Title: Neotropical primates
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Title: Neotropical primates a newsletter of the Neotropical Section of the IUCNSSC Primate Specialist Group
Abbreviated Title: Neotrop. primates
Physical Description: v. : ill. ; 27 cm.
Language: English
Creator: IUCN/SSC Primate Specialist Group -- Neotropical Section
IUCN/SSC Primate Specialist Group -- Neotropical Section
Conservation International
Center for Applied Biodiversity Science
Publisher: Conservation International
Place of Publication: Belo Horizonte Minas Gerais Brazil
Belo Horizonte Minas Gerais Brazil
Publication Date: June Supplement 2002
Frequency: quarterly
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Subject: Primates -- Periodicals -- Latin America   ( lcsh )
Primates -- Periodicals   ( lcsh )
Wildlife conservation -- Periodicals   ( lcsh )
Genre: review   ( marcgt )
periodical   ( marcgt )
Spatial Coverage: Brazil
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Additional Physical Form: Also issued online.
Language: English, Portuguese, and Spanish.
Dates or Sequential Designation: Vol. 1, no. 1 (Mar. 1993)-
Issuing Body: Issued jointly with Center for Applied Biodiversity Science, <Dec. 2004->
General Note: Published in Washington, D.C., Dec. 1999-Apr. 2005 , Arlington, VA, Aug. 2005-
General Note: Latest issue consulted: Vol. 13, no. 1 (Apr. 2005).
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Volume ID: VID00039
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Neotropical Primates 10(Suppl.), June 2002


A TAXONOMIC REVIEW OF THE TITI MONKEYS, GENUS CALLICEBUS THOMAS, 1903, WITH
THE DESCRIPTION OF TWO NEW SPECIES, CALLICEBUS BERNHARDI AND CALLICEBUS
STEPHENNASHI, FROM BRAZILIAN AMAZONIA


Marc G. M. van Roosmalen1, Tomas van Roosmalen2, and Russell A. Mittermeier3

'Instituto Nacional de Pesquisas da Amaz6nia, Caixa Postal 478, Manaus 69083-000, Amazonas, Brazil.
2Center for Environmental Research and Conservation CERC, Columbia University, New York, NY 10027, USA.
3Conservation International, 1919 M St. NW, Washington, DC 20036, USA.


Abstract

This paper provides a taxonomic review of the titi monkeys, genus Callicebus, and describes two new species from central
Brazilian Amazonia, ( .... bernhardi and ( .... stephennashi. Previous revisions include Hershkovitz (1988, 1990),
Kobayashi (1995), Kobayashi and Langguth (1999) and Groves (1993, 2001). Here we arrange the titi monkeys, genus
C .... Thomas, 1903, into five Species Groups or clades, and a total of 28 species. The ( .... donacophilus Group
is represented by the following species: ( .... modestus Ldnnberg, 1939, ( .... donacophilus (d'Orbigny, 1836),
( .... '. .. Thomas, 1907 (treated by Hershkovitz as a subspecies of C. donacophilus), ( .... olallae Ldnnberg,
1939, and ( .... oenantheThomas, 1924. The ( .... moloch Group is represented by the following species: ( .-.
cinerascens (Spix, 1823), ( .... '...rr Thomas, 1908, ( .... baptista Lbnnberg, 1939 (treated by Hershkovitz
as a subspecies of C. '.n rr -. ( .... brunneus (Wagner, 1842), ( .... moloch (Hoffmannsegg, 1807), and the
new species here described as ( .... bernhardi. The ( .... cupreus Group is represented by the following species:
( .... cupreus (Spix, 1823), ( .... discolor (I. Geoffroy & Deville, 1848) (treated by Hershkovitz as a subspecies of
C. cupreus), ( .... ornatus (Gray, 1866) (treated by Hershkovitz as a subspecies of ( .... cupreus), ( .... caligatus
(Wagner, 1842), ( .... dubius Hershkovitz, 1988, and the new species here described as ( .... stephennashi. The
( .... torquatus Group is represented by the following species: ( .... torquatus (Hoffmannsegg, 1807), ( ..
lugens (Humboldt, 1811) (treated by Hershkovitz as a subspecies of C. torquatus), ( .... purinus Thomas, 1927 (treated
by Hershkovitz as a subspecies of C. torquatus), ( .... lucifer Thomas, 1914 (treated by Hershkovitz as a subspecies of
C. torquatus), ( .... regulus Thomas, 1927 (treated by Hershkovitz as a subspecies of C. torquatus), and ( .-.
medemi Hershkovitz, 1963 (treated by Hershkovitz as a subspecies of C. torquatus). The ( .... .. ... .- Group is treated
separately from the ( .... moloch Group and is represented by the following species: ( .... personatus (E. Geoffroy,
1812), ( .... melanochir (Wied-Neuwied, 1820) (treated by Hershkovitz as a subspecies of C. personatus), ( ..,.
nigrifrons (Spix, 1823) (treated by Hershkovitz as a subspecies of C. personatus), ( .... barbarabrownae Hershkovitz,
1990 (treated by Hershkovitz as a subspecies of C. personatus), and ( .... coimbrai Kobayashi & Langguth, 1999.
( .... bernhardi was discovered in the Rio Aripuand basin in 1996 by M. G. M. van Roosmalen, who also discovered
( .... stephennashi in 2001, while traveling on the Rio Purus. The geographic distributions of all hitherto recognized
( .... species are updated, and the validity of the river barrier hypothesis for titis and other Amazonian primates is
discussed, along with their conservation status.

Key Words Primates, Pitheciidae, Callicebus, titi monkeys, distribution, C .... bernhardi new species, .
stephennashi new species, Amazonia, Brazil.


Resumo

Este artigo apresenta uma revisao taxonomica dos macacos 'zogue-zogues', genero Callicebus, e descreve duas esp&cies novas
descobertas na Amazonia central, ( .... bernhardi e ( ... stephennashi. Revis6es recentes desse genero incluem
Hershkovitz (1988, 1990), Kobayashi (1995), Kobayashi e Langguth (1999) e Groves (1993, 2001). Aqui nos classificamos
zogue-zogues da Amaz6nia e sauAs da Mata Atlantica, genero ( .... Thomas, 1903, em cinco Grupos de Esp6cies, ou 28
clados: Grupo C. donacophilus (( .... modestus Lbnnberg, 1939, ( .... donacophilus [d'Orbigny, 1836], ( .-.
' .. Thomas, 1907 [considerada uma subesp&cie de C. donacophilus por Hershkovitz], ( .... olallae Lbnnberg,
1939 e ( ... oenanthe Thomas, 1924); Grupo C. moloch (( .... cinerascens [Spix, 1823], ( .... '...n -
Thomas, 1908, ( .... baptista Ldnnberg, 1939 [considerada uma subep&cie de C. '-.-rr . por Hershkovitz],
( .... brunneus [Wagner, 1842], ( .... moloch [Hoffmannsegg, 1807], e a esp6cie nova aqui descrita como
C .... bernhardi); Grupo C. cupreus (( .... cupreus [Spix, 1823], ( .... discolor [I. Geoffroy & Deville, 1848]
[considerada uma subesp&cie de C. cupreus por Hershkovitz], ( .... ornatus [Gray, 1866] [considerada uma subesp&cie
de C. cupreus por Hershkovitz], ( .... caligatus [Wagner, 1842], ( .... dubius Hershkovitz, 1988, e uma esp6cie





Neotropical Primates 10(Suppl.), June 2002


nova aqui descrita como ( ... -.. p i. i-. ,l-,i-,, Grupo C. torquatus(( .... .-.. .- [Hoffmannsegg, 1807], .
lugens [Humboldt, 1811] [considerada uma subespecie de C. torquatus por Hershkovitz], ( .... purinus Thomas, 1927
[considerada uma subespecie de C. torquatus por Hershkovitz], ( .... lucifer Thomas, 1914 [considerada uma subespecie
de C. torquatus por Hershkovitz], ( .... regulus Thomas, 1927 [considerada uma subespecie de C. torquatus por
Hershkovitz] e ( .... medemi Hershkovitz, 1963 [considerada uma subespecie de C. torquatus por Hershkovitz]; e o
Grupo C. personatus que foi inclufdo no Grupo moloch por Hershkovitz (( .... personatus [E. Geoffroy, 1812], ( ..
melanochir [Wied-Neuwied, 1820] [considerada uma subespecie de C. personatus por Hershkovitz], ( .... nigrifrons
[Spix, 1823] [considerada uma subespecie de C. personatus por Hershkovitz], ( .... barbarabrownae Hershkovitz,
1990 [considerada uma subespecie de C. personatus por Hershkovitz] e ( .... coimbrai Kobayashi & Langguth, 1999).
( .... bernhardi foi descoberta na bacia do rio Aripuana, Amazonas, por M. G. M. van Roosmalen em 1996. ( ..
stephennashi foi descoberta recentemente em 2000 pelo mesmo pesquisador no rio Purus, Amazonas. A probabilidade da
existencia de outras especies de 'zogue-zogues' ate entao desconhecidas na Amaz6nia esti discutida. A distribuigao geogrifica
de todas as especies esti atualizada. Discute-se a validade da hip6tese dos rios da bacia Amaz6nica como barreiras geogrificas
para os 'zogue-zogues' e outros primatas Amaz6nicos.

Palavras-Chave Primatas, Pitheciidae, Callicebus, zogue-zogues, distribuigao, C .... bernhardi especie nova, .
stephennashi especie nova, Amaz6nia, Brasil.




Introduction


The titi monkeys of the genus ( .... -, a diverse group
of Neotropical monkeys found mainly in the tropical forests
of the Amazon and Orinoco basins, but also extending into
the Atlantic forest region of Brazil, and the Chaco and dry
forests of Paraguay and Bolivia as far south as the Rios
Pilcomayo and Paraguay. They are small to medium in size,
roughly between the tamarins and the pitheciines, weighing
1-2 kilograms, and ranging from 270-450 mm in head-
body length (Hershkovitz, 1990). Locomotion consists
mainly of quadrupedal walking, climbing and leaping. All
modern revisions of this genus were by Hershkovitz, the
first being in 1963, followed by two other major papers
in 1988 and 1990. These were followed by a study by
Kobayashi (1995) and the description of anew species from
the Atlantic forest by Kobayashi and Langguth (1999).

Once considered a moderately diverse Neotropical genus,
( .... is now emerging as one of the most diverse of
all primate genera, competing with Saguinus for the largest
number of taxa in the New World. In Hershkovitz's (1963)
revision, he recognized only two species from the Amazon
and Orinoco river basins, ( .... moloch with seven
subspecies and C .... torquatus with three subspecies,
including one, .... .-.., .- .. described in that
paper. He did not treat Atlantic forest ( .... personatus
in 1963, but indicated that the three subspecies there might
be conspecific with ( .... moloch. Consequently, the
number of ( .... r i recognized as of 1963 was 13.

In 1988, Hershkovitz published a more detailed revision
of the ( .... in which he divided the genus into
four species groups (( .... modestus, ( ....
donacophilus, ( .... moloch, and ( .... torquatus),
with 13 species and 24 taxa, nearly double what was
recognized in 1963. This increase resulted mainly from
the resurrection of a number of taxa described earlier in


the 20' century by Thomas (1907, 1908, 1917, 1924,
1927) and Lbnnberg (1939) (modestus, pallescens, olallae,
oenanthe, cinerascens, baptista, lucifer, regulus, purinus),
and one by Wagner (1842; caligatus), but also included
the description of a new form, ( .... dubious. In
addition, four of the taxa recognized as subspecies of
( .... moloch in Hershkovitz (1963) were elevated
to full species status (donacophilus, '-..rr . brunneus,
cupreus), whereas three others continued to be considered
subspecies (baptista, discolor, ornatus). All taxa of ( ..
torquatus were treated as subspecies, including the three
recognized in 1963 torquatuss, lugens, medemi) and the
three resurrected from Thomas' earlier papers (lucifer,
regulus, purinus). Hershkovitz appears to have decided
whether a given taxon was a full species or just a subspecies
based mainly on distributional evidence indicating partial
sympatry.

Finally, in 1990, Hershkovitz published his final contribution
to ( .... taxonomy, a full revision in which he recognized
the same four species groups, 13 species and 24 taxa,
but adding a new subspecies of ( .... personatus,
C. p. barbarabrownae, for a total of 25. Although a
thorough review of the genus, this publication included
only a handful of color illustrations and a number of black
and white photographs of skins and live animals, the latter
mostly by Mittermeier, many of them published again here
in color along with many new color photographs. Given
the importance of coloration in these animals, we believe
it essential that color illustrations be provided for all taxa.
This we have done here by providing both pencil drawings
of every taxon and photographs of live animals wherever
available.

Since 1990, the only addition to our knowledge of
( .... taxonomy has been a review by Kobayashi





Neotropical Primates 10(Suppl.), June 2002


(1995) and a paper by Kobayashi and Langguth (1999),
in which they described a new C .... from the state of
Sergipe in northeastern Brazil, C .... coimbrai. In his
1995 paper, Kobayashi reviewed the taxonomy of the genus
based mainly on cranial measurements, and made several
modifications to Hershkovitz's earlier breakdown of species
into clades.

Groves (2001), although largely following Hershkovitz
(1990), also made some decisions on ( .... taxonomy.
In particular, he decided that C. moloch and C. cinerascens
are distinct; that C. brunneus, C. *'.-rr .. /.r -
C. *'..rr . baptista, and C. moloch are four full species;
that C. caligatus and C. cupreus cupreus are essentially
the same; and that C. dubius is not distinct at all. We
are in agreement with his first two observations, but
disagree with the second two, continuing to recognize
C. caligatus and C. dubius as distinct species. Like
Hershkovitz (1990), he placed them in four species
groups (C. modestus, C. donacophilus, C. moloch (including
C. personatus), and C. torquatus), but increased the number
of species to 15, while reducing the total number of taxa
to 24. He recognized all members of the C. modestus and
C. donacophilus groups and all members of his C. moloch
group except for C. personatus as full species. With
C. personatus, he recognized four of the five taxa as
subspecies, but agreed with Kobayashi and Langguth
(1999) that C. coimbrai is a distinct species. With the C.
torquatus group, he elevated C. medemito full species status,
but continued to list the other five taxa as subspecies.

In this paper, we provide a thorough reanalysis of the genus
Callicebus, complete with distribution maps and color
illustrations of each taxon. In addition, we describe two
new species from central Brazilian Amazonia, ( ...'
bernhardi and ( .... stephennashi. Together with
Kobayashi and Langguth's new taxon, ( .... coimbrai,
these new animals raise the total number of ( .... r. 28,
second only to the 33 taxa of Saguinus (Rylands etal., 2000)
among New World primate genera. Furthermore, since we
increasingly find the concept of subspecies to be of minimal
value in describing the diversity of Neotropical primates,
we have elevated all ( .... to full species status.

To enable the reader to better comprehend the evolution of
( .... taxonomy over the past 40 years, we also provide
a comparative table listing the ( .... r i recognized by
Hershkovitz in 1963, 1988 and 1990, Kobayashi in 1995
(including Kobayashi and Langguth, 1999), Groves in
2001, and in this paper (Table 1).


Geographic Distribution

The genus ( .... has a broad range in western and
central Amazonia, including the drainage basin of the Rio
Amazonas from the foothills of the Andes in Colombia,
Ecuador, Peru and Bolivia east into central Brazilian
Amazonia as far as the Rio Araguaia to the south of the


mainstream and the Rio Branco to the north. It also
occurs in the drainage basin of the Rio Orinoco from the
foothills of the Andes to the Rio Caroni to the south of
the mainstream of the Orinoco, including gallery forests
along Orinoco tributaries in the Colombian llanos. East
of the Rio Araguaia, there is a large gap in distribution in
easternmost Amazonia, the Cerrado and the Caatinga, until
one reaches the Atlantic forest region of eastern Brazil.
There, the ( .... personatus group has a large but highly
fragmented range from the state of Sergipe in tiny remnant
patches of northeastern Atlantic forest south through
Bahia, Minas Gerais, Espfrito Santo, and Rio de Janeiro, as
far as the Rio Tiete in Sao Paulo. South of Amazonia, the
genus extends into the dry forests and Chaco formations
of Paraguay and Bolivia as far as the Rios Pilcomayo and
Paraguay, the southernmost distribution on this side of the
continent being roughly parallel to the southern limits of
C .... personatus in the Atlantic forests of Sao Paulo.
Although present on the eastern slopes of the Andes, it
does not cross over to the Pacific side of the continent,
nor does it come close to extending into Central America.
Interestingly, it is also entirely absent from the northeastern
quadrant of Amazonia and is not found east of the Rio
Branco-Uraricoeira in Brazil nor in the three Guianas.


A New Taxonomic Arrangement for the Genus
Callicebus

Hershkovitz (1990) systematically arranged titi monkeys,
genus Callicebus, in four groupings or clades. Here, we
prefer to follow the more recent phylogenetic study, based
on cranial measurements, by Kobayashi (1995), which
includes ( .... modestus in the C. donacophilus clade,
splits the C. moloch clade into two clades, the C. moloch and
the C. cupreus clades, and puts ( .... personatus in its
own clade because of its long separation from Amazonian
titis both in historical and geographical terms, for a total of
five in all. Each of these is referred to here as a Species Group
named after the first described taxon from each group.

I. C. donacophilus Group
( ... donacophilus (d'Orbigny, 1836)
( .". I '. .. Thomas, 1907
( ... oenanthe Thomas, 1924
( ... modestus Lnnberg, 1939
( ... olallaeLbnnberg, 1939

This group, on average, is comprised of the smallest species
of Callicebus, and is closely related to the moloch group.
Brain-case volume of C. modestus is smallest for cebids,
and the species represents probably the most primitive
form among Cebidae. Diploid chromosome number for
C donacophilus= 50 (De Boer, 1974).

II. C. cupreus Group
( .... cupreus (Spix, 1823)
( .... caligatus (Wagner, 1842)
( .... discolor (I. Geoffroy & Deville, 1848)





Neotropical Primates 10(Suppl.), June 2002


ornatus (Gray, 1866)
dubius Hershkovitz, 1988
stephennashi new species


Diploid chromosome number only known for ( ....
cupreus, C. ornatus and C. discolor = 46 (De Boer,
1974; Schneider et al. 1993). See C moloch Group for
comparisons.

III. C. moloch Group
( .... moloch (Hoffmannsegg, 1807)
( .... cinerascens (Spix, 1823)
( .... brunneus (Wagner, 1842)
( .... '...rr Thomas, 1908
( .... baptista Lbnnberg, 1939
( .... bernhardi new species


The cupreus and moloch clades are composed of the typical
titis that were once regarded as conspecific (as subspecies of
( .... moloch; Hershkovitz, 1963). They belong to the
same so-called "eco-species" in the sense that they occupy
the same ecological niche. Members of this group are
invariably adapted to disturbed habitat and cannot occur
sympatrically. Sympatry only occurs between members
of the moloch cupreus groups and the torquatus group
species, which are adapted to undisturbed terra firme or
high dryland rainforest. Diploid chromosome number
only known for C. moloch and C. brunneus = 48 (Pieczarka
and Nagamachi, 1988; Minezawa et al., 1989; Schneider
et al., 1993).


Table 1. A comparison of the classifications of the titi monkeys, Callicebus, by Hershkovitz (1963), Hershkovitz (1988, 1990),
Kobayashi & Langguth (1999), Groves (2001), and this paper.

Hershkovitz (1988, Kobayashi (1995),
Hershkovitz (1963) 1990) Kobayashi & Langguth Groves (2001) This paper (2002)
(1999)
C. moloch moloch C. modestus Group C. donacophilus Group C. modestus Group C. donacophilus Group
C. m. cupreus C. modestus C. modestus C. modestus C. donacophilus
C. m. donacophilus C. donacophilus Group C. donaophilus donacophilus C. donacophilus Group C. pallescens
C. m. brunneus C. donacophilus donacophius C. d. pallescens C. donacophilus C. oenanthe
C. m. discolor C. d. pallescens C. olallae C. pallescens C. modestus
C. m. ornatus C. oenanthe C. cupreus Group C. oenanthe C. olallae
C. m. .. C. olallae C. cupreus cupreus C. olallae C. cupreus Group
C. torquatus torquatus C. moloch Group C. c. discolor C. moloch Group C. cupreus
C. t. lugens C. moloch C. c. ornatus C. cupreus cupreus C. caligatus
C. t. medemi C. cinerascens C. moloch Group C. c. discolor C. discolor.
C. cupreus cupreus C. moloch C. c. ornatus C. ornatus
C. c. discolor C. cinerascens C. moloch C. dubius
C. c. ornatus C. brunneus C. cinerascens C. stephennashi
C. caligatus C ... .. .. C. brunneus C. moloch Group
C. brunneus C. h. baptista C. C. moloch
C. ... ... C. personatus Group C. baptista C. cinerascens
C. h. baptista C. personatus C. personatus personatus C. brunneus
C. dubius C. melanochir C. p. melanochir C. -....
C. personatus personatus C. nigrifrons C. p. nigrifrons C. baptista
C. p. melanochir C. barbarabrownae C. p. barbarabrownae C. bernhardi
C. p. nigrifrons C. coimbrai C. coimbrai C. torquatus Group
C. p. barbarabrownae C. torquatus Group C. torquatus Group C. torquatus
C. torquatus Group C. torquatus torquatus C. torquatus torquatus C. lugens
C. torquatus torquatus C. t. lugens C. t. lugens C. lucifer
C. t. lugens C. t. lucifer C. t. lucifer C. purinus
C. t. lucifer C. t. purinus C .t. purinus C. regulus
C. t. purinus C .t. regulus C. t. regulus C. medemi
C. t. regulus C. t. medemi C. medemi C. personatus Group
C. t. medemi C. personatus
C. melanochir
C. nigrifrons
C. barbarabrownae
C. coimbrai





Neotropical Primates 10(Suppl.), June 2002


IV. C. torquatus Group
( ... torquatus (Hoffmannsegg, 1807)
( ..* lugens (Humboldt, 1811)
( ..* luciferThomas, 1914
( ... purinusThomas, 1927
( ..'. regulus Thomas, 1927
( ... medemiHershkovitz, 1963

This group is distinguished from all other titis by an overall
dark reddish to blackish color of the fur, by hair that is in
general uniform in color and not banded by a white collar
or throat, and by certain cranial and post-cranial skeletal
characters. All members of the torquatus Group belong
to the same so-called "eco-species," and are undisturbed
high dry-land (terra firme) rainforest habitat specialists
(Kinzey, 1977, 1981; Defler, 1994). Their diet is basically
frugivorous, but includes insects as well, whereas the other
Groups are partial folivores. Sympatry only occurs with
members of the cupreus clade south of the Rios Amazonas,
Solim6es, Marafion, Napo, Aguarico. Average size is larger


than donacophilus, cupreus and moloch Group members, but
slightly less than members of the personatus Group. Diploid
number of chromosomes = 20, the lowest for primates and
among the lowest for mammals in general (Benirschke and
Bogart, 1976; Egozcue et al., 1969).

V. C. personatus Group
( .... personatus (E. Geoffroy, 1812)
( .... melanochir (Wied-Neuwied, 1820)
( .... nigrifrons(Spix, 1823)
( .... barbarabrownae Hershkovitz, 1990
( .... coimbrai Kobayashi & Langguth, 1999

This group is composed of on average the largest species
of Callicebus, and inhabits coastal and inland forests of
southeastern Brazil. It is geographically separated from the
nearest member of the moloch clade (( .... moloch) to
the northwest by at least 1,000 km, and from the nearest
member of the donacophilus clade (C .... pallescens) to
the west by at least 500 km.


Figure 1. Distributions of the Amazonian titi monkeys, genus Callicebus, belonging to the donacophilus, cupreus and moloch Groups. The
asterisk marks the location of the municipality of Valparaiso, Columbia, where Moynihan (1976) recorded the presence of titi monkeys
not identifiable with discolor or ornatus. Map by Stephen D. Nash.




Neotropical Primates 10(Suppl.), June 2002


ALMtM" team$= .


Lm


*4


S'I Titi Monkeys
-- genus Calicebus
SSchemutc Map of the Diributon
o at mopocA, dowMApus,
and acuprvwQrupB


Figure 2. Schematic map of the distribution of Amazonian titi monkeys, belonging to the donacophilus, cupreus and moloch Groups.
Illustration by Stephen D. Nash.


kmmm




Neotropical Primates lO(Suppl.), June 2002


S I,.
n e -!


A

ti


'i n
I


f^v.
p- -W!-

r f----


Figure 3. The Amazonian taxa of titi monkeys, genus Callicebus, more or less geographically arranged. Illustration by Stephen D. Nash.


" 1~





Neotropical Primates 10(Suppl.), June 2002


SPECIES ACCOUNTS

I. C. DONACOPHILUS GROUP


Callicebus donacophilus (d'Orbigny, 1836)


Holotype: The holotype, an adult of unknown
sex, is depicted as "( .-'. donacophilus d'
Orb(igny)" in the "Atlas" of the Mammals of
the "Voyage dans l'Am&rique Mridionale,"
published in 1836 as a separate folio without
text. The animal must have been mounted
and exhibited in the galleries of the Musdum
National d'Histoire Naturelle in Paris, France,
but was not found in the museum collection
(Hershkovitz, 1990).

Type locality: Rio Mamord basin, province of
Moxos, Bolivia.

Geographic distribution: Upper Rios Mamor&-
Grande and San Miguel basins, Beni and Santa Cruz
provinces, west central Bolivia (Figs. 1 and 2).


Figure 4. The white-eared titi, Ci./,,, donacophilus
(d'Orbigny, 1836). An adult female in the Kilverstone Wildlife
Park, Thetford, UK, in 1981. Photograph by R. A. Mittermeier.


Diagnostic characters: Lacking distinct sideburns; upper
and outer parts of head and body, and outer and inner sides
of limbs buff or grayish agouti to dominantly orange agouti,
not contrasting; forehead like crown; blackish superciliary
fringe absent; most of chest and belly uniformly orange;
upper surface of cheiridia buffy or buffy agouti, paler than
forearms; tail buffy mixed with blackish, contrastingly paler
at base; ears very hairy, with whitish tufts.

Distinguished from ( .... .. ... by more saturated
coloration, whitish ear tufts, and less shaggy pelage; from
( .... brunneus by well-developed malar stripe, pale
agouti forehead, forearms, legs, cheiridia, paler under-parts
and lacking distinct sideburns; from ( .... modestus by
well-developed malar stripe, overall buffy to orange agouti
instead of light brownish to reddish agouti coloration
and lack of distinct sideburns; from ( .... olallae by
well-developed malar stripe, overall buffy to orange agouti
instead of reddish brown agouti coloration, lack of distinct
sideburns, and lack of a blackish facial fringe (Fig. 4).



Callicebus pallescens Thomas, 1907


IT^


S Holorvpe A L..tIIr Iv il, skin and skull, in the
ir.I, .! I., ui.. .. N tru. L History, no. 94.3.6.1,
...ll., ,.r, .. ,.r. .-., !v I 1-. J. B ohls.

Type localitr 'c. ,I, north of Concepci6n,


Geographic distribution: West of the Rio
i' iur, ..-.nrli r.. 'i-..r 23S and west to about
61U30'W, in the xeric forest of the northern and
central Chaco boreal in Paraguay, and in the
Pantanal, Mato Grosso do Sul, Brazil (Hershkovitz,
1990; Stallings, 1985) (Figs. 1 and 2).

Diagnostic characters: Trunk shaggy, pelage
extremely long, upper and outer sides of head and body,
and outer sides of limbs pale buff agouti; facial hairs nearly
concealing skin, malar stripe well-developed, blackish
superciliary fringe (almost) absent; tail pale buff agouti not
contrasting with rest of body.

Distinguished from ( .... donacophilus and other titis
by extreme pallor and shagginess of pale buff agouti coat;
tail, cheiridia, forehead, and outer side of limbs uniformly


P





Neotropical Primates 10O(Suppl.), June 2002


pale buff agouti, not contrastingly colored except for the
conspicuous whitish ear tufts (Fig. 5).


Diagnostic characters: Frontal blaze buffy or whitish,
continuous with long cresting whitish hairs bordering
the face; malar stripe present, whitish; sideburns, crown,
outer surface of limbs, cheiridia and tail uniformly and
dominantly to entirely dark brown agouti; inner surface of
limbs, chest, and belly orange; pelage thick, that of the face
longer than usual but not concealing the skin.

Distinguished from ( .... discolor by whitish or buffy
facial fringe or ruff of crested hairs, presence of a malar
stripe, and outer surface of limbs, cheiridia and tail dark
brown agouti.


Callicebus modestus Linnberg, 1939


Figure 5. C.di,, . .. (Thomas, 1907). Photograph by
B.A. Luscombe.


Callicebus oenanthe Thomas, 1924


Holotype: Adult male, skin and skull in the
British Museum of Natural History, London,
U.K., no. 24.7.11.1, collected January 1924 by
L. Rutter.

Type locality: Moyobamba, San Martin, Peru,
at 840 m altitude.

Geographic distribution: Northern Peru,
only known from the upper Rio Mayo valley,
Department of San Martin, altitudinal range
750-950 m (Figs. 1 and 2).


Lectotype: Adult male, skin and skull at Royal
Natural History Museum, Stockholm, Sweden,
no. A612105, collected under no. 135 by A. M.
Olalla in 1937; lectoparatype a sub-adult male,
skin and skull deposited in the same museum
(RNHMS), collected under no. 136 by A. M.
Ollalla in 1937.

Type locality: El Consuelo, Rio Beni, Beni,
Bolivia.

Geographic distribution: As far as is known,
it occurs only in the upper Rio Beni basin, a
tributary of the upper Rio Madeira, Beni, Bolivia. The
species is parapatric with ( .... dubius along the
north bank of the Rio Madre de Dios, with ( ..
donacophilus along the east bank of the Rio Beni, and with
C('" ., '. olallae along the west bank of the upper Rio
Beni (Figs. 1 and 2).

Diagnostic characters: Upper and outer parts of body light
brownish or reddish agouti except for white to whitish ear
tufts, reddish brown agouti forehead and crown, and thin
blackish superciliary fringe; outer surface of limbs reddish
brown agouti; hands and feet blackish or blackish mixed
with reddish; sideburns same color as forehead and crown;
tail blackish agouti, darker than dorsum.

Distinguished from ( .... brunneus by paler coloration,
whitish ear tufts, and dominantly blackish tail; from
( .... .' I.- overall light brownish or reddish agouti
instead of orange coloration, and face not framed with
blackish fur; from ( .... donacophilus by upper and





Neotropical Primates 10O(Suppl.), June 2002


outer parts of body light brownish or reddish agouti instead
of buffy to orange agouti, and dominantly blackish agouti
tail and cheiridia instead of buffy mixed with blackish.


Callicebus olallae Liinnberg, 1939


Distinguished from ( .... donacophilus by blackish
facial fringe, lack of malar stripe, weakly developed whitish
ear tufts, dominantly brownish or blackish cheiridia and
lack of a sharp contrast between coloration of under-parts
and sides of body; from ( .... modestus by blackish
facial fringe, individual hairs of back showing a broad
orange median band, and lack of well-developed whitish
ear tufts (Fig. 6).



II. C. CUPREUS GROUP


Callicebus cupreus (Spix, 1823)


S Holorkvpe Aulr ,il, -kin and skull in the Royal
SNiruL 'il Hi,r.... ILI 11 Ir. of Stockholm, Sweden,
,,.. V.,'!- ,...ll,,.r,d February 1938 by A. M.
1 i l t i ,., ,11 , I I

Type localir\ 1 1 1 i-una, una legua de Santa


Geographic distribution: Upper Rio Beni basin,
S i, ,, 11i; .. l, i l. I 0 .. 2).

S Diagnostic characters Facial fringe (sideburns,
1-, 1.. *in .i -, -, 1i.1. blackish, crown reddish
brown agouti; outer surface of limbs reddish brown;
cheiridia dominantly blackish; blackish suborbital vibrissae
conspicuous; back and limbs uniformly orange (hairs with
extremely broad orange median band); tail entirely dark
agouti sharply contrasting with orange back; whitish ear
tufts weakly developed.


Lectotype: Adult female, mounted, including
skull, Zoologische Staatssammlung, Mtinchen,
Germany, no. 10; lectoparatypes in the same
collection are no. 24, no. 89 a + b, all collected
January, 1820 byJ. B. von Spix.

Type locality: Right bank of Rio Solim6es,
Brazil, near the Peruvian border.

Diagnostic characters: Sideburns, sides of neck,
throat, inner surface of limbs, and underparts
of body uniformly reddish, sharply contrasting
with buff-brown agouti of dorsum and outer
sides of trunk, basal part of tail, and crown; forehead as
crown, reddish-brown agouti, often fringed with blackish
superciliary vibrissae and marginal hair bases.

Distinguished from other members of the cupreus Group,
except for C. caligatus and C. stephennashi, by absence of
distinct pale transverse frontal blaze, and from all members
of the cupreus Group by an overall dark tail with pencil
only white; from ( .... caligatus by lacking a broad
black frontal transverse blaze; from ( .... stephennashi
by lacking a black frontal transverse blaze that contrasts
sharply against a silvery crown, and having an overall
dark tail instead of a three-quarters to entirely buffy to
white tail; from ( .... brunneus by its red sideburns,
forearms, lower legs, cheiridia and under-parts of the body
(Fig. 7).


Figure 6. Olalla Brothers' titi monkey, C.'n!,,, olallae
Lbnnberg, 1939.





Neotropical Primates 10O(Suppl.), June 2002


Callicebus caligatus (Wagner, 1842)


IT Lectotype: Skin and skull said to be collected at
Borba, Rio Madeira, Amazonas, Brazil, restricted
S by Thomas (1908), and two lectoparatypes,
including one skin only from Borba, and one
skin and skull from Manaquirf, right bank of
'., the Rio Solim6es, Amazonas, Brazil, collected by
J. Natterer in December 1832; both deposited in
the Naturhistorisches Museum, Wien, Austria.

Type locality: Borba, right bank of Rio Madeira,
Amazonas, Brazil, restricted by Thomas (1908).
Since only ( .... cinerascens occurs in this
region along the right bank of the Rio Madeira,
"W Hershkovitz (1990) assumed that Natterer should have
ST collected and mislabelled these specimens elsewhere. Since
M.G.M. van Roosmalen found the species to occur in the
lower Rios Purds/Solim6es/Madeira interfluve south as
far as the Rio Ipixuna, the specimens collected at Borba,
right bank of the Rio Madeira, must have originated from
the left bank of the Rio Madeira in the vicinity of Borba.
The other type locality, Manaquirf, right bank of the Rio
Solim6es, falls within the supposed distribution.

Geographic distribution: Central Amazonia, Amazonas
state, Brazil, south of the Rio Solim6es in the interfluve
delineated by the lower Rios Purdis, Solim6es and Madeira,
south as far as the Rio Ipixuna (or Paranapixuna).
Hershkovitz (1990) considered the species sympatric with
( .... dubius, C. brunneus, and C. cupreus, based on,
in his view, mislabelled specimens collected by the Olalla
brothers at Boca Rio Inuya, Iquitos, the Rio Orosa, Rio
Tapiche, and Sarayacu, Rio Ucayali, in the department
of Loreto, Peru. Voss and Emmons (1996) note that
Hershkovitz' report on the sympatry of two members of
the moloch/cupreus Group (C. cupreus and C. caligatus) was
an error caused by "inadvertently listing both original and
revised identifications of the same series from Orosa among
the specimens examined." The correct identification for
the monkeys is ( .... (cupreus) cupreus. Specimens
collected by Peres (1993) along both banks of the Rio
Jurui and deposited in the Musdu Goeldi, Belkm, all were
identified as ( .... (cupreus) cupreus.

The distribution of C. caligatus contradicts that given by
Figure 7. The red or coppery titi, (.i/,,,'n cupreus (Spix, Hershkovitz (1990), who fills in its actual distribution
1823). Top photograph by Chris Zagazzo. Middle and Bottom with C. dubius. Four specimens of real C. caligatus were
photographs by R. A. Mittermeier. caught in July 2001 by locals along the north bank of


All





Neotropical Primates 10(Suppl.), June 2002


Figure 8. The chestnut-bellied titi, (.CAl ,. caligatus (Wagner, 1842), an adult male and female caught from the wild by local people
on the west bank of the Lago Jarf, Rio Purus, on July 27th, 2001. Upper right shows close up of the hands, and lower right, the tail.
Photographs by M. G. M. van Roosmalen.


Lago Jarf at the mouth of Igarapd Bacaba and released at
the same place after being measured and photographed,
and a complete skeleton with some hair found on
the forest floor was collected (private collection no.
MGMR55) on the north bank of Rio Ipixuna near its
mouth, a few kilometers south of the town of TapauA
(Van Roosmalen and Van Roosmalen, in prep.). (Figs.
1 and 2).

Diagnostic characters: Forehead and anterior part of
crown black, rest of crown dark red-brown as neck, back
and sides of body, each hair red-brown with a black tip;
sideburns, under-parts and inner sides of limbs reddish
to red-brown; back reddish-brown agouti; forearms and
lower legs dark red-brown; cheiridia, including wrists
and ankles black as blaze and basal part of tail; proximal
10 cm of tail dark black, next 25 cm blackish mixed with
buffy (hairs blackish with 0.7 cm long white tip), distal
15 cm forming a buffy pencil.

Distinguished from ( .... stephennashi by dark red-
brown agouti instead of bright reddish forearms, forelegs,
sideburns, and under-parts, black-brown to black instead
of buffy to white cheiridia, and forehead and anterior
part of crown not sharply contrasting with rest of
crown, nape, and sides of body; from C. cupreus, with
which it is parapatric along the lower Rio Purds in the
west, by entirely black forehead and anterior part of
crown, dark black-brown cheiridia, and the dominantly
buffy tail mixed with blackish, and the white pencil;
from C. dubius, by black forehead and anterior part of
crown lacking a contrasted pale or whitish frontal tuft
or transverse blaze; from C. cinerascens, with which it is
parapatric along the lower Rio Madeira in the east, by


reddish to dark red-brown sideburns, under-parts and
inner sides of limbs, and the dominantly buffy tail and
white pencil (Fig. 8).



Callicebus discolor (I. Geoffroy & Deville, 1848)


Holotype: Skin and skull of the holotype,
collected in 1847 by Comte Francis de Castelnau
I and Emile Deville, were originally deposited
in the Musde National d'Histoire Naturelle,
Paris, France, but are no longer in the museum
Collection.

Type locality: Sarayacu, left bank ofRfo Ucayali,
Ucayali, Peru.

Geographic distribution: Upper Amazonian
region in Peru, south of the Rfo Maranon in the
entire interfluve delineated by the Rfos Ucayali
and Huallaga, and north of the Rfo Maranon between the
Rfos Napo and Santiago; in Ecuador from the Andean
foothills east to the Rfo Napo/Aguarico basin, and north
to the Rfo Putumayo, and in Colombia to the right bank





Neotropical Primates 10(Suppl.), June 2002


of the Rfo Guamuds (Hernandez-Camacho and Cooper,
1976). Brooks and Pando-Vasquez (1997) recorded
this species just to the north of the Rio Napo on the
left bank of its northern tributary, the Rio Sucusari, in
Peru. Hernandez-Camacho and Cooper (1976) recorded
its range in the Colombian trapezium, between the
Rfos Putumayo and Amazonas. In Figure 1, C. discolor is
given as occurring throughout the interfluvium between
the Rios Putumayo-Iia and Amazonas-Solim6es
extending into Brazil, although this has yet to be
confirmed. In the eastern part of its range the species is
parapatric with C .... cupreus along the Rfo Ucayali
(Figs. 1 and 2).

Moynihan (1976, p.75) mentions the presence of titi
monkeys of the moloch Group in the close vicinity of
the town of Valparaiso, between the Rios Caqueta and
Orteguaza in Colombia. These "lacked the white stripe
above the eyes that is typical of both ornatus to the north
and discolor to the south," and Moynihan indicated that
they could be an unnamed subspecies (Fig. 1).

Diagnostic characters: Forehead with white or buffy tuft,
contrasting with dark-brown transverse band, or blaze,
this sharply contrasting with reddish crown and sideburns;
often small white patches present alongside the lower jaw
sharply contrasted with the reddish sideburns; sideburns,
crown, side of neck, forearms, lower legs, cheiridia, chest


Figure 9. (.inI/,t discolor (I. Geoffroy & Deville, 1848), an
adult female from unknown locality donated to M.G.M. van
Roosmalen in 1988. Photographs by M. G. M. van Roosmalen.


and belly reddish, sharply contrasted with agouti back,
sides ofbody, and tail; tail mixed brownish and buff-agouti,
distal one-third to three-quarters predominantly buffy
or white.

Distinguished from .... oenanthe and C. ornatus by
entirely reddish forearms and cheiridia, from C. cupreus
by one-third to three-quarters of tail buffy, white or
buffy frontal tuft, and dark-brown blaze or transverse
band (Fig. 9).


Callicebus ornatus (Gray, 1866)


Holotype: Skin and skull purchased from Maison
Verraux, Paris, France, and deposited as no.
1859.7.9.4 in the British Museum of Natural
History, London, U.K.

Type locality: Villavicencio region, Rio Meta,
Department of Meta, Colombia.

Geographic distribution: Eastern Colombia,
from the department of Cundinamarca (Medina)
north as far as the lower Rio Upfa/Rio Meta,
and south into the department of Meta, along
the base of the Cordillera Oriental and the
Sierra de la Macarena to the Rio Guayabero/upper Rio
Guaviare. This species is the only member of the cupreus
and moloch Group occurring north of the Rios Amazonas/
Solim6es/Napo axis, in the upper Rio Orinoco basin and
is separated from its nearest other member of the cupreus
Group (( .... discolor) in the south by at least 350 km
(Figs. 1 and 2).

Diagnostic characters: Ears and frontal tuft, transverse
band or blaze whitish, sharply contrasted with reddish
brown crown, the hair bases buffy; sideburns, under-parts,
and inner side of limbs reddish; outer surface of thighs,
upper arms, upper legs and sides of body buff-agouti;
outer side of lower legs and lower part of forearms reddish,
sharply contrasted with pale to whitish feet or toes only,
and hands or digits only, respectively; proximal one-quarter
of tail dark red-brown, distal three-quarters white.

Distinguished from ( .... cupreus and C. discolor by
white ears, and pale or whitish digits and toes sharply
contrasted with reddish wrists and ankles (Figs. 10, 11, and
12).





Neotropical Primates 10(Suppl.), June 2002


Figure 10. The ornate titi monkey, Cu.'/, ,,:, ornatus (Gray,
1866). Photograph by R. A. Mittermeier in 1992.


Figure 11. The ornate titi monkey, C.'/,i,:, ornatus (Gray,
1866). An adult male from north of La Macarena, Colombia.
Photographs by T. Defler at the Capari Biological Station, Rio
Apaporis, in 1992.


Figure 12. The ornate titi, C(./ ,,l : ornatus (Gray, 1866).



Callicebus dubius Hershkovitz, 1988


4 A -
*E. a..


H'lf


Holorvpe-' .lulr ti i.. ile, skin and skull, Field
N.M 1 ur., .'. N ru,-' History, Chicago, no.
.s.s. ...... 11 ,.r, d ..I ., t 1 Lako in June 1931.


I Type locaitv 11,..1 ro be the bank of Lago
'. Ipi r1. i, lI... t rl1I lower Rio Purds, but since
rl,.i ...l. ir. ,,p..... ble (hence it would have
.... n ...] 11' ir H -likovitz thought to be the
distribution of both C. caligatus and C. cupreus),
Hershkovitz (1990) assumed it originated from
the right (east) bank of the Rio Purds, opposite
Lago AyapuA (coordinates approximately 04o20'S,
6200'W). Native hunters must have supplied the
animal collector Carl Lako with specimens from
either side of the Rio Purds and from far upstream, since
M. G. M. and T. van Roosmalen recently located the
species along the Rio Seruinf on the right bank of the
Rio Purds, south of the Rio Ituxf. Some specimens in the
British Museum originated from the vicinity (probably
southwest) of the town of Humaita on the left bank of the
Rio Madeira, the source of the Rio Ituxf.

Geographic distribution: South of the Rio Ituxf, or maybe
even the Rio Mucufm, both right bank tributaries of the
Rio Purds, east as far as the Rio Madeira south of the
town of Humaita, and west to the Rio Purds, southern
limit unknown. The species might be parapatric with
( .... stephennashi along the Rio Ituxf or maybe the Rio
Mucufm in the northern part of its range, with ( .-
brunneus along the upper Rio Madeira in the southern or





Neotropical Primates 10(Suppl.), June 2002


southeastern part of its range, and with ( .... cupreus
along the Rio Purds in the west (Figs. 1 and 2).

Diagnostic characters: Buffy or whitish frontal tuft, blaze,
or transverse stripe, bordered below by blackish superciliary
vibrissae forming narrow black line connecting the blackish
ears, the crown brownish agouti; hairs of crown, nape,
back, and rump with 4-5 narrow pheomelanin bands, each
alternating with a eumelanin band; outer sides of thighs and
upper arms brownish agouti like back; sideburns, sides of
head, and beard deep reddish, outer surface of forearms and
lower legs reddish; hairs of throat, chest, belly, and inner
side of limbs not banded reddish to reddish brown; cheiridia
blackish agouti, the fingers and toes contrasted pale or white;
proximal one-third of tail red-brown agouti, as dorsum, rest
of tail blackish, with a contrasted white pencil.

Distinguished from C. caligatus by its white or buffy frontal
tuft or blaze, and lack of the black forehead and anterior


Figure 13. C.d/, ,;,: dubius Hershkovitz, 1988, pet monkey from
the Rio Seruinf, right bank of the Rio Purts, photographed in
Pauinf, July 2001. Photographs by T. van Roosmalen.


part of crown; from C. cupreus, with which it is parapatric
along the Rio Purds in the west by its white frontal tuft or
blaze, and its white digits; from C. brunneus by its white or
buffy frontal tuft or blaze and the contrasted white or buffy
tail pencil; from C. stephennashi by its white or buffy frontal
tuft or blaze and white digits only instead of entirely white
cheiridia (as far as the wrists and ankles) (Fig. 13).



Callicebus stephennashi new species


Holotype: Adult female, entire animal preserved
in alcohol, Mammal Collection of the National
Institute for Amazon Research, INPA no. 4030,
alive weighing 725 g, brought to M.G.M. van
Roosmalen by fishermen on June 20, 2000.
It died in November, 2000, in Manaus,
Amazonas, Brazil.

Paratypes: Four individuals, brought to
M.G.M. van Roosmalen by fishermen, said to
be caught along the middle or upper Rio Purds,
kept in captivity for several months, then all
died from dengue fever in November 2000.
Adult male weighing 780 g, INPA no. 4031, three females
weighing 740, 480, and 725 g, private collection numbers
MGMR 51, 52 and 53, respectively, Manaus-Amazonas,
Brazil (Table 2).

Type locality: Unknown. Holotype said to be caught
somewhere along the middle to upper Rio Purds together
with the paratypes.


Table 2. Measurements of specimens of Callicebus spp. collected by M.G.M. van Roosmalen.
Callicebus Head & body Tail Hind foot Ear Weight (g) Reference
(mm)
caligatus 290 510 90 31 1000 MGMR 58
stephennashi 270 420 85 35 780 INPA 4031
stephennashi 280 420 90 30 725 INPA 4030
stephennashi ? ? ? ? 740 MGMR 51
stephennashi ? ? ? ? 480 MGMR 52
stephennashi ? ? ? ? 725 MGMR 53
bernhardi 360 550 100 35 1200 INPA 4033
bernhardi 370 ? 90 30 700 INPA 4029
cinerascens 415 ? 95 30 740 INPA 4085




Neotropical Primates 10(Suppl.), June 2002


A


Figure 14. Stephen Nash's titi monkey, C.ii'// stephennashi new species. The adult female holotype, INPA 4030, and adult females
MGMR 51 and MGMR 53. Photographs by M. G. M. van Roosmalen.





Neotropical Primates 10O(Suppl.), June 2002


Geographic distribution: Since the new species seems
to be phenotypically most closely related to both
C. caligatus and C. dubius, we assume ( ....
stephennashi should occur along the right bank of the
Rio Purds in between the distributions of C. caligatus
and C. dubius. As a possible distribution, we suggest the
interfluve delineated by the Rios Purds/Ipixuna/Madeira/
Mucufm (Figs. 1 and 2).

Diagnostic characters: Forehead, superciliary vibrissae,
and anterior portion of crown black, sharply contrasting
with posterior portion of crown, nape, dorsum and rump
which are silvery or buffy mixed with brownish agouti
or brownish-black; ears blackish; lower arms and legs
bright red, sometimes dark red as in C. caligatus and
C. baptista, like sideburns, under parts, and inner sides
of limbs, contrasting with silvery or buffy mixed with
brownish agouti upper and outer parts; upper surface
of cheiridia silvery buffy to white, the proximal third
of foot red like lower legs, the hands entirely silvery




a






h


or white; proximal portion of tail silvery mixed with
brownish agouti to blackish brown, then black mixed
with white or buffy, and distal half to two-third entirely
white or buffy.

Distinguished from C. caligatus by black superciliary
vibrissae, forehead and anterior portion of crown sharply
contrasting with silvery agouti remainder of crown,
nape and upper parts of body, by bright red instead of
dark red-brown lower arms and legs, by white or buffy
instead of blackish cheiridia, and almost entirely white or
buffy tail; from C. dubius by black forehead and anterior
portion of crown, lacking a white blaze, by white or buffy
cheiridia instead of white digits only, and almost entirely
white or buffy tail; from C. cupreus by black frontal blaze
contrasting sharply against a silvery crown and nape,
almost entirely white or buffy tail instead of an overall dark
tail with white pencil, and by white or buffy instead of
brown agouti cheiridia; from C. brunneus by its bright red
sideburns, lower arms and legs, and under parts, white or




d










e


Figure 15. Stephen Nash's titi monkey, Cill // stephennashi new species. Some details of the adult female holotype, INPA 4030. a. left
hand, b. legs, feet and basal part of tail, c. feet aside of tail, d. middle part of tail against background of sides of body, e. detail of pelage on
side of body, f. detail of pelage of underparts and lower legs. Photographs by M. G. M. van Roosmalen.





Neotropical Primates 10O(Suppl.), June 2002


buffy cheiridia, and silvery agouti to silvery brown upper
parts and sides of body (Figs. 14, 15, and 16).

Measurements: See Table 2.

External characters of holotype: Black of forehead and
superciliary vibrissae continuing over anterior portion
of crown as in C. caligatus, but sharply contrasting with
posterior portion of crown, nape, back and rump which
are buffy to silvery mixed with brownish agouti, the lax
hairs cresting against shorter black hairs in front and
longer raised nuchal hairs behind; ears blackish; hairs of
dorsum and sides of body silvery mixed with brownish
agouti, the individual hairs 4 cm long, six-banded as
follows: proximal 1.5 cm brownish agouti to black-brown,
then 3 white or silvery bands each alternating with a brown
or blackish band, the tip of each hair silvery; lower arms
and legs uniformly bright red like sideburns, sides of head,
beard, under parts, and inner sides of limbs, contrasting
with upper arms and thighs which are buffy to silvery
mixed with brownish agouti; upper surface of cheiridia
silvery white to buffy, the proximal (ankle) part of foot
(3 cm) red like lower legs, the remaining 6 cm silvery white
to buffy, upper surface of entire hands including wrists
silvery or white; basal proximall) one-sixth (ca. 8 cm) of
tail same color as lower back and rump, next one-sixth
(ca. 8 cm) black mixed with white or buffy (black hairs
with 1 cm long white tip), remainder two-third entirely
white or buffy; hairs of sideburns, throat, chest, belly,
and inner side of limbs not banded red; blackish face
naked except for fine buffy hairs surrounding lips and
between nostrils.

Origin of the name: This titi monkey is named in honor
of our close friend and colleague, artist Stephen D. Nash,
Technical Illustrator for Conservation International's
President's office and its Center for Applied Biodiversity
Science (CABS). He is based at the State University of


Figure 16. Stephen Nash's titi monkey, (.in,,, stephennashi
new species. The adult female holotype, INPA 4030, and adult
females MGMR 51 and MGMR 53. Photographs by M. G. M.
van Roosmalen.


New York at Stony Brook. For the past 20 years, Stephen
has made major contributions to primate conservation and
the science of primatology through his wonderful scientific
illustrations and his educational materials, which have been
widely distributed around the world.

Vernacular name: Titi monkeys are referred to as zog-zogs
or zogue-zogues by the local people in Amazonia. For an
English name, we suggest Stephen Nash's titi monkey.



III. C. MOLOCH GROUP


Callicebus moloch (Hoffmannsegg, 1807)


A

~I S


S vnrpes: ,\ i in dividuals collected by Mr.
,,. .1 .ii I ,,,,S donated by Count von
H. .-.i o -., __ r.. rlZoologisches Museum der
HuniO-b..l..r-i--ii.i -irt, Berlin, Germany, and
ii, i..t ,r 1i, I.. .. il mounted with skull in skin,
... -.1i22i '. ihisum National d'Histoire
Natulldk, i'aiL, fiance.

Type locality: Near the town of Belkm, Para,
Brazil (Hoffmannsegg, 1807).

Geographic distribution: Brazilian Amazonia
south of the Rio Amazonas in the States of
Para and Mato Grosso. In Para, from the west bank
of Rio Tocantins/Araguaia west as far as the east
bank of Rio Tapaj6s,
south as far as Ilha
do Bananal, north
of the confluence of
Rio das Mortes with
the Rio Araguaia;
in Mato Grosso, as
far west as the Rio


Juruena, including the
headwaters of the Rio
Xingti (M. G. M. van
Roosmalen collected
a specimen shot by a
Waura Indian hunter
along Rio Von den
Steinen) (Fig. 18). In
the northwestern part
of its range, the species


Figure 17. Captive adult red-
bellied titi, ( .... moloch
(Hoffmannsegg, 1807). Photograph
by R. A. Mittermeier.





Neotropical Primates 10O(Suppl.), June 2002


is parapatricwith C. .'...r . il.. i_ the lower RioTapaj6s,
and in the southwestern corner of its range it is parapatric
with C. cinerascens along the upper Rio Juruena (Figs. 1
and 2).

Diagnostic characters: Upper and outer surface of head,
trunk, and limbs buffy or grayish to pale brown agouti;
forehead not sharply defined from grayish crown or
distinctly paler, lacking whitish ear tufts; sideburns, under
parts of body, and inner side of limbs sharply contrasting
light orange to buff-orangish; cheiridia not sharply
contrasting buffy; hairs of tail blackish agouti terminally,
orange or buffy basally, distal half of tail including pencil
buffy.


Distinguished from C. cinerascens by uniformly light
orange instead of grayish sideburns and inner sides of
limbs, chest, and belly; from C. brunneus by grayish agouti
instead of dark-brown to black forehead and crown; from
C. *'...r . by rather contrasting buffy instead of
undefined blackish agouti upper surface of hands and
feet, and distal half of tail buffy instead of entirely blackish
agouti to black; from C. bernhardi by lack of silvery to
whitish ear tufts, light orange to buff-orangish instead of
dark orange sideburns, under parts of body, and inner side
of limbs, upper and outer surface of head, trunk, and limbs
buffy or grayish to pale brown agouti instead of blackish
agouti mixed with brown on the back, the distal half of
the tail buffy instead of an entirely blackish agouti to black
tail with sharply contrasted white pencil, and buffy upper
surface of hands and feet slightly paler than outer side
of arms and legs much less strikingly contrasted than in
C. bernhardi with its white upper surface of hands and feet
(Fig. 17 and 18).


rr m~
r


Callicebus cinerascens (Spix, 1823)


Figure 18. Specimen of the red-bellied titi monkey, C.('/,u,,"
moloch (Hoffmannsegg, 1807) from the left bank of Rio Von den
Steinen, upper Rio Xingd Indigenous Park, Mato Grosso, Brazil,
representing the most southerly limit to its range. The skin was
being used by a Wauri Indian hunter to make a headband. In
the private collection of M. G. M. van Roosmalen (MGMR 56).
Photographs by M. G. M. van Roosmalen.


a


Holorvpe. \ ,, I.. ..II, ctedbyJ.vonSpixbutnot
-_' 1. 1i i d ". i, w' the area during January,
1.s 2 i i,'Li ... ...i n'..ii ding the skull), No. 3,
/, ,I,.., h,. r ,i Lr ammlung, Mtinchen,


SType localir 'pi' '" issued to have collected
I" ,.. ....I. dil..' rhle Rio Putumayo or Rio
i i r' 1 i. I LI iii border, State of Amazonas,
S i 1i bur rl'> u1 no evidence that it was
., i.ruill .. ...11 .r ..I r- ,-e. Specimens from the
i,.. '.!...,1 ., .h.. ith of the Rio Amazonas
perfectly match the description on account of
the holotype. Therefore, the type locality given
by Spix must be wrong.

Geographic distribution: Hershkovitz (1990) includes
three localities in his gazetteer: Prainha, right bank of Rio
Aripuana, Amazonas, Brazil (162a); Sao Joao, right bank of
Rio Roosevelt (Hershkovitz gives here Rio Aripuana, but
his map indicates the Rio Roosevelt, a left bank tributary
of Rio Aripuand), Mato Grosso (205); and Otoho, right
bank of upper Rio Roosevelt (Hershkovitz gives here Rio
Ji-Parana, but his map indicates Rio Roosevelt, Rondonia).
This species has been observed in the wild by M.G.M.





Neotropical Primates 10O(Suppl.), June 2002


van Roosmalen at the following localities along the right
bank of Rio Aripuand: Cipotuba, situated on the east bank
of Lago Cipotuba (0548'23"S, 6012'76"W), Prainha,
Igarape da Prainha (0545'S, 6012'W), Sao Joao, Igarape
Terra Preta (0528'S, 6022'W), and along the right bank
of the Rio Madeira in the vicinity of the town of Novo
Aripuana (0507'08"S, 6022'45"W), left bank flower Rio
Arara (40 km E ofNovo Aripuana, 0512'S, 60004'W), and
in the vicinity of the town of Borba (04o22'S, 5935'W).
Rylands (1982) observed C. cinerascens on the east bank
of the Rio Aripuana at the Nicleo Pioneiro de Humboldt,
Aripuana, (then of INPA) (1010'S, 5927'W). M. G. M.
van Roosmalen kept a live specimen from the left bank of
the Rio Canuma, which was deposited in the Mammal



a f


c h


Figure 19. The ashy black titi, C./ii/ cinerascens (Spix, 1823).
Details of the dorsal (a-g) and ventral (h-j) pelage of an adult
male from the left bank of the Rio Canuma, a right bank affluent
of the Rio Madeira, not far south of the town of Nova Olinda do
Norte (INPA 4085). Photographs by M. G. M. van Roosmalen.


Collection of the National Institute for Amazon Research
(INPA) in Manaus, Amazonas, Brazil, under INPA 4085
(Table 2).

The species is parapatric with C. '..rr . along the east
bank of the Rio Canuma in the interfluve delineated by the
lower Rio Madeira and Rio Canuma, with C. baptista along
the north bank of the Parana do Uraria, at the northern
tip of its range, with C. bernhardi (a new species to be
described below) along the west bank of the lower Rio
Aripuand in the interfluve delineated by the Rios Aripuand
and Roosevelt (its left bank tributary) and Rio Sucundurf,
most likely including the interfluve between the Rios Acarf
and Sucundurf, with C. *'..rr . or a new, still to be
described species of titi along the east bank of the Rio
Sucundurf (east as far as the Rio Juruena) (Figs. 1 and 2).

Diagnostic characters: Forehead, crown, sides of body,
chest, belly, limbs, and tail grayish to blackish agouti,
all contrasting with tawny or reddish brown agouti mid-
dorsum; upper surface of cheiridia blackish mixed with
gray (hair tips grayish); tail predominantly blackish, mixed
with gray, proximal one third mixed with tawny agouti like
the outer surface of legs; arms blackish, the hairs grayish-
tipped as in crown; hairs of dorsum and sides of body with
4 bands, a 2 cm wide blackish tawny proximal one, a 1 cm
wide tawny band, a 1 cm wide black band, and a 0.3 cm
wide, tawny agouti distal tip.

Distinguished from all other titi species by grayish agouti
forehead, crown, sides of body, chest, belly, and limbs;
sideburns and throat usually grayish to grayish agouti.
(Fig. 19).



Callicebus brunneus (Wagner, 1842)











Lector pe -.tIUlr i-, ile, skin and skull, no. 3454,
Nun 1'1ir..l-1 1[.l.'. I museum, Wien, Austria,
collected by Johann Natterer, September, 1829.

Type locality: Cachoeira da Bananeira, Rio
Guapord, upper Rio Madeira, state of Rondonia,
Brazil.

Geographic distribution: Right bank of upper
Rio Madeira, in the states of Rondonia and Acre,
Brazil. In Rondonia, the species is parapatric
with C. bernhardi along the entire Rio Ji-Parand;





Neotropical Primates lO(Suppl.), June 2002


in the north of its distribution, it is parapatric with
C. dubius along the west bank of the Rio Madeira; in the
west of its distribution, it is parapatric with C. cupreus
along the upper Rio Purds; in the south of its distribution,
it is parapatric with C. modestus in the interfluve of the
Rfos Beni and Madre de Dios, and with C. donacophilus in
the upper Rio Mamord and San Miguel basins, Bolivia
(Figs. 1 and 2).

Diagnostic characters: Darkest species of the moloch
group, the forehead, forearms, legs, cheiridia, and base
of the tail blackish to dark reddish-brown, but rest of
tail contrasted pale or dominantly buffy mixed with
blackish; sideburns blackish to dark reddish-brown; upper
parts and sides of body brownish or red-brown agouti,
underparts brownish or reddish, not sharply defined from
sides of body.

Distinguished from C. bernhardi by generally blackish
or dark brown forehead, lack of white ear tufts, blackish-
or reddish-brown instead of contrasted bright orange
sideburns and under parts, and brownish or blackish
not whitish cheiridia; from C. cinerascens by its generally
blackish or dark brown instead of grayish appearance,
blackish or reddish-brown instead of grayish sideburns,
and tail dominantly buffy intermixed with black (Figs. 20
and 21).


Figure 20. The brown titi monkey, C.,'/,,,. brunneus
(Wagner, 1842). Juvenile (left) and adult photographed in the
National Zoo, Washington, DC, August 1988, by M. G. M. van
Roosmalen.


Figure 21. The brown titi monkey, (.i' ,n, brunneus (Wagner,
1842). Top. Adult pair with the tails entwined typical for titis.
Bottom. Adult photographed in the National Zoo, Washington,
DC, August 1988, by M. G. M. van Roosmalen.





Neotropical Primates 10(Suppl.), June 2002


Callicebus hoffmannsi Thomas, 1908
n .. .. .
.........



I ,.





Holotype: Adult male, skin and skull,
British Museum of Natural History no.
1908.5.9.11, collected by W. Hoffmanns
February 1906. ..

Type locality: Urucurituba, Rio Tapaj6s, state of
Pard, Brazil.

Geographic distribution: Central Amazonia,
Brazil, south of the Rio Amazonas in the
states of Amazonas and Pard, from right bank
of Rio Canuma, where it is parapatric with
C. cinerascens, to left bank Rio Tapaj6s, where it
is parapatric with C. moloch, south to the north
(right) bank of Rio Sucundurf, where it is parapatric with
an as yet undescribed species of titi, and north along
the south bank of the Parand do Urarid and Parand do
Ramos, east along the left bank of the Rio Andird and
the right bank of the Rio Ufra-Curupi south of the town
of Parintins where it is parapatric with C. baptista (in
the lower Rios Andir/UirA-Curupi interfluve and north
of the Parand do Urarid and Parand do Ramos) (Figs. 1
and 2).

Diagnostic characters: Basically a two-colored (grayish
and yellowish-white) titi. Upper and outer surface of
head, trunk, and limbs grayish agouti, sometimes light A
gray to almost white; forehead grayish as crown, with
or without a black or blackish coronal band not as far as
the grayish ears; lacking white ear tufts; sideburns, under
parts of body, inner side of limbs sharply contrasted
yellowish to white; mid-dorsum olivaceous-grayish;
tail blackish agouti to black; no whitish cheiridia nor
tail-tip.

Distinguished from C. moloch by upper surface of hands ....
and feet blackish agouti and black tail; from C. baptista by
pale yellowish, not bright reddish or mahogany, sideburns
and under parts, and black tail; from C. cinerascens by
chest, belly, and inner side of limbs uniformly buffy
(yellowish-white); from C. bernhardi by lacking white
ear tufts, white cheiridia and white tip of the tail and
having yellowish-white instead of bright orange sideburns,
beard, under parts of body, and inner side of limbs (Figs.
22 and 23).
Figure 22. Hoffmann's titi monkey, (.du .
Thomas, 1908. Top. Photographs by R. A. Mittermeier in the
Belkm Primate Center, Belem, Pard.





Neotropical Primates O1(Suppl.), June 2002


Callicebus baptista Linnberg, 1939














Holorype N..,, p-..ihed; 17 syntypes in the
i ..i IN N irtL, Hir... Museum of Stockholm,
.. ..ll,,.r,..I 1, 936 by A. M Olalla
1 d..._ i ..i.. i i['.Irr i and Lago do Tapaiuna.
,. i....- ,r '" 1 ..i., -.nated an adult male, skin
r..' .-..ull ,r.. .! 1. From Lago do Baptista, as
t rl,, I,,.r, ,r, p

Type localatv. N.,r p, cified, but restricted to
i ,...... i. q r I, H, ,hkovitz (1963). The lake
1, l.. iN .. i ,- rl, t rl, Parand do Urarid, south
..I th(. Ri.' A . n ..n.i, .md east of the town of
Novo Olinda do Norte on the right bank of the
Rio Madeira.

Geographic distribution: Central Amazonian Brazil,
south of the Rio Amazonas and east of the Rio Madeira
in the state of Amazonas east almost as far as the western
limit of the state of Para, and north of the Parana do
Canuma, Parana do Uraria, and Parana do Ramos. It
was observed in the wild by the first author on the west
(left) bank of the Rio Ufra-Curupa, and is believed
to have crossed over the Parana do Ramos west of the


Figure 23. Hoffmann's titi monkey, (.'/1,,,,, ...
Thomas, 1908. Photographs by Gary Comer.


Figure 24. The distributions of C./, ,,w:, baptista and (.u/ ,,:
... 'between the lower Rios Madeira and Tapaj6s in the
central Brazilian Amazon. Map by Stephen D. Nash.





Neotropical Primates 10(Suppl.), June 2002


town of Parintins, forming an enclave population in
the interfluve delineated by the lower Rio Ufra-Curupa
and lower Rio Andira. M. G. M. van Roosmalen also
observed populations of entirely pale yellowish to
almost white color morphs of ( .... '... .
along the Rio Mamurd, one river further to the east,
and classic yellowish-white and gray ( .... ,'... .
on both banks of the middle and upper Rio Andira
(Fig. 24). These observations confirm the parapatry of
( .... '...r and C. baptista, and therefore they
are elevated to full species here, whereas Hershkovitz
(1990) considers them subspecies of C .... hoffmannsi
(Figs. 1, 2, and 24).

Diagnostic characters: Sideburns, under parts, and inner
side of limbs bright to dark reddish, or reddish brown
(saturate pheomelanin); upper and outer surface of head,
trunk, and limbs grayish to blackish agouti; forehead like
crown, whitish ear tufts lacking; tail dominantly blackish


Figure 25. The Baptista Lake titi monkey, CA.'//, baptista
L6nnberg, 1939. A skin obtained from the left bank of the Rio
Ufra-Curapi, a right bank affluent of the Rio Amazonas: a. whole
skin, b. crown and nape, c. tail on the background of the dorsum.
In the private collection of M. G. M. van Roosmalen (MGMR
50). Photographs by M. G. M. van Roosmalen.


agouti to entirely blackish, often intermixed with buff and
gray hairs.

Distinguished from C. cinerascens and C. .'...rr
by uniformly reddish or reddish brown sideburns, and
underparts and inner surface of limbs; from C. bernhardi
by lack of white ear tufts, white cheiridia and white tail tip;
from C. moloch by dark brownish or grayish agouti upper
and outer parts of trunk, limbs, crown and forehead, reddish
or reddish brown instead of bright orange sideburns, and
lacking the buffy upper surface of the cheiridia and buffy
pencilled tip of tail (Fig. 25).



Callicebus bernhardi new species











Holotype: Complete adult skeleton and skull
of unknown sex, found on the forest floor. The
specimen apparently died from natural causes
and was collected by M. G. M. van Roosmalen
in November 1998. It was deposited as INPA
no. 3929 in the Mammal Collection of the
National Institute for Amazon Research (INPA),
Manaus, Amazonas, Brazil.

Paratypes: In 1996, two juvenile males were
obtained alive along the Rio MariepauA at
Santa Cruz, not far from its confluence with the
Rio Madeira, and were kept in M. G. M. van
Roosmalen's Breeding Center for Endangered
Amazonian Monkeys in Manaus, Brazil. They died in
April and August 2001 and are deposited as INPA no.
4029 and INPA no. 4033, respectively, in the Mammal
Collection of the National Institute for Amazon Research
(INPA), Manaus, Amazonas, Brazil.

Type locality: West bank of the lower Rio Aripuana, at the
edge of the settlement of Nova Olinda, 41 km southwest
of the town of Novo Aripuana, Amazonas state, Brazil.
This region is located in south-central Amazonia, south of
Rio Amazonas and east of Rio Madeira. Coordinates for
the type locality are: 0530'63"S, 6024'61"W. Altitude
45 m. (Fig. 26).

Geographic distribution: Interfluve delineated by the
Rios Madeira-Jf-Parana and Rios Aripuana-Roosevelt,
in the states of Amazonas and Rondonia, Brazil. In
Rondonia, the species is parapatric in the west with
C. brunneus along the entire Rio Jf-Parana, and in the east





Neotropical Primates 10(Suppl.), June 2002


Figure 26. Collecting localities for (.i'l,,,., bernhardi new
species.



with C. cinerascens along the Rio Roosevelt; in Amazonas,
the species is parapatric with C. dubius in the west along
the middle Rio Madeira, and with C. cinerascens in the
east along the Rio Aripuand (Figs. 1 and 2). Ferrari et al.
(1996) observed a grey titi monkey at Pimenta Bueno in
Rondonia, on the west bank of the Rio Ji-Parana. They
noted that it was not the distinctively brown-colored
C. brunneus and were unable to identify it. It may have
been C. bernhardi, which would extend its range a little
to the west across the upper the Rio Ji-Parana, but this
requires confirmation.

This species has been observed in the wild by M.G.M. van
Roosmalen at the following localities: west bank of lower
Rio Aripuana, Nova Olinda, Amazonas state, 0530'01"S,
6024'27.4"W; west bank of lower Rio Aripuana, Monte
Alegre, Reserva Florestal Getal, 0534'68"S, 6023'40"W;
west bank of lower Rio Aripuana, Novo Oriente
(Capimtuba), 0543'41"S, 6017'09"W; east bank of
middle Rio Madeira, seringal Sao Luis, ca. 5 km south of
the town of Manicord, 0550'28"S, 6118'19"W, altitude
45 m.

Diagnostic characters: Upper and outer surface of head,
trunk, and limbs grayish black, on the back mixed with
brownish agouti or red-brown; forehead not defined
from crown, grayish black to gray; ears black with
conspicuous whitish tufts; sideburns, under parts of body,
and inner side of limbs sharply contrasted dark orange;
cheiridia sharply contrasted white against grayish black
lower limbs; tail black except for a sharply contrasted
white pencil.

Distinguished from C. cinerascens by uniformly dark
orange instead of grayish sideburns and inner sides of


Figure 27. Prince Bernhard's titi monkey, (.,,,',:, bernhardi
new species. The adult male paratype INPA 4033 from the Rio
Mariepaud, affluent of the Rio Madeira, in 1998. Photographs by
M. G. M. van Roosmalen.


limbs, chest, and belly, and tail with white pencil; from
C. brunneus by grayish instead of dark brown to black
forehead and crown, and dark orange sideburns, inner
sides of limbs, chest, and belly; from C. .'..rr . and
C. baptista by strikingly contrasting white ear tufts,
cheiridia and tip of the tail (pencil); from C. moloch
by grayish forehead and crown, white ear tufts, and
blackish tail with a distinct white pencil; from C. dubius
by lack of black vibrissae and white blaze (Figs. 27,
30-34).

External characters of holotype: Forehead, crown, sides of
body, and outer sides of limbs grayish; rump, mid-dorsum,





Neotropical Primates 10O(Suppl.), June 2002


back, and nape grayish mixed with brownish agouti to
reddish brown, the hairs 5 cm long, with 5 blackish bands
alternating with 4 narrow brownish agouti to red-brown
ones, the most proximal (2 cm long) and the distal one (tip)
black; face black with some white hairs around mouth and
nostrils; ears black with white tufts contrasted with light


Figure 28. Skull and mandible of the adult (unknown sex)
holotype of C.d'/,,,., bernhardi (INPA 3929). Photographs by
M. G. M. van Roosmalen.


grayish forehead and crown; tail ca. 55 cm long, the distal
7 cm forming a white pencil (Table 2).

Cranial measurements: See Table 3 and Figures 28
and 29.

Habitat: Titi monkeys of the cupreus and moloch Group
essentially are secondary and disturbed forest specialists.
The various groups of C. bernhardi observed in the wild
by M. G. M. van Roosmalen were invariably in naturally
or anthropogenically disturbed forest, such as blow-
downs (large patches of secondary forest after massive
deforestation caused by dramatic rainstorm events),
liana forest on abandoned terras pretas, secondary forest
surrounding plantations and fields, and so-called seringais,
an early form of agroforestry, in which areas of high
riverbank forests along Amazonian white-water rivers were
turned into productive multi-species forests. These forests
are rich in a number of commercially valuable and edible
fruit-producing native trees, including rubber (Hevea
brasiliensis), Brazil nut (Bertholletia excelsa), wild cocoa
(Theobroma spp.), 'bacurf' (Rheedia spp., Platonia insignis),
'ingi' (Inga spp.), 'taperebi' (Spondias mombin), 'biribi'
(Rollinia squamosa), soursopp' (Annona spp.), and a number
of tree species belonging to the sapotilla family (Sapotaceae)
and palms (Palmae or Arecaceae). These seringais seem to



Table 3. Cranial and dental measurements of the holotype
Callicebus bernhardi (INPA 3929) (mm) (measurements taken
by R. Voss).
Cranial measurements (mm)
Greatest skull length 64
Condylobasal length 53
Zygomatic breadth 40
Biorbital breadth 36
Postorbital constriction 31
Brain-case length 52
Brain-case width: 34
Greatest skull length 64
Condylobasal length 53

Dental measurements (mm
Outside crown-to-crown dimensions
I -M3 23.19
C -M3 18.06
PM2 M_ 3 15.62
M -M3 9.76
12- 2 10.08
C1- C1 14.47
M1-M1 21.31
M-- M3 20.49


Mandible measurements (mm)
Mandible length 42.76
Mandible height 34.44





Neotropical Primates 10(Suppl.), June 2002


offer optimal habitat and a year-round food supply for titi
monkeys, marmosets, tamarins, and night monkeys. The
highest densities of titi monkeys of the cupreus and moloch
Group can be found in these seringais, if the local people do
not hunt them. Often before dawn in the early morning,
all pairs living in the area can be heard performing duet
calls and, therefore, local densities of titis can be easily
estimated.

Origin of the name: This titi monkey is named in honor
of His Royal Highness Prince Bernhard of the Netherlands,
who for half a century has been a global leader in nature


Figure 29. Skull and mandible of the adult male paratype of
(C.///i,,; bernhardi (INPA 4029). Photographs by M. G. M.
van Roosmalen.


conservation. In particular, the authors are grateful to him
for having created the Order of the Golden Ark, a highly
prestigious award, equivalent to knighthood, which is
presented every year to a select group of conservationists
from around the world. Two of the authors (RAM in 1995;
MGMR in 1997) have been thus honored and are naming
this new titi as a small token of their appreciation.

Vernacular name: This titi monkey is referred to as zog-zog
or zogue-zogue by the local people. For an English name, we
suggest Prince Bernhard's titi monkey.


Figure 30. Dorsal views of the adult male paratype of Ci.'/i ,,,;,
bernhardi (INPA 4029): a. crown, forearms and hands, b. close
up of hairs of the nape, and c. close up of the hairs of the mid-
dorsum. Photographs by M. G. M. van Roosmalen.





Neotropical Primates 10(Suppl.), June 2002


c












d



























Figure 31. Details of the adult male paratype of C.'/,/,,,
bernhardi (INPA 4033): a. dorsal surface of feet, b. inner side of
right arm and tail tip, c. dorsal surface of left hand, d. close-up of
dorsal pelage, e. crown, ear, and part of sideburn. Photographs by
M. G. M. van Roosmalen.


Figure 32. Prince Bernhard's titi monkey, C.'// ,,,, bernhardi
new species. Views of the adult male paratype (INPA 4029).
Photographs by Gary Comer.





Neotropical Primates 10(Suppl.), June 2002


Figure 33. Prince Bernhard's titi monkey, C. //,, bernhardi new species. Views of the adult male paratype (INPA 4029). In the top left
photograph it is with a Saterd marmoset, Mico saterei (Sousa e Silva & Noronha, 1998). Photographs by M. G. M. van Roosmalen.




Neotropical Primates 10(Suppl.), June 2002


e1


Figure 34. Prince Bernhard's titi monkey, C.ii// bernhardi new species. Views of the adult male paratype (INPA 4029). Photographs
by M. G. M. van Roosmalen.





Neotropical Primates 10O(Suppl.), June 2002

IV. C. TORQUATUS GROUP


Callicebus torquatus (Hoffmannsegg, 1807)


Syntypes: One, perhaps the holotype, mounted
in the Museum fur Naturkunde der Humboldt-
Universitat, Berlin, Germany; another syntype,
no. 687(522), in the Museum National d'Histoire
Naturelle, Paris, collected before 1806 by FW.
Sieber, donated in 1808 to the Lisbon Museum,
Portugal, by Count von Hoffmannsegg, and later
removed to Paris, France.

Type locality: Interior of state of Pard (before
1850 including the state of Amazonas), Brazil, re-
determined by Hershkovitz (1963) as Codajas, a
town on the north bank of the Rio Solim6es, state
of Amazonas.


Geographic distribution: State of Amazonas, Brazil; left
(north) bank of Rio Solim6es/Rio Japura, west as far as
Rio Apaporis/upper Rio Vaupes. The range in the north is
delineated by the Rio Negro/Rio Uaupes, east at least as far
as the town of Codajis, maybe even as far as the town of
Manacapurd, both on the north bank of the Rio Solimoes,
west of Manaus (Figs. 35 and 37).

Diagnostic characters: Forehead, crown, sideburns, back,
trunk and outer surface of limbs uniformly dark reddish
to blackish brown; underparts, chest and belly reddish or
reddish brown; throat collar weakly defined, buffy, not
extending to ear base; hands and feet whitish or buffy; tail
blackish mixed with reddish.

Distinguished from C. purinus by blackish forehead and
dark reddish crown not sharply demarcated from mahogany
nape, hairs of back uniformly colored or faintly banded,
and white cheiridia; from C. regulus by reddish brown
instead of brown to blackish underparts, back of crown not
markedly differentiated from forehead and nape, and white
cheiridia; from C. lucifer and C. lugens by overall reddish
brown or mahogany coloration and white hands and feet;
from C. medemi by buffy or white instead of black cheiridia
(Fig. 36).


Figure 35. Distributions of the Amazonian titi monkeys, genus Callicebus, belonging to the torquatus Group. Map by Stephen D. Nash.


. 1 4 1
NZWr





























Figure 36. The collared titi monkey, C./i,t : torquatus
(Hoffmannsegg, 1807). A pet seen in 1973 in Tefe, Amazonas,
Brazil. Photographs by R. A. Mittermeier.


Neotropical Primates 10(Suppl.), June 2002

Callicebus lugens (Humboldt, 1811)










Holotype: None; the name was based on a captive
animal observed by A. Humboldt in 1811 during
his journey on the upper Rio Orinoco, Amazonas
state, Venezuela.

Type locality: In the vicinity of San Fernando de
Atabapo, at the confluence of the Rios Orinoco
and Guaviare, Amazonas state, Venezuela.

Geographic distribution: Eastern Colombia,
departments of Vichada, Meta east of the Rio
Ariari, GuainiA, Guaviare, Vaupes, and Caqueta


Titi Monkeys
genus Callicebus


Schematic Map 4 the stbuioun
o the tuonrau Group


: WTM' tnff-s


Figure 37. Schematic map of the distribution of the Amazonian titi monkeys belonging to the torquatus Group. Illustrations by by
Stephen D. Nash.


K -"





Neotropical Primates 10(Suppl.), June 2002


east of the Rio Caguan, between the Rio Tomo in the north
and Rio Caguan-Caqueta in the south; southern Venezuela,
state of Amazonas south of the Rio Ventuari, and state of
Bolivar between the Rios Caura, Caroni, and Orinoco; and
bordering parts of northwestern Brazil, Amazonas state
north of the Rios Uaupes-Negro, and the state of Roraima
west of the Rio Branco, north as far as the foothills of
Mount Roraima (personal observations by M.G.M. van
Roosmalen) (Figs. 35 and 37).

Diagnostic characters: Feet, tail, head, sideburns, and
under parts except throat entirely blackish, hairs of back
and sides of body blackish intermixed with dark chestnut,


the hairs uniformly colored or faintly banded, hands and
throat contrasted white.

Most blackish of the C. torquatus Group, distinguished
from C. torquatus, C. purinus, and C. regulus by blackish
chest and belly, and white hands combined with black
feet; from C. lucifer by lack of contrast between blackish
crown and reddish brown or blackish nape, hairs of back
uniformly colored or faintly banded, and white instead of
orange hands; from C. medemi by white instead of blackish
hands (Fig. 38).


Callicebus lucifer Thomas, 1914


Figure 38. Cu./i/ lugens (Humboldt, 1811). An adult female
from the interfluve of the Rio Caqueti (left bank) and the Rio
Apaporis (right bank), west of the main portage between the two
rivers and downriver from La Pedrera, near the border between
Colombia and Brazil. Photographed at the Capart Biological
Station, Rio Apaporis, by R. A. Mittermeier in August, 1995.


9 x-ey


Holotype: Adult male, skin and skull, in the British
Museum of Natural History, London, U.K., no.
14.3.1.2, collected August 1913 by J.J. Mounsey.

Type locality: Yahauas Territory, in the vicinity of
Pebas, Department of Loreto, Peru.

Geographic distribution: Interfluve delineated
by the Rio Solim6es and Rio Napo in the south,
and the Rio Japura and Rio Caqueta in the
north; in Brazil between the Rios Solim6es and
Japuri; in Colombia between the Rios Caqueta
below mouth of Rio Caguan, and Rios Putumayo
and Amazonas in the departments of Caqueti,
Putumayo and Amazonas; in Ecuador between the upper
Rios Aguarico and Putumayo, Napo province; and in Peru
in northern Loreto, between the Rios Putumayo, Nanay,
and Amazonas. Campos et al. (1992) and De la Torre et
al. (1995) report on the presence of C. torquatus, which
we presume to be C. lucifer (but may, alternatively, be C.
medemi) in the Cuyabeno Reserve, Rio Aguarico, province
of Sucumbios, in northeastern Ecuador (Figs. 35 and 37).

Diagnostic characters: Feet, tail, head, sideburns, and
under parts except throat entirely blackish, hairs of back
and sides of body brownish or reddish brown, the hairs
distinctly to weakly banded, throat white, hands orange.

Distinguished from C. lugens by brownish agouti upper
parts and orange instead of white hands; from C. torquatus
and C. purinus by orange instead of white hands and
blackish under parts; from C. regulus by blackish (except
white throat) head and hairs surrounding ears uniformly
blackish, and orange hands; from C. medemiby dominantly
or entirely orange instead of black hands.


IP-.,;
r *"^ih '"k





Neotropical Primates 10O(Suppl.), June 2002


Callicebus purinus Thomas, 1927


Holotype: Adult male, skin
British Museum of Natural
U.K., no. 26.5.521, collected
Ehrhardt.


and skull, in the
History, London,
May 1925 by W.


Type locality: Lago AyapuA, left bank of lower Rio
Purds, state of Amazonas, Brazil.

Geographic distribution: In the state of
Amazonas, Brazil, south of the Rio Solim6es
between the Rios Purds and Jurua. It extends
south as far as the Rio Tapaua or even the Rio
Pauinf if the species reported to occur between the
Rios Tapaua and Pauinf, left bank of tributaries
of the Rio Purds, does not represent a new form (Figs.
35 and 37).


Figure 39. (.di/ ,n .* Thomas, 1927. An adult female, a pet
monkey photographed at the seringal/castanhal Camarua, at the
north bank near the mouth of the Rio Tapaud with the Rio Purts,
August 2, 2001. Photographs by M. G. M. van Roosmalen.


Figure 40. (.d/i/,,." purinus Thomas, 1927. An adolescent
male from seringal/castanhal Camarua, near the mouth of the
Rio Tapaud with the Rio Purts, August, 2001. Photographs by
M. G. M. van Roosmalen.





Neotropical Primates 10(Suppl.), June 2002


Diagnostic characters: Hands whitish, lower arms
and feet black, hairs of back and sides dark red-brown,
strongly to faintly banded, tail blackish with mixture of
reddish, under parts except throat dark reddish brown
or reddish, throat collar contrastingly colored buffy,
yellowish, or whitish, the collar well developed and
extending to ear base, sideburns, forehead (blaze) and ears
black, sharply contrasting with white whiskers and bright
red crown.

Distinguished from C. torquatus by bright reddish crown
sharply contrasting with black forehead and sideburns,
marked strongly to faintly banded agouti pattern of
back, throat collar more developed and sharply defined
from surrounding parts, and black instead of white
or buffy feet; from C. regulus, C. lugens and C. lucifer
by reddish brown instead of blackish under-parts
(chest and belly) and by more reddish coloration
throughout, the crown always sharply defined from
nape; and from C. medemi by more reddish coloration
throughout, and white or yellowish instead of black hands
(Figs. 39 and 40).


more or less banded, and orange instead of white hands;
from C. torquatus by inner side of arms entirely blackish,
more developed white throat collar, a strongly contrasting
reddish crown, and blackish instead of white feet; from
C. luciferbyindistinctlybandedhairs ofbackandsides ofbody,
and a contrasting reddish instead of blackish crown; from
C. lugens by paler back, sides of body, and under-parts, and
a strongly contrasting reddish crown; and from C. medemi
by orange instead of blackish hands, and contrasting
reddish instead of blackish crown.



Callicebus medemi Hershkovitz, 1963


0


Callicebus regulus Thomas, 1927


,1 h*m.


Holotype: Adult female, skin and skull, in the
British Museum of Natural History, London,
U.K., no. 27.3.6.8, collected August 1926 by
W. Ehrhardt.

Type locality: Fonte Boa, right bank of upper
Rio Solim6es, state of Amazonas, Brazil.

Geographic distribution: Brazil, state of
Amazonas, between the upper Rio Solim6es, the
lower Rio Javarf, and the left (west) bank of the
Rio JuruA from mouth at the Rio Solim6es to
about 70 S (Figs. 35 and 37).

Diagnostic characters: Hands orange, tail blackish, inner
side of arms entirely blackish, under parts (chest and
belly) except throat brown or blackish, hairs above and
behind ears more or less banded, sideburns brownish,
crown strongly contrasted reddish, and throat collar well
developed, white.

Distinguished from C. purinus by dark brown chest and
belly, brownish sideburns, hairs above and behind ears


Holotype: Adult female, skin and skull, in the Field
Museum of Natural History, Chicago, no. 70699,
collected March, 1952 by P. Hershkovitz.

Type locality: Rio Mecaya, near mouth, at right
bank of Rio Caqueta, Putumayo, Colombia.

Geographic distribution: The Colombian
Amazon between the Rios Caqueta and Putumayo
in the Intendencia del Putumayo and the southern
part of the Intendencia de Caqueta (Figs. 35 and
37).

Diagnostic characters: Head, sideburns, hands,
feet, tail, and under parts except throat entirely or
predominantly blackish, throat white.

Distinguished from C. torquatus, C. regulus, and
C. purinus by much darker (blackish) coloration
throughout, including the blackish hands, legs, and under
parts; from C. lucifer and C. lugens by upper surface of
hands uniformly or dominantly blackish instead of orange
and white, respectively.











Undescribed form of the
Callicebus torquatus group


Neotropical Primates 10(Suppl.), June 2002

V. C. PERSONATUS GROUP


Callicebus personatus (E. Geoffroy, 1812)


Lv


Figure 41. This individual was being kept in captivity at Boca
Manu, Rio Manu, Peru, in 1983 and probably represents an
undescribed form of the C.'/,,,,. torquatus group. Photographs
by A. Young and R. A. Mittermeier.


Holotype: Mounted 1..Ir ,iI......'.
originally in the Ajuda Museum, Lisbon,
Portugal, but seized by the French in
1808 during the Napoleonic invasion
and transferred to the Musdum National
d'Histoire Naturelle, Paris, France, from where it
disappeared around 1820.

Type locality: Restricted by Hershkovitz (1990) to
the lower Rio Doce, Espirito Santo, Brazil.

Geographic distribution: In the Atlantic forest of
southeastern Brazil, in the state of Espirito Santo, south
from the region of the lower Rio Itadnas (Kinzey, 1982;
Oliver and Santos, 1991). The Rio Mucurf, to the north
of the Rio Itadnas was marked as the limit by Hershkovitz
(1990), but Oliver and Santos (1991) reported that
C. melanochir may occur south of the lower Itadnas as far
as Barra Nova (18054'S, 39047'W). Oliver and Santos
(1991) indicated that the region of the Rios Itadnas and
Mucurf may be a zone of intergradation between the
personatus (to the south) and melanochir (to the north).
C. personatus occurs further inland into northwestern
Minas Gerais, east at least as far as Te6filo Otoni (Kinzey,
1982; Hershkovitz, 1990) and the east (right) bank of the
Rio Jequitinhonha (Rylands et al. 1988). South of the
Rios Mucurf/Itadnas, they occur throughout the state of
Espirito Santo and in the north of Rio de Janeiro (M. C. M.
Kierulff in Rylands, 1988), although now extremely scarce
(Oliver and Santos, 1991). C. personatus extends west along
the Rio Doce valley into Minas Gerais as far as the Serra
da Mantiqueira (Serra do Brigadeiro) (Cosenza, 1993),
and at least as far south as Juiz de Fora, Minas Gerais. It
remains unclear if C. personatus, or another species, occurs
northwest of the Rio Jequitinhonha. Hershkovitz (in litt. to
A. B. Rylands, January 1988) listed Buen6polis, near the




Neotropical Primates 10(Suppl.), June 2002


Titi Monkeys
genus Callicebus
Schematic Map of the Distribution
of the eastern Brazillan txa






If


wk- .


6Ar'nGMi I


ATLAAMf OCFAM


Figure 42. Schematic map of the distribution of the southern Brazilian titi monkeys of the personatus Group, genus
Callicebus.





Neotropical Primates 10(Suppl.), June 2002


Figure 43. (.r/, A,:, personatus E. Geoffroyi, 1812, from Minas
Gerais, Brazil. Photographs by R. A. Mittermeier.


Serra do Cabral (17054'S, 4411'W), northwestern Minas
Gerais, as a locality for C. personatus, but it was not included
as a locality in his publication in 1990 (Figs. 42 and 44).

Diagnostic characters: Throat, sideburns, forehead
and crown blackish to plane of ears, rest of body
sharply defined uniformly buffy to orange like nape except
blackish hands, lower arms and feet. Two color variations,
body and tail uniformly buffy or orange (Figs. 3 and 37).

Distinguished from C. nigrifrons, C. melanochir, and
C. barbarabrownae by blackish forehead, crown to line of
ears, cheeks, ear tufts, and throat, the hairs not banded; and
back of crown sharply contrasting orange; from C. coimbrai
by black instead of buffy cheeks, sideburns, chest, back of
head, and nape (Fig. 43).


Callicebus melanochirWied-Neuwied, 1820











Holorvpe .l]ulr t> i [de, mounted without skull,
S .. /.... I.. .I-, staatssammlung, Mtinchen,
., ,- .i i r. .,. rwo in Rijksmuseum van
N rNui.I iI..l H,r. ... Leiden, Holland, of which
in. I -, .' .... i-] ., -,i r d lectotype by Hershkovitz
Sl'i', ... i"i ,.u'I um fur Naturkunde der
-J. Huil.. I.Ir- 1 I. -....ir u Berlin, Germany, one in
\ 1 uni.'.,, N u'..i'il ..I Histoire Naturelle, Paris,
i i, J.... I.. ...S 'kull no. A2.815, and one in
S I'r., -ir-.i ir. .- Wied-Neuwied Museum
,'. I-.l. ..i.... "..r , .r mymore), all mounted with
skull in skin, collected April 1816.

Type locality: Morro d'Arara or Fazenda Arara,
state of Bahia, Brazil.

Geographic distribution: Hershkovitz (1990) gives the
range as the Atlantic coastal forest of eastern Brazil, north
from the Rio Mucurf in the state of Espfrito Santo to the
Rio Paraguaqi in Bahia. As discussed for C. personatus, it
would seem that the southern limit is not clearcut, and is
possibly marked by a zone of intergradation in the valleys
of the Rios Itatnas and Mucurf in northern Espfrito Santo.
To the north, C. melanochir extends as far as the Rio
Paraguagi, where it meets the range of C. barbarabrownae
(Hershkovitz, 1990; Oliver and Santos, 1991; Flesher,
1999). Inland it would appear that its range is limited by
inhospitable liana forests and dry forests of the interior of
the state of Bahia. South of the Rio Jequitinhonha, it is
restricted to coastal forest, being replaced by C. personatus
further inland (Figs. 42 and 44).

Diagnostic characters: Forehead, crown, and throat
dominantly grayish agouti, buffy or pale brownish agouti,
the hairs finely banded, cheiridia and facial fringe blackish,
overall the least colorful member of the personatus Group.

Distinguished from C. nigrifrons and C. personatus by
the forehead not being sharply defined blackish, entire
crown blackish agouti or grayish agouti like nape, and
sides of neck and throat grayish or blackish agouti; from
C. barbarabrownae by overall much darker coloration; from
C. coimbrai by grayish agouti, buffy or pale brownish agouti
forehead, crown, and throat.





Neotropical Primates 10O(Suppl.), June 2002

Callicebus nigrifrons Spix, 1823


Lector pe A.tulr unknown, mounted skin
only, no. 88, Zoologische Staatssammlung,
Miinchen, Germany, collected 1817 by the Spix
and Martius expedition.

Type locality: Interpreted by Hershkovitz (1990)
to be the Rio Ongas, municipality of Campos, state
of Rio de Janeiro, Brazil.

Geographic distribution: Southeastern Brazil, in
the states of Rio de Janeiro, Sao Paulo north from
the Rio Tiete, east of the Rio Parana but restricted
to the right bank of the Rio Paranafba in western
Minas Gerais. It occurs on both sides of the uppermost
reaches of the Rio Sao Francisco, extending to the east as
far as the Serra da Mantiqueira and Serra do Espinhago
in Minas Gerais, where it meets the range of C. personatus
(Figs. 42 and 44). As in all of the Atlantic forest titi
monkeys, although widespread, the extreme fragmentation
and urbanization of the forests within its range means that
today populations are isolated and generally very small, and
in many places they are locally or regionally extirpated even
where forests patches remain.


/-
f P &



Spa, lo -0i aM









Figure 44. Distributions of the Atlantic forest titi monkeys,
genus Callicebus, belonging to the personatus Group. Map by
Stephen D. Nash.


Figure 45. Top. The southern masked titi, (.I'/ tt,,:, nigrifrons
(Spix, 1823). An individual from the south-east of the state of
Minas Gerais, Brazil, 1979. Bottom. A titi monkey from Guapi-
Mirim, Sao Paulo, Brazil. This is within the recognized distribu-
tion of (.nl ,;,. nigrifrons, but this individual had a darker mask
and mantle and a more orange, rather than rusty-brown, tail.
Photographs by R. A. Mittermeier.


I





Neotropical Primates 10O(Suppl.), June 2002


Diagnostic characters: Forehead and crown blackish to
about halfway plane of ears, rest of crown grading into
coarsely banded brownish agouti or orange-brown of nape;
throat pale brownish agouti like chest; cheiridia and ears
black; tail orange.

Distinguished from C. melanochir and C. barbarabrownae
by blackish forehead, and anterior portion of crown blackish
thinly mixed with buff-banded hairs; from C. personatus by
blackish front of crown grading into agouti of nape without
line of demarcation, and throat pale brownish agouti like
chest; from C. coimbrai by pale brownish agouti sideburns,
throat, and chest (Fig. 45).



Callicebus barbarabrownae Hershkovitz, 1990


r


"A .f


Holotype: Skin and skull, no. 3.9.5.7, British
Museum of Natural History, London, U.K.,
collected June 1903 by Alphonse Robert.

Type locality: Lamarao, Bahia, Brazil, altitude
about 300 m above sea level.

Geographic distribution: According to
Hershkovitz (1990), the coastal highlands of the
north-central part of the state of Bahia, Brazil,
between the Rios Paraguaid, just north of the
city of Salvador, and Itapicurd. The western
limits to its range are unknown but probably,
at least historically, the middle reaches of the
Rio Sao Francisco. Ricardo B. Machado and the late
A. Brandt (pers. comm. 1988) recorded a population of
( .... at the Serra da Quixaba in the municipalities of
Canudos and Monte Santo, north of the Rio Vaza-Barris,
northern Bahia (39o20'W, 1015'S). Ilmar B. Santos also
recorded ( .... at Jeremoabo, a little to the east, on
the northern margin of the Rio Vaza-Barris in 1990 (pers.
comm.). Marinho-Filho and Verissimo (1997) confirmed
the westernmost locality. They observed it in gallery
forest at Miror6s, municipality of Ibipeba, western Bahia
(1124'S, 4217'W). C. barbarabrownae was evidently once
widespread in forests east and south of the Rio Sao Francisco


Figure 46. The blond titi, C.Y ,i n barbarabrownae Hershkovitz,
1990. In captivity at the Federal University of Minas Gerais, Belo
Horizonte, in the mid-1980s before the species was described.
Photograph by R. A. Mittermeier.

(see Coimbra-Filho and Camara, 1996), but today survives
only in small forest enclaves in what is now predominantly
caatinga (dry thorn scrub) (Figs. 42 and 44).

Diagnostic characters: Superciliary vibrissal line black,
forehead and crown to anterior plane of ears dominantly
buffy; raised hairs of rest of crown buffy, the fine tips
blackish; sideburns, nape, and shoulders pale buff; hairs
of back and sides of body banded pheomelanin and
eumelanin; thighs and upper arms paler, forearms and legs
like back; cheiridia blackish; throat, chest, and belly nearly
entirely buffy; tail dominantly orange, base of tail yellowish;
ears and skin blackish.

Distinguished from C. melanochir by dominantly buffy
crown, sideburns, throat, trunk, and limbs with the
subterminal pheomelanic (buffy) bands of hairs paler; from
C. nigrifrons and C. personatus by forehead buffy instead of
blackish; from C. coimbrai by buffy forehead and crown.
(Fig. 46).





Neotropical Primates 10(Suppl.), June 2002

Callicebus coimbrai
Kobayashi & Langguth, 1999


* V -' ...-


Holotype: Adult female, UFPB no. 1599,
mammal collection of the Departamento de
Sistematica e Ecologia, Universidade Federal
da Parafba, Joao Pessoa, state of Parafba, Brazil,
collected January, 1994, by S. Kobayashi and A.
Langguth.

Type locality: Proximity of the small village
Aragao, in the region of Santana dos Frades
about 11 km south-west of Pacatuba, state of
Sergipe, Brazil (coordinates 10032'S, 36o 41'W),
altitude 90 m. The locality is south of the estuary
of the Rio Sao Francisco.

Geographic distribution: First described from three
localities in eastern Brazil, along the coast of the state
of Sergipe, between the Rio Sao Francisco in the north
and the Rio Real in the south (the southern border of
Sergipe). Oliver and Santos (1991) obtained reports of the
occurrence of titi monkeys in the vicinities of Umbauba,
Estancia, and AruA in coastal southern Sergipe, and
also at Cachoeira da Abadia and Jandaira in north-east
Bahia, which are probably referable to C. coimbrai. They
indicated that the Rio Sao Francisco was the northern
limit to the range of the genus in the Atlantic forest.
Sousa (2000) reported two further localities in Sergipe:
Mata do Crasto in the municipality Santa Luzia do
Itanhy, and the Mata do Dira in the municipalities of
Itaporanga and Laranjeiras. Sousa (2000) also reported
hearing vocalizations of titi monkeys in the Matas do
Conde, municipalities of Conde and Jandaira in extreme
northern Bahia in 1996. The western limits of its range are
unknown, but Ricardo B. Machado (pers. comm. 1989),
Marinho-Filho and Verfssimo (1997) recorded ( ....
in forest patches in the Caatinga, inland between Monte
Santo and UauA in the upper valley of the Rio Vaza-
Barris, and Jeremoabo and Canudos in northern Bahia.
Marinho-Filho and Verfssimo (1997) identified them as
the form barbarabrownae, although they are at about
the same latitude as coimbrai. The evidence obtained
by Kobayashi and Langguth (1999) indicated that
C. coimbrai is today restricted to the humid coastal
Atlantic forest of Sergipe, and that its southern limit is
the Rio Itapicuri in Bahia, the northern limit to the range
of C. barbarabrownae (Hershkovitz, 1990). Considering
the widespread and rapid destruction of the forests
in northern Bahia and Sergipe even in the early 16t'


Century (Coimbra-Filho and Camara, 1996), C. coimbrai
undoubtedly had a much broader range in the past (Figs.
39 and 40).

Diagnostic characters: Forehead, crown, and ears black;
trunk buffy; cheiridia blackish; tail orange; sideburns,
cheeks, back of head, and nape pale buffy; anterior half of
dorsum saddle-backed (with striped pattern).

Distinguished from all other titis of the C. personatus Group
by black forehead, crown, and ears sharply contrasting with
buffy sideburns, cheeks, back of head, nape, and trunk.


Discussion

In this review, we elevate all currently known taxa of
titi monkeys, genus Callicebus, to full species status
using the same arguments as De Vivo (1991), Van
Roosmalen et al. (1998), and Van Roosmalen et al. (2000)
in their respective reviews of Amazonian marmosets,
genus ( .-'. As in Amazonian ( .-'. (or Mico,
following Rylands et al., 2000), titi monkeys in lowland
Amazonia invariably have their distributions confined by
river barriers. These rivers may fall in the 'black-water',
'clear-water' or 'white-water' category. Both Amazonian
( .... and ( .-'. (Mico) are restricted to dry-land
(terra firme) rain forests, and only titis of the C. moloch and
C. cupreus Groups tend to seasonally venture into flooded
forest habitat along black-water and clear-water streams,
to feed on certain fruits only available there during high
water. Individuals belonging to both of these genera are
unable to swim, meaning that an accidental fall into the
water will quickly result in drowning. Passive transfer to
the other side of a river barrier through river bend cut-offs
has probably not occurred either because populations of
Amazonian titis and marmosets are not found in white-
water flooded forest (vdrzea), the only kind of habitat
that is usually subjected to cross-river transfers. Also, they
probably cannot survive on temporary igapd (black-water
flooded forest) islands long enough to be transferred to the
other side of the river.

Most of the lowland tributaries of the Amazon River are
fringed with vdrzeas and igapds several kilometers wide
and sometimes as many as 10 kilometers, which usually
stretch from the mouth to the headwaters, effectively
isolating populations of terra firme-dwelling primates such
as ( .-'. Callicebus, Saguinus, Chiropotes, Lagothrix
and Ateles. For these monkeys, the only way to colonize
adjacent interfluves is to circumvent river barriers near
their headwaters, where the floodplain ends and the river
becomes narrow enough to cross. In the vicinity of these
headwaters, there should be or have been contact zones
between populations of the taxa inhabiting the adjacent
interfluves. If interbreeding takes place between the two taxa
and broad natural hybrid zones of intergradation occur, we
should treat them as races or subspecies. However, in most
taxa of Amazonian Callicebus, ( .-'. and Saguinus,





Neotropical Primates 10O(Suppl.), June 2002


these regions seem to be narrow contact zones (and not
regions with broad clinal variation), where the colonizing
taxon successfully excludes the resident one. In such cases of
narrow zones of hybridization, the animals should continue
to be considered distinct species, following Mayr (1970),
the very same criteria used nearly 30 years ago to recognize
most Atlantic forest ( .-'. as good species (Coimbra-
Filho and Mittermeier et al., 1973).

Presently, we know of three examples of this ongoing
natural process in which one monkey species, after crossing
a geographic barrier, outcompetes another belonging to the
same 'ecospecies' (meaning they occupy the same ecological
niche). These include the following:

The central-Amazonian titi monkey, C .... baptista,
classified as a subspecies of ( .... '-...- .. by
Hershkovitz (1990), ranges south of the Rio Amazonas,
east of the lower Rio Madeira, north of the Paranas do
Canuma, Uraria, and Ramos, and east as far as the western
limit of the state of Para. However, it also occurs south of
the Parana do Ramos and west of the town of Parintins,
forming a propagule population in the small interfluve
delineated by the lower Rio Ufra-Curupa and the lower
Rio Andira amidst the much larger interfluve that is
occupied by ( .... '..rn . which stretches south
of the Paranas do Uraria and Ramos from the lower Rio
Canuma in the state of Amazonas as far as the lower Rio
Tapaj6s in the state of Para. Field surveys conducted by M.
G. M. and T. van Roosmalen revealed that no hybrid zones
of intergradation occur. It seems that ( .... baptista
managed to cross the Parana do Ramos and successfully
replace ( ...., '...r . in the interfluve delineated
by the Rios Andira and Ufra-Curupa, and the Parana do
Ramos. Their parapatry being confirmed in the field, we,
therefore, elevated the two taxa to full species status.

The second example concerns the golden-handed tamarin,
Saguinus midas, which originated in the Guianas and
largely replaced the pied bare-face tamarin, Saguinus
bicolor after possibly having colonized the entire northern
lower Amazon basin delineated by the Rio Branco in the
west, the Atlantic Ocean in the east, the Guayana Shield in
the north, and the Rio Amazonas in the south. This area
was probably formerly occupied by Saguinus bicolor before
S. midas managed to cross one of the geographic barriers
(either the Guayana Shield highlands and savannas at the
border between Brazil and the Guianas, or some river
barriers in the Brazilian state of Amapa). This natural
process of species replacement reaches its dramatic final
stage in the vicinity of Manaus, the capital of Amazonas
state, where Saguinus bicolor is headed toward extinction.
Saguinus midas is now penetrating the remaining territory
of bicolor, a small enclave on the northern shore of the Rio
Amazonas/Rio Negro and west of the Rio Trombetas. The
contact zone in this case is very narrow, and interbreeding
between the two species, which belong to the same
'ecospecies', has not been observed. Instead, since tamarins
are extremely territorial, violent encounters between


groups of both species are not uncommon (M. G. M. van
Roosmalen, pers. obs.).

The third example concerns Saguinus mystax pluto and
Saguinus mystax pileatus, classified as subspecies by
Hershkovitz (1977), with pileatus occupying the entire
Rios Jurua/Purds interfluve south of Rio Tapaua and
north of Rio Pauinf, which are both left-bank tributaries
of the Rio Purds fringed with extensive igapds along their
entire course. These rivers have their source very close to
the vdrzeas of the Rio Jurua. It is assumed that S. m. pluto
originally inhabited the entire interfluve delineated by the
Rios Jurua, Solimoes, Purds, and Tapaua, where it evolved
away from S. m. pileatus due to its separation by the strong
barrier represented by the Rio Tapaua. S. m. pileatus must
have crossed the headwaters of the Rio Tapaua somewhere
near the Rio Jurua in relatively recent geological times,
and colonized the territory of S. m. pluto from the north
and the east. Today, S. m. pileatus has replaced S. m. pluto
throughout the entire lower Rios Jurua/Solimoes/Coarf
interfluve and has already passed the headwaters of the Rio
Coarf into S. m. pluto's last stronghold, the Rios Solimoes/
Coarf/Purds interfluve as far as the Igarapd Pauapixuna.
As in the case of Saguinus midas and S. bicolor, no
interbreeding is taking place and there seems to be a sudden
transition between populations of S. m. pileatus and S. m.
pluto. These field observations by M. G. M. and T. van
Roosmalen during a recent survey of the entire Rio Purds,
therefore, would justify elevating these two tamarins to full
species status, Saguinus pileatus and Saguinus pluto, distinct
from Saguinus mystax, west of the Rio Jurui.

These arguments apply as well to Callicebus, which exhibit
extreme territorial behavior centered on family groups (a
pair with its offspring of subsequent years), as they do to
Saguinus (Peres, 1989). In both genera, one can distinguish
two 'ecospecies'. In the central-Amazonian ( .-
taxa belonging to the C. moloch and C. cupreus Groups
and taxa belonging to the C. torquatus Group, sympatry
between members of each of these two Groups is common,
because they exhibit different habitat preferences, dietary
requirements, and foraging behavior. In contrast, C .-
cupreus, C. caligatus and C. brunneus, for example, can never
occur sympatrically, as Hershkovitz (1990) suggested,
because they ecologically exclude one another.

The research on which a large part of this paper is based
indicated that the biogeography of Amazonian primates
continues to be poorly known. For example, the recent
survey of the Rio Purds, conducted by M. G. M. and
T. van Roosmalen, revealed two species of ( .... new
to science (including C. stephennashi described here), one of
Saguinus, and one of Ateles, and significant modifications
in a number of distributions given by Hershkovitz in his
various reviews. Zoological collections from the Rio Purds
are few and far between, and many of them are quite old.
Collections used to be acquired by purchasing live and dead
animals from animal dealers who sent natives into the bush,
often far up- or downstream from where they camped.





Neotropical Primates 10(Suppl.), June 2002


Consequently, errors were made in labeling and recording
exact localities. Moreover, museum skins with the skull and
skeleton removed often do not clearly show characteristics
of head, hands, and feet, which are so important in
identification, especially for Callicebus, Mico, Saguinus and
Ateles. Another complication is the under-representation
of monkey species occurring on dry-land or terra fire
rainforests, among them Saguinus, Mico, Callicebus,
Chiropotes, Ateles, and Lagothrix. In part this is because it is
hard to reach the terra firme hinterland along most of the
major Amazonian rivers. Rivers such as the Solim6es, Jurua,
Purds, and Madeira are fringed with almost impenetrable
stretches, often as wide as 10 kilometers or more, of white-
water flooded forests (vdrzeas), the kind of habitat where
many primate species do not venture. Alouatta, Cacajao,
Cebus, Saimiri, and Aotus do occur in these habitats, but
many of the others do not even go there seasonally.

Given the relative lack of study of terra fire areas, it is
understandable that the river barrier hypothesis of Wallace
(1852) is not always believed to apply to primates. Wallace,
while conducting fieldwork and collecting specimens in
the mid-nineteenth century in the upper Rio Negro and
Rio Uaupes, noticed the isolating effect of rivers such
as the Rio Amazonas and its major tributaries (Tapaj6s,
Madeira, Branco, Negro, Purds, Jurua) on the distribution
of monkeys and described it in his paper "On the Monkeys
of the Amazon", published seven years before Charles
Darwin's "The Origin of Species" (1859). Monkeys from
the Amazon provided the data, along with many later
collected in the Malay Archipelago, that led Wallace to his
theory of evolution by natural selection. Only systematic
field studies will test the hypothesis that rivers in lowland
Amazonia can act as effective barriers to the dispersal of
organisms and, as such, induce speciation and generate
allopatric species. Few such studies to date include those of
Capparella (1987, 1988) on the distribution of understory
birds, Peres et al. (1996) on the primate fauna along the Rio
Jurua, and M. G. M. and T. van Roosmalen's systematic
surveys of the Rios Madeira/Tapaj6s, Madeira/Purds, and
Purds/JuruA interfluves (1998, 2000, this paper). In the
coming years, the latter researchers intend to systematically
survey all major rivers of lowland Amazonia and their main
tributaries by boat, collecting dung, tissue and/or hair
samples from both wild and captive specimens of monkeys,
and some other megafaunal elements such as manatees,
giant otters, tapir, deer, and large cats. This project also
includes several terra firme tree species that have their seed
dispersal adapted to terra fire dwelling gut-dispersers,
such as Ateles and Lagothrix.

In our review of Amazonian marmosets (formerly genus
C,//.rI, now Mico), we note that all distributional
boundaries for marmosets coincide with rivers that flow
from the Central Brazilian Plateau to the Rios Madeira and
Amazonas (Van Roosmalen etal., 2000). These rivers mostly
fall within the black-water or clear-water category, which
do not meander much and have not changed their courses
over considerable geological times. In the current paper,


the same pattern is shown for distributions of titi monkeys,
genus Callicebus, at least in the recently inventoried basins
of the Rios Madeira and Purds, conforming to Wallace's
hypothesis of river barrier-assisted speciation.


Conservation Status of the Two New Species

As with all members of the C. moloch and C. cupreus Groups,
( .... bernhardi and ( .... stephennashi prefer
lightly to heavily disturbed terra fire forest, liana forest,
and black-water and clear-water flooded gallery forest.
Densities in undisturbed matrix high dry-land rainforest are
generally low and, when present, the titis seem to frequent
natural edge habitats, such as tree-fall clearings, fringes of
streams and lakes, and liana forest. However, wherever the
rain forest has been disturbed, both naturally as well as
anthropogenically, these 'ecospecies' of titi monkey seem to
thrive. The highest densities of these monkeys are usually
found close to human habitations, along roads, and along
banks of rivers and larger creeks, where human settlements
are mostly situated in this part of Amazonia. Therefore,
there is no reason to suspect that ( .... bernhardi and
( .... stephennashi are threatened. The forests in the
northern part of ( .... bernhardi's range, between the
lower and middle Rio Aripuana and Rio Madeira are still
in almost pristine condition, although selective logging
has taken place along the navigable rivers. There are no
major towns or cities in the area, except Manicord and
Auxiliadora, both located on the right bank of the Rio
Madeira. Indian tribes, which usually hunt relatively small
mammals, including titis, only live in the southern part
of its range. Elsewhere, the local people (caboclos) do not
normally hunt small game and are widely scattered in small
settlements of one to a few families along the major rivers,
the Rios Madeira, Aripuana, Roosevelt, and Ji-Parana, and
along the lower courses of minor rivers, such as the Rios
Matauri, Urui, Maripaui, Araud, Atininga, Manicord, and
Rio dos Marmelos. The interfluvial basins of these black-
and clear-water rivers are practically uninhabited.

The main threat to the habitat of ( .... bernhardi comes
from the Transamazonian Highway that crosses its range in
the south connecting the city of Humaita on the left bank
of Rio Madeira with Apuf on the Rio Sucunduri, Itaituba
on the Rio Tapaj6s, and the states of Para and Mato Grosso.
Given that we still do not know the precise distribution of
( .... stephennashi, we cannot say anything more about
its current conservation status than the general comments
given above, which apply to any member of the C. moloch
and C. cupreus Groups.


Acknowledgments

The authors thank Stephen Nash for producing the
high-quality drawings and maps that illustrate this paper,
Robert Voss for his kind assistance in measuring the skull
of the holotype ( .... bernhardi, Anthony Rylands,





Neotropical Primates 10O(Suppl.), June 2002


William Konstant and Kim Meek for assistance in editing
and lay-out, Ella Outlaw for support from the office in
Washington, DC, and Betty van Roosmalen-Blijenberg for
taking care of the captive primate specimens in Manaus.
Support for fieldwork and for publication of this paper was
provided by the Margot Marsh Biodiversity Foundation
and Conservation International, Washington, DC, and
Conservation International do Brasil, Belo Horizonte.


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Figure 47. Array of faces of titi monkeys, genus Callicebus, arranged by group. Illustration by Stephen D. Nash.


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modestus olallae




















dufluuuplui/lu eonanthe


Figure 48. Titi monkeys, genus Callicebus, donacophilus group. Illustration by Stephen D. Nash.





Neotropical Primates 10(Suppl.), June 2002


II. cupreus Group


cupreus


dubius


stephennashi


Figure 49. Titi monkeys, genus Callicebus, cupreus group. Illustration by Stephen D. Nash.





Ill. moloch Group



I











hoffmannsi







baptista


'i


Figure 50. Titi monkeys, genus Callicebus, moloch group. Illustration by Stephen D. Nash.


Neotropical Primates 10(Suppl.), June 2002


bernhardi





Neotropical Primates 10(Suppl.), June 2002


IV torquatus Group


torquatus


lugens


lucifer


Figure 51. Titi monkeys, genus Callicebus, torquatus group. Illustration by Stephen D. Nash.


medemi


regulus


purinus





Neotropical Primates 10(Suppl.), June 2002


V. personatus Group


~' ~I-jEEa


melanochir


Figure 52. Titi monkeys, genus Callicebus, personatus group. Illustration by Stephen D. Nash.


barbarab






V barbarabrownae


, ',imit'r vi




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