Janeiro,
24-29 July, 1994. (Abstract).
Gomes, M. T. 1994. Aspecto de contefido nutricional
da dieta e comportamento alimentar de Brachyteles
arachnoides no Parque Estadual de Carlos Botelho -
SP. In: Resumos. VI Congresso Brasileiro de
Primatologia, p. 62. Universidade Federal do Rio de
Janeiro, Rio de Janeiro, 24-29 July, 1994. (Abstract).
Mittermeier, R. A., Valle, C. M. C., Alves M. C., Santos,
I. B., Pinto, C. A. M., Strier, K. B., Young, A. L.,
Veado, E. M., Constable, I. D., Paccagnella, S. G. and
Lemos de Sa, R. M. 1987. Current distribution of the
muriqui in the Atlantic forest region of eastern Brasil.
Primate Conservation (8): 143-148.
Moraes, P. L. R. 1992a. Dispersao de sementes pelo
mono-carvoeiro (Brachyteles arachnoides E.
Geoffroy, 1806) no Parque Estadual de Carlos Botelho.
Rev. Inst. Florest., Sdo Paulo 4: 1193-1198.
Moraes, P. L. R. 1992b. Esp6cies utilizadas na
alimentagao no mono-carvoeiro (Brachyteles
arachnoides E. Geoffroy, 1806) no Parque Estadual
de Carlos Botelho. Rev. Inst. Florest., Sao Paulo, 4:
Moraes, P. L. R. 1994. Disponibilidade alimentar e
padres de distribuig o espacial de espdcies utilizadas
pelo muriqui no Parque Estadual de Carlos Botelho,
Sao Paulo. In: Resumos. VI Congresso Brasileiro de
Primatologia, p. 61. Universidade Federal do Rio de
Janeiro, Rio de Janeiro, 24-29 July, 1994. (Abstract).

Neotropical Primates 4(2), June 1996

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Page 63 Neotropical Primates 4(2), June 1996

Paccagnella, S. G. 1991. Censo da populagao de monos
(Brachyteles arachnoides) do Parque Estadual Carlos
Botelho, estado de Sao Paulo. In: A Primatologia no
Brasil 3, A. B. Rylands and A. T. Bemardes (eds.),
pp.225-233. Fundag9o Biodiversitas and Sociedade
Brasileira de Primatologia, Belo Horizonte.
Strier, K. B. 1992. Faces in the Forest: The Endangered
Muriqui Monkeys of Brazil. Oxford University Press,


The four International Management Committees for the
Brazilian lion tamarins held their annual meeting on 28-
29 May 1996, in Brasilia, hosted by Maria lolita Bampi,
Head of the Faunal Division of the Wildlife Department
of the Brazilian Instiute for the Environment (Ibama).
Over the two days the committees discussed the status
of the captive and wild populations. The captive breeding
program for the golden lion tamarin, Leontopithecus
rosalia, supervised by Jon Ballou and Devra Kleiman
of the National Zoological Park, Washington, D. C., has
now reduced the captive population to a little under 500
animals, while still maintaining a good part of the genetic
diversity of founders. Golden-headed lion tamarins, L.
chrysomelas, now number over 600 in captivity, and
although the populations in Europe and the USA are now
stable, those in Brazil and Asia continue to grow. There
was considerable discussion regarding the future of the
black lion tamarin, L. chrysopygus, in captivity (and in
the wild). The captive population now numbers over 80
animals, but the very few founders available would
require the maintenance of an unacceptably large
population in captivity to preserve genetic diversity.
Having decided on the merits of maintaining a captive
population, a metapopulation management plan drawn
up by Claudio Valladares-Padua (University of Brasilia)
and Jon Ballou, which involves the regular introduction
of new founders from the five wild populations, was
presented, discussed and approved. The various teams
working on research and conservation measures in the
wild presented reports on their progress. Notable is the
highly successful translocation program being carried
out by Cecilia Kierulff (University of Cambridge) and
Paula Procopio (Golden Lion Tamarin Association) for
the isolated and threatened groups of L. rosalia. James
Dietz (University of Maryland) reported on his ongoing
ecological studies ofL. chrysomelas at the Una Biological
Reserve. Claudio Valladares-Padua informed on the
continued monitoring of the wild groups of L.
chrysopygus, and also the progress concerning an
ecological behavioral study which his team has begun
on a group of L. caissara on the island of Superagiii,
Parand. Guadalupe Vivekananda, Director of the

Superagili National Park, and S6rgio Brant (Ibama)
reported on the progress made over the last year
regarding efforts to improve the status and infrastructure
for the Park, as well as to increase its size to include
further important populations of L. caissara. Faigal
Simon (Sao Paulo Zoo) resigned as co-chairman of the
committee for the black lion tamarin. He was warmly
thanked for his work on behalf of the committee,
especially for his role in establishing the current captive
population of the black lion tamarin. Alcides Pissinatti
(Rio de Janeiro Primate Center CPRJ/FEEMA) was
appointed as co-chair in his place.

The 1997 meeting will be held in Belo Horizonte, Minas
Gerais. It will be organized by the Funda9ao
Biodiversitas and Conservation International do Brasil,
and will involve besides, a two-day symposium for state-
of-the-art reviews on research and conservation, as well
as two days devoted to Population and Habitat Viability
Analyses for the four lion tamarin species.


One of the recommendations made at the Saimiri
Workshop held in 1994 was to have a second meeting
in the area where they live. From 5-7 June 1995, 48
people met at the Manuel Antonio National Park, on the
Central Pacific coast of Costa Rica, to discuss the status
and recommendations for the conservation of the species.
Five working groups were formed: Biology discussed
the biological data available and ran the VORTEX
population simulation for the species; Distribution -
analyzed the past and current distribution of the species
and mapped the areas of remaining Saimiri populations;
Translocation and Captive Breeding developed
protocols in case these measures were necessary for the
survival of the species, helped by Dr. Cheryl Asa, St.
Louis Zoo, and Dr. Larry Williams, University of South
Alabama; Public Education discussed the need for an
education program; and Community analyzed current
and future community actions, with the collaboration of
local authorities and five campesinos.

The final report is still being prepared. It includes
recommendations for more specific research to support
an adequate management of the species. The most urgent
topics were given as: 1) Determination of the distribution
and demographics of the populations of the two
subspecies, S. o. oerstedi and S. o. citrinellus; 2) the
establishment of a public education program directed to
the local communities and tourists; and 3) the
establishment of a community action plan to protect the
species. Dr. Ulysses Seal, Chairman of the IUCN/SSC

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Neotropical Primates 4(2), June 1996

Neotropical Primates 4(2), June 1996

Conservation Breeding Specialist Group (CBSG), met
with government authorities, who indicated their
willingness to implement the recommendations of the
workshop. From a report to CBSG News 1995, 6(1): 23,
by Yolanda Matamoros, ALPZA-AMAZOO.


Bristol Zoo has been exhibiting primates since it opened
in 1836. At the present time, the cebid collection
comprises 2.2 black howlers (Alouatta caraya), 2.2 long-
haired spider monkeys (Ateles belzebuth belzebuth), 3.9
squirrel monkeys (Saimirisciureus) and 3.1 white-faced
sakis (Pithecia pithecia). Other primate groups are
represented in the Zoo and include two species of lemur,
three species of callitrichids and five species of Old
World monkeys.

As well as exhibiting the above cebids, the zoo
demonstrates its commitment to their captive
management in other ways. The UK Taxonomic
Advisory Group (TAG) subgroup for cebids is chaired
by Sian Waters. This group is subordinate to the
European subgroup which is chaired by Bert de Boer of
Apenheul Zoo in the Netherlands. Sian Waters took over
the leadership of the cebid group in 1994, but prior to
that Neil Bemment of Paignton Zoo, UK, ran it and much
work was accomplished by him. For example, Neil
Bemment began a programme to karyotype the British
population of spider monkeys, which is now well
underway. This work will now be extended to the rest
of Europe. Plans for the future include investigating the
possibility of initiating breeding programs for more
endangered cebid species, bearing in mind the
recommendations of the forthcoming Global Captive
Action Plan for primates.

The zoo other contributions involve two breeding
programmes for cebid species. One is a European
Breeding Programme (EEP) for the white-faced saki,
with Sian Waters as the Species Co-ordinator and the
other is a European studbook for black howlers compiled
by Darren Webster, Bristol Zoo.

Two editions of the European black howler studbook
have been published (Webster, 1995, 1996). Black
howlers have a small founder base of 16 (7.9) in Europe,
and only Twycross Zoo, UK, has had success with
sustained breeding. We plan to carry out a husbandry
survey for black howlers in Europe and an exchange of
potential founder males has already taken place between
Bristol and Apenheul. Between them both zoos hold 6
(2.4) potential founders.

The first European studbook for white-faced sakis will

be published in 1996. Many zoos are interested in
exhibiting the species and there is even an EEP
participant in South Africa. Although the white-faced
saki is not an endangered species an EEP was deemed
necessary because the species is the only representative
of its genus in any numbers in captivity in Europe. A
species committee has been elected and comprises
representatives from all over Europe. Husbandry
guidelines will be formulated in the future, although
some information on management is available in Waters

Although the long-haired spider monkeys are the only
species exhibited ate Bristol Zoo which are classed as
vulnerable in the wild, we feel it important to exhibit
cebids which are not endangered to improve techniques
and as a way to inform the public about primate biology
and the threats faced by more endangered species in
South America.

Sian S. Waters and Darren A. Webster, Bristol Zoo
Gardens, Clifton, Bristol BS8 3HA, England, UK.


Waters, S. S. 1995. A review of social parameters which
influence breeding in white-faced saki Pithecia
pithecia in captivity. Int. Zoo. Yb. 34:147-153.
Webster, D. A. 1995. European Studbook for Black
Howlers (Alouatta caraya). No. 1. Bristol Zoo Gar-
dens, UK.
Webster, D. A. 1996. European Studbook for Black
Howlers (Alouatta caraya). No. 2. Bristol Zoo Gar-
dens, UK.


On the 12th July 1996, the Governor of the Brazilian
state of Amazonas, Amazonino Mendes, signed the law
approved by the State Assembly to transform the
1,124,000 ha Mamiraua Ecological Station, located 600
km upriver from Manaus on the Rio Solimies, into
Brazil's first Sustainable Development Reserve. This
action, following four years of work by the Sociedade
Civil Mamiraua and the local communities in and around
the reserve, legitimizes a unique participatory approach
to biodiversity conservation, research and management,
and provides the legal framework for the creation of
similar reserves throughout the Amazon.

The transformation of Mamiraud into a sustainable
development reserve removes the conflicts caused by

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the previous legislation, which implied the removal of
local residents. Quite to the contrary, the new law will
promote their active participation in the management of
the reserve, protect their access to the natural resources,
and make them principal partners in this conservation

While fishing and timber production within the reserve
is of crucial importance to local residents and regional
markets, the maze of lakes, channels and forests in this
inland delta in the upper Amazon is also home to a wide
range of endangered or endemic species of wildlife.
These include a large number of aquatic birds, the
Brazilian manatee (Trichechus inunguis), the giant
Amazon river otter (Pteroneura brasiliensis), the black
caiman (Melanosuchus niger), tapirs (Tapirus
brasiliensis), and jaguars (Panthera onca), and the
Reserve covers the entire known geographic ranges of
the blackish squirrel monkey (Saimiri vanzolinii), and
the white-faced uakari (Cacajao calvus calvus). The
conservation value of MamirauA's biodiversity and its
importance for studying the intricate aspects of vdrzea
ecology have resulted in the area being included in the
Ramsar list of Internationally Important Wetlands, and
also its proposal as a future Biosphere Reserve under
UNESCO's Man and the Biosphere Program.

The Mamiraud Project has concentrated its efforts so
far in the eastern fifth of the reserve, and has supported
local initiatives to protect fishery resources, vital to the
lifestyle of the local people (caboclos) in the flooded
vdrzea forests. In addition to helping residents organize
the enforcement of regulations regarding the judicious
use of natural resources, the project has developed a wide
range of socioeconomic and ecological studies aimed at
understanding the workings of this unique ecosystem
and developing guidelines for biodiversity conservation
and sustainable resource use aimed at improving the
living standards and reducing the impact of subsistence
and small scale commercial activities in agriculture,
forestry and fishing.

Results of the first four years of the project include a
wide variety of research projects carried out by more
than 80 Brazilian and foreign researchers in
anthropology, epidemiology, fisheries management and
ecology, terrestrial ecology, agroforestry, soils, and
limnology, among other fields. Extension efforts have
supported the development of environmental education
in local schools, participation of local communities in
policy formulation and resource protection, and
increasing interest in agroforestry and traditional
agricultural techniques.

The Project has been coordinated since it's inception by
Dr. Jos6 MArcio Ayres of the Sociedade Civil Mamiraua

and the Wildlife Conservation Society (WCS).
Institutional agreements have been signed with both state
and national authorities, including the Brazilian Institute
of the Environment and Renewable Natural Resources
(IBAMA), the Brazil Science Council (CNPq), and the
Institute for Protection of the Amazon Environment
(IPAAM). Financial support has come from a wide range
of organizations, including the British Overseas
Development Administration (ODA), the World Wide
Fund for Nature (WWF), the Wildlife Conservation
Society (WCS), the European Union (EU), and the Brazil
Science Council (CNPq).

Donald Masterson, Projeto Mamiraud, Sociedade Civil
Mamiraud, Caixa Postal 38, 69470-000 Tefd, Amazonas,


Ayres, J. M. 1993. As Matas de Vdrzea do Mamiraud.
Conselho Nacional de Desenvolvimento Cientifico e
Tecnol6gico (CNPq), Brasilia. 123pp.
Queiroz, H. L. de. 1995. Preguigas e Guaribas: Os
Mamiferos Folivoros Arboricolas do Mamiraud.
Conselho Nacional de Desenvolvimento Cientifico e
Tecnol6gico (CNPq), Brasilia. 161pp.


We are very pleased to announce the creation of the
Margot Marsh Biodiversity Foundation, a new charitable
foundation dedicated exclusively to primate
conservation. This foundation was created by the late
Margot Marsh of La Jolla, California, a long-time
supporter of a wide variety of primate research and
conservation efforts, who died in May 1995.

I had the great privilege of knowing Margot Marsh for
13 years, and was able to enjoy her company on various
trips, including one to Madagascar to see lemurs and
another to Kenya and Rwanda to see mountain gorillas
and some of savanna-dwelling species of Kenya's Masai
Mara Reserve. Margot was extremely knowledgeable
about primates and human evolution, not to mention
many other aspects of biodiversity, and was a great friend
and supporter of many of our organizations. We should
all be honored that she saw fit not only to remember us
in her will, but also to ensure that the kinds of projects
that she supported during her life would continue
receiving support in the future.

The Primate Specialist Group was specifically mentioned
in Margot's will, as were some of our newsletters,
journals and action plans, so she clearly recognized the

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Neotropical Primates 4(2), June 1996

Neotropical Primates 4(2), June 1996 Page 66

value of our group and the critical role that it plays in
global primate conservation activities. In recognition of
this, some of the first projects supported by the
Foundation have been aimed at ensuring the continuity
of publications such as Neotropical Primates. We are
extremely grateful to this wonderful friend, and will
miss her all very much.

The mission of the Margot Marsh Biodiversity
Foundation is straightforward: "To contribute to global
biodiversity conservation by providing strategically
targeted, catalytic support for the conservation of
endangered nonhuman primates and their natural

Project guidelines are as follows, with preference being
given to projects that have one or more of the following
characteristics: 1) Projects focusing on endangered
nonhuman primates living in their natural habitats; 2)
primate projects being conducted in areas of high overall
biodiversity and under great threat (e.g., "threatened
hotspots", "megadiversity countries") to ensure
maximum multiplier effect for each project; 3) projects
being carried out by nationals from the tropical countries
to increase local capacity for implementing biodiversity
conservation; 4) projects that strengthen international
networks of field-based primate specialists and enhance
their capacity to be successful conservationists; and 5)
projects that result in publication of information on
endangered primate species in a format that is useful
both to experts and the general public.

Projects should contribute to at least one, and preferably
more, of the following themes: 1) Enhancement of
scientific understanding/knowledge of the target species/
ecosystem; 2) improved protection of a key species,
habitat, or reserved area; 3) demonstration of economic
benefit achieved through conservation of a species and
its habitat, as compared to loss thereof; 4) increased
public awareness or educational impact resulting from
the project in question; and 5) improved local capacity
to carry out future conservation efforts through training
or practical experience obtained through project

The Board of Directors of the Margot Marsh Foundation
consists of three members, and an Advisory Group has
also been created with an additional three members, all
of them selected on the basis of their past relationship
with, and knowledge of the interests of, Margot Marsh.
I currently serve as President of the Board of Directors,
and inquiries about how to apply for support from the
foundation can be sent to me at the address below.

Russell A. Mittermeier, Margot Marsh Biodiversity

Foundation, 432 Walker Road, Great Falls, Virginia
22066, USA. Fax: + 1 703 759 6879.


An international studbook for the pied tamarin, Saguinus
bicolor bicolor, has been endorsed by the World Zoo
Organization (IUDZG) and the Species Survival
Commission (SSC) of the World Conservation Union
(IUCN). The studbook keeper is Dr. Andrew J. Baker,
Curator of Primates and Small Animals, Philadelphia
Zoological Garden, 3400 West Girard Avenue,
Philadelphia, PA 19104-1196, USA. Dr. William
Langbauer Jr., also of the Philadelphia Zoological
Garden, has taken over as studbook keeper of the cotton-
top tamarin, Saguinus oedipus. Reported in CBSG News,
December 1995, 6(1):15.

A Conservation International do Brasil, Fundagao
Biodiversitas, Fundacao SOS Mata Atlintica e Fundagao
Andre Tosello/Base de Dados Tropicais, promoveram
nos dias 23 e 24 de maio, o Workshop "Padrbes de
Distribuigio da Biodiversidade da Mata Atlantica do Sul
e Sudeste Brasileiro", em Campinas, Sgo Paulo. 0
encontro 6 parte de uma sdrie de Workshops para a
definigo de areas prioritArias para a conservagdo dos
biomas brasileiros.

A reuniao de Campinas teve dois objetivos principals:
apresentar e discutir com os especialistas em Mata
Atlantica das regimes sul e sudeste a metodologia e as
ferramentas utilizadas para realizar diagn6sticos de Areas
prioritArias para conservagao sob a 6tica regional; e
identificar as principals subunidades biogeogrAficas e
seus problems de conservagAo. Para tal, reuniram-se
cerca de 40 especialistas provenientes de instituigoes
academicas, organizag6es nao-governamentais
ambientalistas, 6rgaos estaduais e federais de meio
ambiente, e pesquisadores com larga experiencia sobre
a Mata Atlantica do sul e sudeste brasileiro.

Os especialistas trabalharam em grupos tematicos -
r6pteis e anfibios, mamiferos, aves, peixes e
invertebrados, flora e estrat6gias de conservaao para
fazer um pr6-diagn6stico das informag9es existentes e
discriminar os dados mais importantes a serem obtidos
para definir as Areas prioritarias para conservagAo,
segundo o grupo biol6gico em discussAo.

Neotropical Primates 4(2), June 1996

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Page 67 Neotropical Primates 4(2), June 1996

Os resultados da reuniao serao divulgados em breve
via Internet, que apresentard um mapa das principals
regi6es-alvo para diagn6sticos de biodiversidade da
Mata AtlIntica do sul e sudeste, uma avaliagao das
lacunas de conhecimento para as diversas parties da
Mata Atlintica, e definir um program de trabalho para
agregar as informaq6es visando uma base dados que
fornecera suporte para uma Politica Nacional de
conservagao para o bioma.

Luiz Paulo de Souza Pinto, Conservation
International do Brasil, Avenida Ant6nio, Abrahao
Caram 820/302, 31275-000 Belo Horizonte, Minas
Gerais, Brasil.


The European editorial office of Animal Behaviour,
the journal of the Association for the Study of Animal
Behaviour (ASAB), has moved. With the Executive
Editorship of the journal passing from Tim Roper to
Chris Barnard as from 1 May 1996, the office is now
at the University of Nottingham, UK. All
communications relating to manuscripts and editorial
matters should be directed to the Managing Editor:
Dr. Angela Turner, Department of Life Science,
University of Nottingham, Nottingham NG7 2RD, UK,
Tel/Fax: +44 (0)115 9513249, e-mail:
a.k.turner@plnl .life.nott.ac.uk.

The latest table of contents and other information about
Animal Behaviour can be found at Academic Press'
IDEAL site, an online scientific journal library
containing content lists and abstracts of articles from
the 178 journals they publish. IDEAL has mirror sites
in North America and Europe. http://www.idealibrary.
com (San Jose, CA, North America), and http://www.
europe.idealibrary.com (Bath, UK).


SOCIOBIO is a moderated discussion list dealing with
the biological bases of the social behavior of animals,
including aggression, territoriality, social systems, and
mate selection. It welcomes contributions from
scholars currently engaged in any kind of research
project linked with the issues of socialization, altrusm,
and animal cooperation. To subscribe to SOCIOBIO
send an e-mail message to Listserv@sjuvm.
stjohns.edu containing the following command in the
body of the message < subscribe sociobio
yourfirstname yourlastname >. The list is run by Juan
C. Garelli, e-mail: garelli@ attach.edu.ar.

Primate Societies


Estamos preparando o Volume 5, no qual
serao incluidos os resumos dos trabalhos
apresentados no VI. Congress Brasileiro de
Primatologia que nao foram submetidos para publicagao.
Para a atualizacao e padronizagao destes resumos,
pedimos aos autores destes resumos o fornecimento das
seguintes informag6es: (a) os nomes completes de todos
os autores; e (b) o enderego e/ou vinculo atual do autor
para correspondencia (inclusive de e-mail, se tiver).

Stephen F. Ferrari, Departamanto de Genetica,
Universidade Federal do Para (UFPA), Caixa Postal
8607, 66.075-150 Bel6m, Pard, Brasil. Fax: 091-211-
1568, e-mail: ferrari@cuxiu.cbio.ufpa.br


The International Primatological Society (IPS) is holding
a workshop on Methods of Primate Conservation from
11-17 August 1996, immediately preceding the XVIth
International Primatological Congress being organized
jointly by IPS and the American Society of
Primatologists (ASP), in Madison, Wisconsin. The
workshop is for primatologists from developing
countries. Dr. Jeanne Altmann, Chairman of the
Conservation Committee of IPS and Research Curator
at the Brookfield Zoo, Chicago, will head the Workshop.
Arrival will be the evening of 9 August. Workshop
sessions will commence on 10 August. and will include
lectures, laboratory and observational work and
discussions. Special topics will include genetics,
nutrition, and behavior. The course will be given mainly
by Brookfield Zoo staff, but will also count on Dr. Anna
Feistner, Jersey Wildlife Preservation Trust, Jersey, UK,
and a number of other invited scientists. The Workshop
will end on 11 August, with bus transport to the IPS
Congress at Madison, Wisconsin. The Workshop will
aim to take on 24 participants from developing countries,
preferably eight from each southern continent.

Contact: Dr. Jeanne Altmann, Vice President for
Conservation, International Primatological Society,
Chicago Zoological Society, Brookfield Zoo, Chicago,
Illinois 60513, USA, Fax: 708 485 3532, e-mail:
bzconbio@ix.netcom.com, or Dr. Alison Jolly,

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Neolropical Primates 4(2), June 1996

Neotropical Primates 4(2), June 1996

President, International Primatological Society,
Department of Ecology and Evolutionary Biology,
Princeton University. Princeton, NJ 08544, USA, Fax:
609-258-5381, e-mail: ajolly@arachne.princeton.edu.


The Publications Committee of the American Society
of Primatologists (ASP) are accepting book proposal
applications for their newly established book series. Each
volume will be based on an organized symposium from
an ASP meeting and/or special topic in primatology.
Income from books published by ASP will be used for
fundamental purposes of the Society (e.g., Conservation
Fund, educational development, etc.). The inaugural
volume, entitled "Primate Conservation: The Role of
Zoological Parks", is based on a symposium presented
at the Congress in 1995, and will hopefully be ready for
sale at the ASP/IPS meeting in Madison in August 1996.
For details on submitting a book proposal, contact:
Janette Willis, Dept. OB/GYN, OUHSC, Box 26901,
Oklahoma City, OK 73190, USA, Tel: (405) 271 4229,
Fax: (405) 271-8547, e-mail: janette-
wallis@uokhsc.edu. From ASP Bulletin 20(2):5, June

Recent Publications


The most recent issue of the Boletin Primatol6gico
Latinoamericano, Volume 5, Number 1, 1995, 33pp.,
has been published by the Grupo Argentino de
Primatologia (GADEP). The editors are Gabriel E.
Zunino, Museo Argentino de Ciencias Naturales
"Bernardino Rivadavia", Buenos Aires, and Julio C.
Ruiz, Centro Argentino de Primatologia (CAPRIM),
Corrientes. This issue includes four articles and a note.
Alternative reproductive behaviors in the mantled howler
monkey (Alouatta palliata Gray): Testing Carpenter's
hypothesis Clara B. Jones; Labilidad cromos6mica: una
possible explicaci6n en el origen de los reordenamientos
cromos6micos en cebideos Marta D. Mudry et al.;
Deforestation, selective cutting, and habitat
fragmentation: the impact on a black howler monkey
(Alouatta caraya) population in northern Argentina -
Anneke DeLuycker; Dispersi6n y germinaci6n de
semillas de Ficus monckii por el mono aullador negro
(Alouatta caraya) Susan P. Bravo, Martin M.
Kowalewski and Gabriel E. Zunino; and (note) Monos
Cai (Cebus apella) en cautiverio: composici6n de grupos
- Aldo M Giudice and Marta D. Mudry.

Contributions to and correspondence regarding the
Boletin Primatol6gico Latinoamericano should sent to
Dr. Gabriel E. Zunino, Div. Mastozoologia, Museo
Argentino de Ciencias"Naturales, Avenida Angel
Gallardo 470, 1405 Buenos Aires, Argentina, e-mail:


Lista Anotada dos Mamiferos do Brasil/Annotated
List of Brazilian Mammals, by Gustavo A. B. da
Fonseca, Gisela Hermann, Yuri L. R. Leite, Russell A.
Mittermeier, Anthony B. Rylands and James L. Patton,
1996, 38pp. Occasional Papers in Conservation Biology
No. 4, Conservation International, Washington, D. C.,
and Fundacgo Biodiversitas, Belo Horizonte. ISBN 1
881173 17 8. In Portuguese and English. This booklet
presents an updated list of the Brazilian mammal fauna,
including terrestrial, aquatic and marine species.
According to existing data, the country harbors 483
continental and 41 marine mammals, totaling 524
species, distributed among 11 orders, 46 families and
213 genera. These estimates indicate that Brazil has the
most diverse mammalian fauna in the world, with
Indonesia a close second. This annotated list of Brazilian
mammals includes information on each species'
occurrence in Brazil's major biomes, along with body
weights, diet and locomotor adaptations, as well as
common names in Portuguese. Available from
Conservation International, Department of Conservation
Biology, 1015 Eighteenth Street, N. W., Suite 1000,
Washington D. C. 20036, USA. Fax: 202 887 0193.

A Belizean Rain Forest: The Community Baboon
Sanctuary, by Robert H. Horwich and Jonathan Lyon,
1990, 420pp. illustrated. Orang-utan Press. ISBN 0-
9637982-0-0. Price US$14.95 + $2 postage and handling
in the U.S., $4 outside the U.S. This is a composite of
information about the Baboon Sanctuary, its flora and
fauna, and Belizean natural history. Begun as an eight-
page booklet distributed to subsistence farmers, it has
developed into a 420 page description of a rain forest
and an alternative method of saving it. It has become a
rain forest primer for college and high school classes as
well as fortourists travelling to Belize or adjacent Central
American forests. To order: Community Conservation
Consultants, RD 1, Box 96, Gays Mills, WI 54631, USA,
Tel: (608) 735 4717, e-mail: ccc@mwt.net. Profits from
book sales go toward community conservation programs.

Primate Ontogeny: International Symposium, 10-
15 September 1995, Trest, Czech Republic. Meeting
Abstracts, by the Primatological Group of Czech
Republic of the Czech Anthropological Society, 1995,
22pp. Contact: Dr Marina Vancatova, Primate Research

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Neotropical Primates 4(2), June 1996

Group, VUFB Konarovice, 28125 Konarovice, Czech

Lemurs of the Lost World: Exploring the Forests
and Crocodile Caves of Madagascar, by Jane
Wilson, 1995, 2nd edition, 216pp. Impact books,
London. Paperback. ISBN 1 874687 43 9. Price 5.00
or US$11.00. This is a fascinating account of expeditions
to the Crocodile Caves at the AnkArana Massif, northern
Madagascar, in the late 1980's, the difficulties
confronted by the expedition members, the remarkable
paleaontological discoveries, and of the observations of
the fauna and especially the living lemurs in the sunken
forests there. Besides the crocodiles living in the complex
of more than 60 miles of caves, six subfossil lemurs have
been found there, including the remarkable sloth lemur,
Babakotia radofilai, the giant Megaladapis, Pachylemur,
Mesopropithecus, Palaeopropithecus, and
Archaeolemur. Ten living lemur species occur there
today, and a further two occurred there in the recent
past. This book is illustrated with beautiful color
photographs and provides an insight to the social,
economic, and conservation problems of Madagascar.
An excellent read, and highly recommended. In the UK,
write to Impact Books, 22 Glen Dale, Rowlands Castle,
Hants. P09 6EP, or Impact Books, 70 Newcomen Street,
London SE1 1YT, and in the USA it is available from
Impact Books, P. 0. Box 287, Great Falls, VA 22066,
USA. All profits from the sale ofthis book will be donated
to the Jersey Wildlife Preservation Trust for their work
in Madagascar.

Animais Ameagados de Extingdo, by Milton Thiago
de Mello, 1996, 67pp. Comitd Brasileiro da Associacao
Mundial de Veterindria, Rio de Janeiro. In Portuguese.
Chapters include: Extinct species; Demographic
explosion and extinction; Development and biodiversity;
Causes of threat; Conservation of threatened species;
Protected Areas; Lists of threatened species; Legislation;
Universities and research; Eco-92; Conclusions; and
Summary. For more information contact the author: Prof.
Milton Thiago de Mello, SHIN Q14, Conjunto 2, Casa
19, Lago Norte, 71510-225 Brasilia, D. F., Brazil.

BIODOC, Centro de Documentaci6n e
Informacidn. Catalogo de Publicaciones
Periodicas Existentes en el BIODOC hasta 1994,
edited by Nidia DurAn Villalobos and Enrique Quesada
Dobles, 1995, 105pp. 2nd edition. Program Regional
en Manejo de Vida Silvestre (PRMVS), Universidad
Nacional, Costa Rica. A catalogue of periodicals
maintained by BIODOC, sponsored by the U. S. Fish
and Wildlife Service and the Universidad Nacional,
Heredia, Costa Rica. Available from: BIODOC, Centro
de Documentaci6n en Vida Silvestre, Universidad
Nacional, Apartado 54-3000, Heredia, Costa Rica,

America Central. Tel: 2773-472, Fax: 237-7036.


Anderson, J. R., Degiorgio, C., Lamarque, C and Fagot,
J. A. 1996. A multi-task assessment of hand lateral-
ization in capuchin monkeys (Cebus apella). Primates
37(1): 97-103.
Ayres, J. M., Lima-Ayres, D. de M., Albernaz, A. L.,
Alves, A. R., Moura, E., Queiroz, H. L., Santos, P.,
Barthem, R. B. and Silveira, R. de. 1996. Mamiraud:
Um novo modelo de Estago Ecol6gica. Cidncia Hoje,
20(118): 24-33.
Baker, M. 1996. Fur rubbing: Use of medicinal plants
by capuchin monkeys (Cebus capucinus). Am. J.
Primatol. 38(3): 263-270.
Bard, K. A. 1995. Parenting in primates. In: Handbook
of Parenting, M. Bornstein (ed.), pp.27-58. Lawrence
Erlbaum Assoc., Mahwah, NJ.
Beck, B. B. 1995. Reintroduction, zoos, conservation,
and animal welfare. In: Ethics of the Ark: Zoos, Ani-
mal Welfare, and Wildlife Conservation. B. G. Norton,
M. Hutchins, E,. F. Stevens and T. L. Maple (eds.),
pp.155-163. Smithsonian Institution Press, Washing-
ton, D.C.
Bilgener, M. 1995. Chemical factors influencing food
choice of howler monkey (Alouattapalliata). Turkish
J. Zool., 19(4): 291-303.
Brice, S. 1995. Screening a New World monkey colony
for Yersinia and investigations of Y. pseudotubercu-
losis and soil. Dodo, J. Wildl. Preserve. Trusts 31:139-
Bravo, S. P., Kowalewski, M. M. and Zunino, G. E. 1995.
Dispersi6n y germinaci6n de semillas de Ficus monckii
por el mono aullador negro (Alouatta caraya). Bol.
Primatol. Lat. 5(1):25-27.
Butler, B. H. 1996. The enrichment challenge: A new
primate rain forest at Auckland Zoo. Shape of Enrich-
ment 5(1): 9-10.
Cartelle, C. and Hartwig, W. C. 1996. A new extinct
primate among the Pleistocene megafauna of Bahia,
Brazil. Proc. Nat. Acad. Sci. 93:6405-6409.
Casanova, C. and Vaz, C. 1995. [Do you know the
Callitrichidae?] Ciencia S6r VI(6): 28-34. (In Portu-
Chapman, C. A. 1995. Primate seed dispersal: Coevo-
lution and conservation implications. Evolutionary
Anthropology 4(3): 74-82.
Coates, K. W., Gibson, S., William. L. E., Brady, A.,
Abee, C. R., Shull, B. L. and Kuehl, T. J. 1995. The
squirrel monkey as an animal model of pelvic relax-
ation: An evaluation of a large breeding colony. Am.
J. Obstetr. Gynecol. 173(6): 1664-1670.
Coutinho, P. E. G. and Correa, H. K. M. 1995. Polygyny
in a free-ranging group of buffy-tufted ear marmo-
sets, Callithrix aurita. Folia Primatol. 65:25-29.

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Neotropical Primates 4(2), June 1996 Page 70

Defler, T. R. 1996. Aspects of the ranging pattern in a
group of wild woolly monkeys (Lagothrix lagotricha).
Am. J Primatol. 38: 289-302.
DeLuycker, A. 1995. Deforestation, selective cutting,
and habitat fragmentation: the impact on a black howler
monkey (Alouatta caraya) population in northern Ar-
gentina. Bol. Primatol. Lat. 5(1):17-24.
Fleagle, J. G. and Reed, K. E. 1996. Comparing primate
communities: a multivariate approach. J. Hum. Evol.
Forster, F.C. 1996. Novel objects and learning as en-
richment for captive common marmosets Callithrix
jacchusjacchus. Australian Primatology 10(3): 2-10.
French, J. A., Pissinatti, A. and Coimbra-Filho, A. F.
1996. Reproduction in captive lion tamarins
(Leontopithecus): Seasonality, infant survival, and sex
ratios. Am. J. Primatol. 39: 17-33.
Frisch, D. and Monska-Wright, I. 1994. Animal enrich-
ment at the Columbus Zoo. Proc. Nat. Conf AAZK
(American Assocation Zoo Keepers., Inc.) (1994): 55-
Froehlich, J. W. 1996. Primate Species: The irrevers-
ible units in the evolution of our mammalian diver-
gence. Am. J. Primatol. 38: 271-279. (Article + re-
view of Species, Species Concepts, and Primate Evo-
lution, W. H. Kimbel and L. B. Martin (eds.), Plenum
Publishing, New York, 1993, combined).
Giudice, A. M. and Mudry, M. D. 1995. Monos Cai
(Cebus apella) en cautiverio: composici6n de grupos.
Bol. Primatol. Lat. 5(1): 29-33.
Goldschmidt, B., Cabrera, M. A. A., Weinzetl, M. and
Llerena, J. C. 1995. Cytogenetic study in a specimen
of Cebus apella with an atypical phenotype. Folia
Primatol. 65:54-58.
Harcourt, A. H. 1995. Sexual selection and sperm com-
petition in primates: What are male genitalia good for?
Evol. Anthropol. 4(4): 121-129.
Hartwig, W. C. and Cartelle, C. 1996. A complete skel-
eton of the giant South American primate
Protopithecus. Nature, Lond. 381: 307-311.
Heistermann, M. 1995. Dominance status and behaviour
of captive-housed female cotton-top tamarins
(Saguinus oedipus) in motherless families. Primate
Report 42: 65-78.
Hook-Costigan, M. A. and Rogers, L. J. 1995. Hand,
mouth and eye preferences in the common marmoset
(Callithrix jacchus). Folia Primatol. 64(4): 180-191.
Jacobs, S. C., Larson, A. and Cheverud, J. M. 1995.
Phylogenetic relationships and orthogenetic evolution
of coat color among tamarins (genus Saguinus). Syst.
Biol. 44(4): 515-532.
Jones, C. B. 1995. Alternative reproductive behaviors
in the mantled howler monkey (Aloualta palliata
Gray): Testing Carpenter's hypothesis. Bol. Primatol.
Lat. 5(1): 1-5.
Jungers, W. L. and Burr, D. B. 1994. Body size, long

bone geometry and locomotion in quadrupedal mon-
keys. Zeit. Morph. Anthropol. 80(1): 89-47.
Jurke, M. H. 1996. Behavioral and hormonal aspects of
reproduction in captive Goeldi's monkeys (Callimico
goeldii) in a comparative and evolutionary context.
Primates 37(1): 109-119.
Jurke, M. H., Pryce, C. R. and Doebeli, M. 1995. An
investigation into sexual motivation and behavior in
female Goeldi's monkey (Callimico goeldii): Effect
of ovarian stat, mate familiarity, and mate choice.
Horm. Behav. 29(4): 531-553.
Kerr, J. and Currie, D. 1995. Effects of human activity
on global extinction risk. Conservation Biology,
9(6): 1528-1538.
Koontz, F, Koontz, C., Westrom, W., Glander, K.,
Horwich, R., Saqui, E. and Saqui, H. 1994. Reintro-
duction of black howler monkeys (Alouattapigra) into
the Cockscomb Basin Wildlife Sanctuary, Belize. Proc.
Nat. Conf AAZK (American Assocation Zoo Keep-
ers., Inc.) (1994): 104-111.
Laska, M. 1996. Taste preference thresholds for food-
associated sugars in the squirrel monkey (Saimiri
sciureus). Primates 37(1): 91-95.
Ludes, E. and Anderson, J. R. 1995. 'Peat-bathing' by
captive white-faced capuchin monkeys (Cebus
capucinus). Folia Primatol. 65:38-42.
MacLatchy, L. M. and Ross, C. F. 1995. Primate pale-
ontology. Evol. Anthropol. 4(2): 41-42.
Mello, M. T. de. 1995. Treinamento em Primatologia
no Brasil. Rev. Bras. Cignc. Vet. 2(3): 69-74.
Morrison, C. 1996. Enrichment for tree shrews and tama-
rins. Shape of Enrichment 5(1): 8.
Mudry, M. D., Fundia, A., Hick, A., Gorostiaga, M. A.
1995. Labilidad cromos6mica: una possible explicaci6n
en el origen de los reordenamientos cromos6micos en
cebidos. Bol. Primatol. Lat. 5(1): 7-15.
Nagamachi, C. Y., Pieczarka, J. C., Barros, R. M. S.,
Schwarz, M., Muniz, J. A. P. C. and Mattevi, M. S.
1996. Chromosomal relationships and phylogenetic
and clustering analyses on genus Callithrix, group
argentata (Callitrichidae, Primates). Cytogenet. Cell
Genet. 72: 331-338.
Norconk, M. A. 1996. Remembrance of Warren G.
Kinzey (1935-1994). Am. J. Primatol. 38:281-283.
Oerke, A.-K., Einspanier, A. and Hodges, J. K. 1996.
Noninvasive monitoring of follicle development, ovu-
lation, and corpus luteum formation in the marmoset
monkey (Callithrixjacchus) by ultrasonography. Am.
J. Primatol. 39: 99-113.
Oliveira-Filho, A. T. and Galetti, M. 1996. Seed preda-
tion of Cariniana estrellensis (Lecythidaceae) by black
howler monkeys, Alouatta caraya. Primates 37(1): 87-
Pieczarka, J. C., Nagamachi, C. Y. Barros, R. M. S. and
Mattevi, M. S. 1996. Analysis of constitutive hetero-
chromatin by fluorochromes and in situ digestion with

Neotropical Primates 4(2), June 1996

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Page 71 Neotropical Primates 4(2), June 1996

restriction enzymes in species of the group Callithrix
argentata (Callitrichidae, Primates). Cytogenet. Cell
Genet. 72: 325-330.
Poleschuk, V. F., Balayan, M. S., Sobol, A. V., Gulyaeva,
T. V., Titova, I. P. and Dokin. V. P. 1995. A breeding
colony of common marmosets (Callithrixjacchus) for
experiments with hepatitis. Primate Report 42:79-81.
Rendell, D. and Di Fiore, A. 1995. The road less trav-
eled: Phylogenetic perspectives in primatology. Evol.
Anthropol. 4(2): 43-52.
Saltzman, W., Schultz-Darken, N. J. an Abbott, D. H.
1996. Behavioral and endocrine predictors of domi-
nance and tolerance in female common marmosets,
Callithrix jacchus. Anim. Behav. 51(3): 657-674.
Sampaio, I, Schneider, M. P. C. and Schneider, H. 1996.
Taxonomy of the Alouatta seniculus group: Biochemi-
cal and chromosome data. Primates 37(1): 65-73.
Silvius, R. Park protection fragmented. Wildlife Con-
servation 99(1): 16.
Stoner, K. E. 1996. Habitat selection and seasonal pat-
terns of activity and foraging of mantled howling
monkeys (Alouatta palliata) in northeastern Costa
Rica. Int. J. Primatol. 17(1): 1-30.
Takai, M. and Anaya, F. 1996. New specimens of the
oldest fossil platyrrhine, Branisella boliviana, from
Salla, Bolivia. Am. J. Phys. Anthropol., 99(2): 301-
Townsend, W. R. and Ramirez, V. M. 1995. Cultural
teaching as an ecological data base: Murui (Witoto)
knowledge about primates. Latinamericanist, Center
for Latin American Studies, University of Florida,
Gainesville, 31(1): 1-7.
Torii, R., Abbott, D. H. and Nigi, H. 1996. Morphologi-
cal changes of the ovary and hormonal changes through
the ovarian cycle of the common marmoset (Callithrix
jacchus). Primates 37(1): 49-56.
Trtkova, K., Mayer, W. E., O'hUigin, C. and Klein. J.
1995. Mhc-DRB genes and the origin of New World
monkeys. Molec. Phylogenet. Evol. 4(4): 408-419.
Valencia, R. 1995. How will highly diverse Amazonian
forests change over time? Center for Tropical Forest
Science Fall: 3.
Waters, S. S. 1995. A review of social parameters which
influence breeding in white-faced saki Pithecia
pithecia in captivity. Int. Zoo. Yb. 34:147-153.
Wright, N. 1995. The challenge of hand-rearing a
Geoffroy's tamarin at the Cleveland Metroparks Zoo.
Proc. Nat. Conf AAZK (American Assocation Zoo
Keepers., Inc. (1994): 49-50.
Ximenez, M. F. F. M. and Sousa, M. B. C. 1996. Family
composition and the characteristics of parental care
during the nursing phase of captive common marmo-
sets (Callithrixjacchus). Primates 37(2): 175-183.
Zunino, G. E. 1996. AnAlisis de nacimientos en Alouatta
caraya (Primates, Cebidae), en el noreste de la Ar-
gentina. Museo Argentino de Ciencias Naturales "Ber-

nardino Rivadavia" e Instituto Nacional de
Investigaci6n de las Ciencias Naturales, Extra, Nueva
Serie(133): 1-10.


Webster, D. A. 1995. European Studbook for Black
Howlers (Alouatta caraya). No. 1. Bristol Zoo Gar-
dens, UK.
Webster, D. A. 1996. European Studbook for Black
Howlers (Alouatta caraya). No. 2. Bristol Zoo Gar-
dens, UK.


Savage, A. 1996. Proyecto Titi: Saving Colombia's en-
dangered cotton-top tamarin (Saguinus oedipus). Proc.
Nat. Conf AAZK (American Assocation of Zoo Keep-
ers., Inc.) 22: 196.
Zhong, F., Shi, J., Brady, A., Abee, C. etal. 1996. Iden-
tification of 56 short tandem repeat polymorphisms
for the squirrel monkey (Saimiri boliviensis). Invest.
Med. 44(l): 64A.

Selected abstracts of the 10th Congress of the
Associazione Primatologica Italiana (API), 26-28
Maggio 1994, Bussolengo (VR), edited by Gemma
Perretta, in Primate Report, 1995, 42.

Carli-Campa, M. C., Ardito, G., Crovella, S. Montagnon,
D. and Bulatti, M. Comparative analysis of alphoid
DNA restriction patterns in selected species of New
and Old World primates. p.4.
Casetti, R., Taglioni, A. and Perretta, G. Baseline blood
values in the marmosets (Callithrix jacchus): Preg-
nancy related changes. p.16.
Derme, V., Mudry, M., Romano, E. Nicolai, F. and
Ferrucci, L. Recent developments of cytomolecular
techniques applied to the study ofphiloevolutive rela-
tionships in primates. pp.21-22.
Gozzo, S., Perretta, G., Casetti, R., Taglioni, A. and
Rossiello, E. Morphological aspects of the nucleus
basalis of Meynert in the Callithrixjacchus. p. 19.
Guidi, C. and Visalberghi, E. Play behaviour in juvenile
tufted capuchin monkeys (Cebus apella). pp.8-9.
Melotto, S., Poffe, A. and Gerrard, P. A. Flight behaviour
of the captive common marmoset (Callithrixjacchus)
in different social situations. pp.11-12.
Poffe, A., Melotto, S. and Gerrard, P. A. Comparison of
four environmental enrichment strategies in captive
common marmosets (Callithrixjacchus). pp.24-25.
Riviello, M. C. The use of a feeding board as an envi-
ronmental enrichment device for tufted capuchin mon-
keys (Cebus apella). pp.23.
Riviello, M. C. and Misiti, A. An alternative to woodchip
as a foraging substrate for yufted capuchin monkeys

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Neotropical Primates 4(2), June 1996 Page 72

(Cebus apella). p.24.
Romano, E., De Stefano, G. F., Derme, V., Nicolai, F.
Ferrucci, L. and Mudry, M. Cytomolecular
characterisation of the New World primates: a study
of natural populations. pp.4-5.
Scalfari, F. and Chiarelli, B. Biochemical aspects of ol-
factory communication in primates: review of data and
perspectives of research. p. 17.
Schnell, C. R., Scott, L. and Pryce, C. The European
Marmoset Research Group an example of interdisci-
plinary communication. p.21.
Valente, M., Limongelli, E. and Visalberghi, E. Feed-
ing alone and feeding in group: When does company
make a difference? pp.10-11.
Vitale, A. and Santamaria, F. Response to a novel ob-
ject by socially-housed common marmosets (Callithrix
jacchus). pp.15-16.


International Primatological Society, Workshop on
Methods of Primate Conservation, 9-11 August 1996,
Chicago Zoological Society (Brookfield Zoo). For
primatologists from developing countries attending the
XVIth IPS/ASP International Congress of Primatology,
Madison, Wisconsin. Contact: Dr. Jeanne Altmann, Vice
President for Conservation, International Primatological
Society, Chicago Zoological Society, Brookfield Zoo,
Chicago, Illinois 60513, USA, Fax: 708 485 3532, e-
mail: bzconbio@ix.netcom.com, or Dr. Alison Jolly,
President, International Primatological Society,
Department of Ecology and Evolutionary Biology,
Princeton University. Princeton, NJ 08544, USA, Fax:
609-258-5381, e-mail: ajolly@arachne.princeton.edu.

XVIth Congress of the International Primatological
Society & 19th Conference of the American Society
of Primatologists, 11-16 August 1996, University of
Wisconsin, Madison, hosted by the Wisconsin Regional
Primate Research Center. Contact: Edith Chan,
Coordinator/Information, Wisconsin Regional Primate
Research Center, 1220 Capitol Court, Madison,
Wisconsin 53715-1299, USA. Tel: (608) 263-3500, Fax:
(608) 263 4031, e-mail: ipsasp-info@primate.wisc.edu.

Meeting of the Association of Primate Veterinarians,
16-17 August 1996, University of Wisconsin, Madison.
Contact: Edith Chan, Coordinator/Information,
Wisconsin Regional Primate Research Center, 1220
Capitol Court, Madison, Wisconsin 53715-1299, USA.
Tel: (608) 263-3500, Fax: (608) 263 4031, e-mail:

Ecological Summit 96, 19-23 August 1996,
Copenhagen, Denmark. Organized by Elsevier Science,

Journal Editors Robert Costanza (Ecological
Economics), Sven E. Jorgensen (Ecological Modelling),
William J. Mitsch (Ecological Engineering) and David
Rapport (Ecosystem Health). In collaboration with the
International Society of Ecological Modelling,
International Ecological Engineering Society,
International Society of Ecosystem Health, International
Society of Ecological Economics, SAS Institute
Denmark, and International Lake Environmental
Committee. For information contact: Ecological Summit
96, Conference Secretariat, Elsevier Science Ltd., The
Boulevard, Langford Lane, Kidlington, Oxford OX5
1GB, UK. Tel: +44 (0)1865 843643, Fax: +44 (0)1856
843958, e-mail: g.spear@elsevier.co.uk.

1996 Annual Meeting of the Conservation Breeding
Specialist Group CBSG/SSC/IUCN, 22-25 August
1996, Denver, Colorado, USA. Hosted by the Denver
Zoological Garden. Immediately preceding the Annual
Meeting of the World Zoo Organization IUDZG.
Registration contact: Angela Baier, Marketing Director,
Denver Zoological Foundation, City Park, 2300 Steele
Street, Denver, Colorado 80205-4899, USA. Tel: 1-303-
331-5805, Fax: 1-303-331-4125.

Conservation in a Changing World: Integrating
Processes into Priorities, 24-25 September, 1996,
Zoological Society of London, Regent's Park, London.
There is a widespread recognition of the urgent need to
identify priorities for conservation action. The last five
years have seen enormous progress in the development
of quantititative methods for identifying priority areas
based on what we know about species' distributions and
the relationships between areas and species. The
challenge now is to expand on this approach by building
an understanding of biological processes into
conservation planning. In particular, how can a
knowledge of past extinctions help to predict current
vulnerablility, and how ecological and evolutionary
processes can be incorporated into priorities for action?
Confirmed speakers include: Nick Nichols, Brian
Huntley, Jeremy Jackson, Andrew Balmford, Chris
Thomas, William Bond, Dan Janzen, Russ Lande, Stuart
Pimm, Jon Fjeldsa and David Pearce. The two-day
meeting costs 50 (25 for students). For further
information: Assistant Editor, Zoological Society of
London, Regent's Park, London NW1 4RY, UK. Tel:
+44 (0)171 448 6272, Fax: +44 (0)171 586 5321.

6th International Behavioural Ecology Congress, 29
September -4 October 1996, Canberra, Australia. Details
from: Andrew Cockburn, Division of Botany and
Zoology, Australian National University, Canberra ACT
02000, Australia. Fax: 61 6249 5773, e-mail:

Neotropical Primates 4(2), June 1996

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III Congresso de Ecologia do Brasil, 6-11 October
1995, Centro de Convenrges Ulisses Guimarles,
Brasilia. Deadline for submitting preliminary abstracts:
30 March 1996. Deadline for submitting final version
of abstracts: 30 June 1996. Contact: Comissio
Organizadora, III Congresso de Ecologia do Brasil,
Departamento de Ecologia, Universidade de Brasilia
(UNB), Caixa Postal 04355, 70919-970 Brasilia, D. F.,
Brasil. Tel: +55 (0)61 348-2326,348-2592, & 348-2282,
Fax: +55 (0)61 272-1497 & 273-4571. E.mail:

68th IUCN Species Survival Commission Full
Meeting, 11-12 October 1996, Montreal, Canada.
Theme: Communicating the value of the SSC its
worldwide presence, scientific knowledge, expert advice,
and ongoing work, and its relevance to the conservation
of biodiversity. Plenary sessions: SSC advice to
intergovernmental bodies; Biodiversity conservation
information system; SSC Specialist Group Reports.
Round table discussion: SSC at the regional and country
levels. Workshops: IUCN categories of threat; SSC
communications strategy; Fund-raising strategies.
Registration fee $25. For more information: World
Conservation Congress Coordinator, IUCN, Rue
Mauverney 28, 1196 Gland, Switzerland, Fax: + 41 22
999 0020.

IUCN World Conservation Congress, 13-23 October
1996, Montreal Conference Centre, Montreal, Canada.
Four distinct parts: Special Members' Session (13-14
October) to consider revised statutes accredited
delegates of IUCN voting members; Members' Business
Session (15-16, 22-23 October) to discuss and approve
IUCN's future strategy, programme and budget, elect
the officers and Council of the Union, and debate and
adopt resolutions and recommendations invited
observers may also attend; Open Session of Workshops
(17-18, 20-21 October) under the overall theme of
"Caring for the Earth" open to the public; A major
environmental exhibition open to the public. 19 October
set aside for excursions. Registration fee $50 if paid
before 31 July 1996, $100 after that date. Contact: John
Burke, Director of Communications, IUCN The World
Conservation Union, 28 rue Mauverney, 1196 Gland
Switzerland. Tel: +41 22 999 0123.

XIV Encontro Anual de Etologia, 16-19 October 1996,
Uberlandia, Minas Gerais, Brazil. Organized by the
Sociedade Brasileira de Etologia (SBEt) and the
Departamento de Biociencias, Universidade Federal de
Uberlandia. Includes symposia on: Human ethology;
insect behavior; behavior, animal production and
conservation; defensive behavior; reproductive behavior;
and aquatic mammals. Deadline for abstracts: 30 July
1996. For more information: XIV Encontro Anual de

Etologia, Coordenador Prof. Dr. Kleber Del-Claro,
Departamento de Biociencias, Universidade Federal de
Uberlandia, Caixa Postal 593, 38400-902 Uberlandia,
Minas Gerais, Brazil. Fax: (034) 232 8620, e-mail:

Measuring Behavior '96 International Workshop
on Methods and Techniques in Behavioral Research,
16-18 October 1996, Rudolf Magnus Insititute for
Neurosciences, Utrecht University, The Netherlands.
Registration fee: before 1 August 1996 is NLG 200
(students: NLG 50), after 1 August 1996 is NLG 300
(students: NLG 75). Submission of abstracts: Those who
wish to present an oral paper, poster or demonstration
should submit the title and abstract of their contribution.
All submissions should be received before 1 May 1996.
Notification of acceptance of abstracts 1 July 1996.
For program booklet and registration/abstract forms:
Measuring Behavior '96, Workshop Secretariat, Attn:
Rosan Nikkelen, P.O. Box 268, 6700 AG Wageningen,
The Netherlands. Tel: +31 (0)317-497677, Fax: +31
(0)317-424496, e-mail: mb96@noldus.nl. (Information
on the workshop is also available on the World Wide
Web: http://www.diva.nl/noldus/mb96.html).

I Congress APE and European Workshop on Primate
Research, 16-19 October 1996, Hotel Escuela, Madrid,
Spain. Organized by the Asociaci6n Prmatol6gica
Espafiola (APE). The European Workshop on Primate
Research, consisting of a panel of invited speakers and
free poster contributions, will take up the last two days
of the meeting. The objective of the Congress is to
provide a forum to assess the current situation and
perspectives on primate research in Spain and the rest
of Europe to facilitate the exchange of information
among European primatologists and to promote the
establishment of co-operative links between European
institutions and research groups working in primatology.
Confirmed speakers include: B. Deputte (Paimpont,
France), B. Thierry (Strasbourg, France), R. Vercauteren
Drubbel (Bruxelles, Belgium), R. D. Martin (Zuirich,
Switzerland), E. Visalberghi (Rome, Italy), P.
Timmermans (Nijmegen, Holland), L. Sterck (Utrecht,
Holland), W. Kaumanns (G6ttingen, Germany), M.
Vancatova (Konarovice, Czech Republic), H. Preuschoft
(Bochum, Germany), R. Crompton (Liverpool, UK), H.
0. Box (Reading, UK), G. Norton and D. Hawkins
(Cambridge, UK). Prof. Hans Kummer will also be
elected an Honorary Member of APE and will give a
talk entitled "Through the fieldglasses: a primatologist's
retrospective". For further information: Dr. Fernando
Colmenares, Departamento de Psicobiologia,
Universidad Complutense de Madrid, Campus de
Somosaguas, 28223 Madrid, Spain. Tel: +34 1 3943073,
Fax: +34 1 3943189, e-mail: pspsc06@sis.ucm.es.

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Neotropical Primates 4(2), June 1996

Page 74

XV Congress Panamericano de Ciencias
Veterin:irias, 21-25 October 1996, PalAcio Popular de
Cultura, Campo Grande, Mato Grosso do Sul, Brazil.
Associaqao Panamericana de Ciencias Veterindrias,
Sociedade Brasileira de Medicina Veterindria. Contact:
XV PANVET, Comissao Organizadora, Avenida Afonso
Pena 2386, Sala 84, 79002-074 Campo Grande, Mato
Grosso do Sul, Brazil. Tel: +55 (0)67 724-7071, Fax:
+55 (0)67 383-4371.

PSGB Winter Meeting 1996 Social Learning Among
Mammals, 29-30 November 1996, Meeting Rooms.
London Zoological Society, London. Organized by the
Primate Society of Great Britain (PSGB), in association
with the Mammal Society and the Zoological Society of

Biodiversity, Conservation and Management at the
Beni Biosphere Reserve, Bolivia, 3-6 December 1996,
La Paz, Bolivia. Organized by the Beni Biological
Station, Bolivian Academy of Sciences, and the
Smithsonian/MAB Biodiversity Program. The objective
is to provide a complete overview of the last ten years
of research on biodiversity, conservation and
management at the reserve. Papers and posters are
requested. Proceedings will be published. For additional
information, contact: Carmen Miranda, Academia
Nacional de Ciencias de Bolivia, Av. 16 de Julio 1732,
Casilla 5829, La Paz, Bolivia. Tel./Fax: (591-2) 350612,
e-mail cmiranda@ebb.bo, or Francisco Dallmeier,
Smithsonian/MAB Biodiversity Program, 1100
Jefferson Drive SW, Suite 3123, Washington, D. C.
20560, USA. Tel: (202) 357 4793, Fax: (202) 786 2557,
e-mail: icfgd@ic.si.edu.

ASAB Winter Meeting, Behaviour and Speciation,
5-6 December 1996, Zoological Society of London
Meeting Rooms, London Zoo. Organizer: Roger Butlin.
For further information contact: Dr Roger Butlin,
Ecology and Evolution Programme, Department of
Genetics, University of Leeds, Leeds LS2 9JT, UK.

Australian Primate Society XVIth Annual
Conference, 6-8 December 1996, Wellington Zoo,
Wellington, New Zealand. Conference Organizer:
Graeme Strachan, Wellington Zoo. Contact: Graeme
Crook, CSIRO Division of Human Nutrition, Animal
Services, Majors Road, O'Halloran Hill, South Australia
5158. Tel: +61 82980336, Fax: +61 83770004, e-mail:


1997 Meeting of the American Society of
Primatologists, 27 June 1 July, 1997, Bahia Hotel,
San Diego, California. For more information, contact:

Nancy Caine, Psychology Department, California State
University, San Marcos, California 92096, USA. Tel:
(619) 752-4145, Fax: (619) 752-4111, e-mail:


We would be most grateful if you could send us
information on projects, research groups, events
(congresses, symposia, and workshops), recent
publications, activities of primatological societies and
NGOs, news items or opinions of recent events and
suchlike. Manuscripts should be double-spaced and
accompanied by the text in diskette for PC compatible
text-editors (MS-Word, Wordperfect, Wordstar).
Articles, not exceeding six pages, can include small
black-and-white photographs, figures, maps, tables and
references, but please keep them to a minimum.

Please send contributions to: ANTHONY RYLANDS, c/o
Conservation International do Brasil, Avenida Ant6nio
Abrahao Caram 820/302, 31275-000 Belo Horizonte,
Minas Gerais, Brazil, Tel/Fax: +55 (31) 441 17 95 or
Fauna Silvestre Tropical, Universidad Veracruzana,
Apartado Postal 566, Xalapa, Veracruz 91000, Mexico,
Fax: 52 (28) 12-5748.

LILANA CORTES-ORTIZ (Universidad Veracruzana)
provides invaluable editorial assistance.

Correspondence, messages, and texts can be sent to:



NEOTROPICAL PRIMATES is produced in collaboration
with Conservation International, 1015 18th Street
NW, Suite 1000, Washington DC 20036, USA, and
FundalAo Biodiversitas, Av. do Contorno, 9155/11.
andar Prado, Belo Horizonte 30110-130, Minas
Gerais, Brazil.

Design and Composition ALEXANDRE S. DINNOUTI -
a.dinnouti@conservation.org.br Conservation
International do Brasil.

Neotropical Primates 4(2), June 1996

ISSN 1413-4703
^ \

Anthony Rylands/Ernesto Rodriguez Luna, Editors
Conservation International
Avenida Ant6nio Abrahlo Caram 820/302
31275-000, Belo Horizonte
Minas Gerais, Brazil

A Dmsion of the Houston
Paris and Recreation Department

This issue of Neotropical Primates was kindly sponsored by the Margot Marsh Biodiversity
Foundation, 432 Walker Road, Great Falls, Virginia 22066, USA, the Houston Zoological Gar-
dens Conservation Program, General Manager Donald G. Olson, 1513 North MacGregor, Hous-
ton, Texas 77030, and the Penscynor Wildlife Park, Manager Rob Colley, Cilfrew, Nr. Neath,
West Glamorgan, Wales SA10 8LF, U. K.

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